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Genetic Variations of

DAN L. LINDSLEY
and

E. H. GRELL

BIOLOGY DIVISION, OAK RIDGE NATIONAL LABORATORY


operated by
UNION CARBIDE CORPORATION
for the
UNITED STATES ATOMIC ENERGY COMMISSION

Carnegie Institution of Washington Publication No. 627


Library of Congress Catalog Card Number 68-15915
ISBN 0-87279-S38-8
First Printing-1968
Second Printing - 1 9 7 2
revised from

THE MUTANTS OF DROSOPHILA MELANOGASTER

Calvin B. Bridges and Katherine F. Brehme

Carnegie Institution of Washington Publication 552, 1944


We dedicate this book to Alexander Hollaender on his retirement as director
of the Biology Division of the Oak Ridge National Laboratory and in recognition
of his continuous encouragement and support-
EDITOR Isabel L. Norton
CONSULTING EDITOR Eileen J. Slaughter
INDEXING Elizabeth S. Von Halle
PROGRAMMING Charles E. Price
TYPING Neva P. Hair
COMPOSITION Ethel Cagle
DESIGN, LAYOUT AND Technical Publication Department
PRODUCTION Technical Information Division
Oak Ridge National Laboratory
Oak Ridge, Tennessee
Atomic Energy Commission
Division of Technical Information Extension
Oak Ridge, Tennessee
PREFACE

The last exhaustive compilation of genetic ance; and (7) wild-type stocks. All except the
variations of Drosophila melanogaster was "The first two groups have relatively few entries.
Mutants of Drosophila melanogaster" prepared by Several new categories of effects unknown or
Calvin B. Bridges and Katherine S. Brehme; it nearly so in 1942 are included here. (1) Pseudo-
was published in 1944 and was complete through alleles: the intensive investigations into pseu-
1942. This volume is a revision of their work; doallelism and complementation dating from the
it contains new information on variants already pioneering work of E. B. Lewis on Star and as-
described and descriptions of variants dis- troid (1945, Genetics 30: 137-66) have produced
covered since 1942; it is reasonably complete information on the genetic fine structure of many
through 1966. The new material was extracted loci. (2) Isozymes: a series of genetically con-
from the literature, from Drosophila Information trolled enzyme polymorphisms and deficiencies
Service, and from voluminous contributions of described mostly in the last decade; their dis-
unpublished material supplied by Drosophila covery was made possible by the development of
geneticists throughout the world. The revision gel electrophoresis. (3) Compound chromosomes:
describes genetic material currently available formed by the attachment of two doses of one
to Drosophila geneticists and extinct material chromosome arm to a single centromere; repre-
that may be encountered in earlier literature on sented by only the attached-^ chromosome in the
the subject. original edition, the various classes of compound
The work of Bridges and Brehme was divided chromosomes now available occupy an entire sub-
into two sections, one describing wild-type section. (4) Marked Y chromosomes: Y chromo-
stocks and the other describing the known depar- somes marked by the genes carried on small at-
tures from the normal genotype. Our revision is tached euchromatic segments derived from the X
divided into seven sections: (1) mutants, with or an autosome. (5) Reciprocal translocations
about 3000 entries; (2) chromosome aberrations, between the X and Y chromosomes. (6) Attached
more than 1500 entries; (3) special chromosomes, XY chromosomes: chromosomes with the portions
i.e., multiply marked chromosomes, balancers, of the X and Y chromosomes necessary for male
compound chromosomes, Y derivatives, and X-Y viability and fertility attached to a single centro-
combinations; (4) cytological markers; (5) depar- mere.
tures from diploidy; (6) nonchromosomal inherit-
Development of the system of nomenclature The conventions adopted fdr naming and sym-
designating genetic variations of Drosophila mel- bolizing different types of genetic changes are
anogaster has been rather haphazard; conse- discussed at the beginning of the different sec-
quently, the system is not a logical structure but tions of the book. Symbols of all genetic vari-
is replete with relics, redundancies, and incon- ants both normal and abnormal are always itali-
sistencies. Revision into a consistent scheme cized but their names in text are printed in roman.
is not practicable, creating as it would a chaotic We are grateful to our colleagues throughout
discontinuity in the literature. Even were such the world for their cooperation in making avail-
a revamping considered desirable, design of such able to us their unpublished observations and
a system is not obvious, since a change pro- in responding to our numerous queries. Special
posed to obviate one inconsistency would likely thanks are due Doctors E. B. Lewis, the late
create more conflicts than it alleviated. There- H. J. Muller, J. Schultz, and A. H. Sturtevant,
fore, with few exceptions, we have adhered to who served as an informal board of consultants.
the conventions established by Bridges and They have contributed a measure of success to
Brehme in the original volume. Some changes this effort but bear no responsibility for its
were made to correct glaring inconsistencies shortcomings.
and others to facilitate automatic handling of
Drosophila symbols.
D. L. L.

July 1967 E. H. G.
guide to

Genetic Variations of MUTATIONS

CHROMOSOME ABERRATIONS

Deficiencies

Duplications

Inversions

Rings

Translocations

Transpositions

SPECIAL CHROMOSOMES

Balancers

Compound Chromosomes

Multiply Marked Chromosomes

X—Y Combinations

Y Derivatives

CYTOLOGICAL MARKERS

DEPARTURES FROM DIPLOIDY

NONCHROMOSOMAL INHERITANCE

WILD TYPE STOCKS

Vn/T(Y;3)1 GENETIC AND CYTOLOGICAL MAPS


Mohr *nd Mossige, 1943, Norske Vldenskaps-Akad. 7: 1—51.
MUTATIONS

A variant exists when there are two or more sive and its name begins with a lower-case
alternative phenotypes. Usually, one is des- letter.
ignated as normal or wild-type because it is the For convenience, a symbol is assigned to each
phenotype characteristic of wild-type flies; the mutant type. This symbol is an abbreviation of
other or others are considered mutant alternatives the name that uniquely designates the mutant in
because they represent departures from normal. question; it combines brevity with information.
Distinction between normal and mutant may be- It usually begins with the same letter as the
come blurred or disappear where both alternatives name, is always italicized, and never contains
are characteristic of wild-type strains, as in iso- Greek letters, subscripts, or spaces; e.g., r for
alleles. The pair or group of alternatives de- rudimentary, R for Roughened, ro for rough, rs
fines a locus, which is given a name that sug- for rose, and ry for rosy. In designations of
gests the main diagnostic features of the mutant genotypes with several mutant genes, symbols
form of the locus without regard to secondary of genes on the same chromosome are separated
characters. The name is concise and is pref- by spaces (e.g., y w i B); symbols of genes on
erably a simple adjective such as black or a homologous chromosomes are separated by a
noun such as Bar. When the main character is slash bar (e.g., y H- / B); symbols of genes on
recognized in the heterozygote, the mutant is noohomologous chromosomes are separated by
considered dominant and its name begins with semicolons and spaces (e.g., bw; v; ey). Names
an upper-case letter; when it is recognized in are not italicized in text.
the homozygote, the mutant is considered reces-
GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

MULTIPLE ALLELES. The alternatives or al- B+. The plus symbol alone implies the normal
leles at a particular genetic locus are designated (wild-type) allele in any context, such as y/+ or
by the same name and symbol and are differenti- y m //+. Absence of a particular locus may be
ated by distinguishing superscripts. At publication noted by use of a superscript minus sign with
of the original volume, heterozygotes for allelic the symbol; e.g., bb~.
mutants were thought always to show a mutant Loci controlling electrophoretic mobility of
phenotype and segregation of the mutants at mei- enzymes and other proteins require special con-
osis. Since that time, however, discoveries of ventions. Since electrophoretic variants can be
complementation and intra-allelic recombination scored equally well in heterozygotes and homo-
have revealed the widespread existence of com- zygotes, the genes controlling them are consid-
plex pseudoallelic series, with consequent com- ered dominant; e.g., Adh. Alleles specifying
plications in the definition of allelism. By the the variants are differentiated by arbitrary super-
criteria that they occupy virtually identical po- scripts; e.g., AdhF, AdhD, and those specifying
sitions on the genetic map and have similar phe- the absence of a particular enzyme or other pro-
notypic effects, mutants formerly thought to be tein by an appropriate superscript, such as n
at different loci may now be considered changes (negative), a zero, or a minus sign, rather than
in the same pseudoallelic complex; e.g., (a and by a lower-case symbol; e.g., Adhnl. The sole
spl, Iz and amx, and m and dy. The locus will exceptions to the rule that the genetic determi-
probably be defined ultimately as the unit of nants of electrophoretic protein variants be sym-
transcription or, more likely, of translation; but bolized as dominant genes are loci originally
in the current state of knowledge, we have been recognized by recessive phenotypes and so
content to adhere to the historic terminology. named; e.g., v and ry. For proteins with unde-
One school of thought names every recombina- termined activity, we use the symbol Pt- fol-
tionally separable element; another gives all lowed by an arbitrary designation specifying the
members of the same complex a single name with particular protein; e.g., Pt-1. Abbreviations for
arbitrary superscripts. The latter is our pref- the protein and the gene are frequently identical,
erence. and both are used in most discussions. The
The superscript notation designating alleles gene symbol may be differentiated from the pro-
has a number of different forms. A common de- tein symbol by having only its initial letter cap-
vice is an abbreviation that further characterizes italized and by being italicized, whereas the
the particular allele or that was used as the protein symbol is in roman capitals; e.g., ADH.
locus symbol before allelism was established. In several instances where two members of the
This practice is avoided because it has the dis- same allelic series were formerly given different
advantage of preempting useful symbols and locus names, both are here included under one
names from use as locus designations. Another name; e.g., Pm = bwvl. In other cases, we as-
unacceptable device is the use, as superscripts, sume allelism of mutants with similar phenotypes
of elements of the genotype in which the allele and genetic positions even though they have not
arose, since such a designation implies some- been tested for phenotypic interaction. In such
thing more than a trivial connection between instances, the basis for the assumption is usu-
allele and element. More-acceptable super- ally noted. Since the practice has not been con-
scripts for allelic designations are arbitrary sistent, some alleles may be described as dif-
numbers, experiment numbers, capitalized initials ferent genes. We make special effort to infer
of the finder or laboratory, or the date of dis- allelism for Minute loci and for factors causing
covery. The numeral 1 is the implied superscript production of melanotic pseudotumors. Bridges
of nonsuperscripted symbols. Whereas genes in and Brehme made few such inferences. Except
the same allelic series are designated by the in special cases, investigation of allelic inter-
same symbol but with different superscripts, mu- action of sex-linked recessive lethals is not
tants with similar phenotypes at different loci possible; consequently, they are often given dis-
are not given the same symbol and differentiated tinctive symbols where allelism may actually
only by a superscript; this was done extensively exist.
in the past; for example, for genes causing for- MIMICS. Mutants at different loci sometimes
mation of melanotic pseudotumors. have similar phenotypic effects. Such loci may
For a recessive allele of a preponderantly dom- be handled in several ways. The simplest is to
inant series or a dominant allele of a predomi- give each a distinctive name (e.g., vermilion,
nantly recessive series, the superscripts used cinnabar, scarlet, karmoisin, cardinal); this
are r and D, respectively; e.g., Hnr, Hnr2, and method has the effect of scattering such mimics
hwD. Finally, for the normal allele in a series, throughout the alphabetical listing. Or a com-
a superscript plus sign may be used, e.g., 6+ or mon symbol separated by a hyphen from a dis-
MUTATIONS

tinguishing symbol may be used (e.g., tu-la, e.g., 3.0 represents a more accurate location
tu-lb, tu-2 for genes controlling production of than 3. Map units are not computed to the
melanotic pseudotumors). Distinctive suffixes second decimal place. Accuracy of a map
are also useful (e.g., rough, roughoid, roughish, position determination is of course dependent
roughex; plexus, Plexate; dachs, dachsous; on the accuracy of the positions assigned to
maroon, maroonlike). The latter two schemes the reference markers; i.e., on the accuracy
frequently have the virtue of placing like pheno- of the map. We treat the map as a rough
types in sequence in an alphabetical listing. guide to the relative positions of loci but,
Some phenotypes result from mutation at many considered on a refined level, it may be inac-
loci in all chromosomes; these are given a com- curate with respect to both position and order
mon symbol followed by a parenthetical desig- of genes. (We have abandoned the ± used by
nation of the chromosome and then by a distin- Bridges and Brehme to indicate a particu-
guishing designation. Examples of this type of larly low level of reliability.)
mutant are the female steriles, the lethals, the origin: For induced mutants, the agent is
Minutes, and the male steriles [e.g., fs(2)B, given; mutants recovered from untreated par-
1(1)Jl, M(l)n, ms(2)E4, respectively]. Conven- ents or a wild population are listed as spon-
tions for formulating distinguishing symbols are taneous. Isoallelic variants found as major
similar to those for superscripts; use of infor- components of stocks or populations are
mation about the cytological or genetic location listed as naturally occurring alleles. The
is avoided to allow updating such information stock or chromosome in which each mutant
without changing the symbol. arose was listed by Bridges and Brehme;
MODIFIERS. The primary effect of some mutants unless the new mutant is virtually insepa-
is to cause another mutant to exhibit a more- rable from some element of the stock of ori-
extreme departure from normal (enhancer) or a gin (e.g., y31d in In(l)sc8), we omit mention
more nearly normal phenotype (suppressor). Such of the original stock.
mutants are symbolized e or E and sti or Su, fol- discoverer: Name, date of discovery.
lowed in parentheses by the gene modified. Des- synonym: Alternative symbol or name or both,
ignation of the particular allele modified appears mostly obsolete terminology.
as a superscript within the parentheses and references: Sources of the major descriptive
alleles of the modifier gene as superscripts out- material are listed, but bibliographic material
side the parentheses; e.g., su(lz34) and su(Hw)2. may also appear in some of the other cate-
Terms such as dilutor, exaggerator, inhibitor, gories.
intensifies and modifier were also formerly used, phenotype: The most important departures from
but we have usually attempted to classify such normal, which are usually those suggested by
genes as enhancers or suppressors. the name, are described first. Other infor-
FORMAT. Mutants with their descriptions mation about the phenotype follows, and
are now listed alphabetically according to sym- finally there may be data on viability and fer-
bol and cross-indexed according to name. Cur- tility. The last item in the phenotypic de-
rent terminology is listed in bold face. All scription is the rank, abbreviated RK. Mu-
cases of synonymy are also listed in body type tants are classified into three different ranks
with cross-references to current usage. Mutants according to their utility in experiments in
no longer existing in published stock lists or in which counts are made: RK1 mutants are
private stock lists that we have examined are easily scored; RK2 mutants are usable but
considered lost and are preceded in the list by less convenient; RK3 mutants have limited
a star. Each mutant is described according to usefulness. An RK3 mutant may be one with
the following format: good expression and viability but simply not
symbol: name convenient to use in counting experiments;
location: The location is indicated by the e.g., enzyme polymorphisms. The letter A
chromosome number, separated by a hyphen follows the rank of mutants associated with
from the genetic position on the chromosome. chromosome aberrations.
Two levels of accuracy of the genetic loca- other information: This category contains
tion are indicated, those carried to tenths of miscellaneous information that does not fit
a unit being the more accurately determined; into one of the other categories.
GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

a: arc phenotype: Nearly normal; distinguished by its


location: 2-99.2. interaction with certain svr alleles (see Gold-
discoverer: Bridges, 12e24. schmidt, 1945, table 74). RK3.
references: Bridges and Morgan, 1919, Carnegie cytology: Salivary chromosomes normal (Kodani).
Inst. Wash. Publ. No. 278: 202 (fig.). £>a2
Morgan, Bridges, and Sturtevant, 1925, Bibliog. origin: Spontaneous.
Genet. 2: 212 (fig.). discoverer: Goldschmidt.
Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol. synonym: bran2.
2: 745-55. references: 1945, Univ. Calif. (Berkeley) Publ.
phenotype: Wings broader, bent downward in slight, Zool. 49: 363-73, 388-89.
even arc, and edges drawn down to diamond shape. phenotype: Wings somewhat more angular than aba.
Sometimes in stock, wings are bent upward instead Interactions with other genes shown in table 74 of
of downward. Crossveins closer together. RK2. Goldschmidt (1945). RK2A.
cytology: Placed between 57F11 and 58E4 on the cytology: Associated with Df(2R)aba2 = Df(2R)58D5-
basis of its inclusion within Df(2R)M-l = 6;58D7-8 (Goldschmidt, 1945).
Df(2R)57Fll-58Al;58F8-59Al but not Dp(2;3)P =
Dp(2;3)58E3-4;60D14-E2;96B5-Cl (Bridges, 1937). origin: Spontaneous.
Likely in band 58D6 or 7 based on Df(2R)ab"2 - discoverer: Goldschmidt.
Df(2R)58D5-6;58D7-8. synonym: bran^.
references: 1945, Univ. Calif. (Berkeley) Publ.
Zool. 49: 386-89.
phenotype: Normal; distinguished by its interaction
with certain svr alleles (see Goldschmidt, 1945,
table 74). RK3.
cytology: Salivary chromosomes normal (Kodani).

origin: Spontaneous. Probably a derivative of aba3.


discoverer: Goldschmidt.
synonym: bran4.
references: 1945, Univ. Calif. (Berkeley) Publ.
Zool. 49: 389-90, 490.
phenotype: Like aba; distinguished by its interaction
a: arc with certain svr alleles (see Goldschmidt, 1945,
From Bridges and Morgan, 1919, Carnegie Inst. Wash. table 74). RK2.
Publ. No. 278: 148. cytology: Salivary chromosomes normal (Kodani).

aba: arc-broad angular *abadb: arc-broad angular dumpy blistered


origin: Spontaneous. origin: Spontaneous.
discoverer: Goldschmidt, 1934. discoverer: Goldschmidt.
synonym: Always referred to as bran: broad angular synonym: brandb.
by Goldschmidt, but shown by him to be an allele references: 1945, Univ. Calif. (Berkeley) Publ.
of arc. Zool. 49: 370-71, 388-89.
references: 1945, Univ. Calif. (Berkeley) Publ. phenotype: Like aba; distinguished by its interaction
Zool. 49: 351-56, 388-89, 519. with certain svr alleles (see Goldschmidt, 1945,
phenotype: Wings broader and shorter than wild type, table 74). RK2.
blunt at the tip. Frequently shows upturned pos- cytology: Salivary chromosomes normal (Kodani).
terior scutellar bristles. In combination with *abadp; arc-broad angular dumpy
svrPoi, produces soft blistered wing. Other inter- origin: Spontaneous.
actions described by Goldschmidt, 1945, table 74. discoverer: Goldschmidt.
Wing grows in pupal stage to full length and then synonym: brandP.
retracts, possibly with histolysis [Goldschmidt, references: 1945, Univ. Calif. (Berkeley) Publ.
1935, Z. Induktive Abstammungs- Vererbungslehre Zool. 49: 373-86, 388-89.
69: 38-131 (fig.)J. RK2. phenotype: Normal; distinguished by its interaction
cytology: Salivary chromosomes normal (Kodani). with certain svr alleles (see Goldschmidt, 1945,
other information: Claimed to recur repeatedly in table 74). RK3.
certain lines (Goldschmidt, 1945). cytology: Salivary chromosomes normal (Kodani).
other information: Claimed by Goldschmidt to recur
origin: Spontaneous, repeatedly in certain lines.
discoverer: Goldschmidt. *ai>or; arc-broad angular rudimentary
synonym: bran*. origin: Spontaneous derivative of abadP.
references: 1945, Univ. Calif. (Berkeley) Publ. discoverer: Goldschmidt.
Zool. 49: 364-69, 388-89. synonym: branr.
MUTATIONS

references: 1945, Univ. Calif. (Berkeley) Publ. cytology: Salivary chromosomes normal (Hannah-
Zool. 49: 378-79, 388-89. Alava).
phenotype: Wing broad, round and dp-like. Interacts *aM60: arc of Meyer
with certain svr alleles (see Goldschmidt, 1945, origin: X ray induced.
table 74). RK2. discoverer: Meyer, 60f.
references: 1963, DIS 37: 50.
* o Bo ; arc-Broad angular Dominant phenotype: Homozygous lethal. RK3A.
origin: Spontaneous derivative of aba. cytology: Associated with In(2LR)aM6°; breakpoints
discoverer: Goldschmidt. unknown.
synonym: Bran. *A: Abnormal abdomen
references: 1945, Univ. Calif. (Berkeley) Publ. location: 1-4.5.
Zool. 49: 360-63, 388-89. discoverer: Morgan, l l g .
phenotype: aBa/+ resembles aba/aba. aBa/aba synonym: Abnormal.
shows Minute bristles. RK2. references: 1915, Am. Naturalist 49: 384-429 (fig.)-
* o 8 o C : arc-Broad Angular in Canton Morgan and Bridges, 1916, Carnegie Inst. Wash.
origin: Spontaneous. Publ. No. 237: 27 (fig.).
discoverer: Goldschmidt. phenotype: Tergites and sternites raggedly incom-
synonym: Bran^a. plete, exposing a thin crinkled cuticle; bristles
references: 1947, J. Exptl. Zool. 104: 197-221. and hairs on abdomen correspondingly eliminated.
phenotype: aBaC/+ is normal; aBaC/aba ig broad Highly variable, wild phenotype in old dry cultures.
angular, but overlaps wild type. aBa^ is dominant A/+ less extreme than A/A and A male; homozy-
in its interaction with certain svr alleles. RK3.. gous female fully viable and fertile. RK2 in well-
cytology: Salivary chromosomes normal (Hannah- fed cultures.
Alava). other information: Lost by reversion to wild type.
*aBapl: arc-Broad angular in silver-pointed A: see bwA
origin: Spontaneous.
discoverer: Goldschmidt, 1947. location: 1- (just to the right of w; judged to be
synonym: BranP0*47"1. allelic to .4).
references: 1947, J. Exptl. Zool. 104: 197-221. origin: Spontaneous.
phenotype: a13^1/+ resembles aba/ab*. RK2. discoverer: Hillman, 53g.
cytology: Salivary chromosomes normal (Hannah- references: 1953, DIS 27: 56.
Alava). Hillman and Barbour, 1963, Proc. Intern. Congr.
*aBop2 Genet., 11th. Vol. 1: 170.
origin: Spontaneous. phenotype: Highly variable; ranging from extreme
discoverer: Goldschmidt, 1947. expression in young cultures to normal in old cul-
synonym: BranPoi47-2. tures. Expression in AS36/AS36 females > A536/Y
references: 1947, J. Exptl. Zool. 104: 197-221. males > AS3&/+ females. Expression varies from
phenotype: Phenotype normal in combination with loss of tergites 2—8 in extreme cases to loss of
aba and +; homozygous lethal. Dominant in inter- lateral part of tergite in one or more segments.
actions with certain svr alleles. RK2 as lethal. RK2A in young cultures.
cytology: Salivary chromosomes normal (Hannah- cytology: Associated with rearrangement of 2-5
Ala va). bands in 3C-D.
* o BoX ; arc-Broad angular from X irradiation a-3: seea(3)26
origin: X ray induced. *A-p; Abnormal abdomen-polygenic
discoverer: Goldschmidt. location: Polygenic.
synonym: Branx. discoverer: Sobels, 49j.
references: 1945, Univ. Calif. (Berkeley) Publ. references: 1950, DIS 24: 62.
Zool. 49: 521-22. 1951, DIS 25: 75-76.
phenotype: Resembles abo2 and abadb, but more or 1952, Genetica 26: 117-279 (fig.).
less dominant. Homozygote never obtained. Inter- 1952, Trans. Intern. Congr. Entomol., 9th. Vol. 1:
actions listed by Goldschmidt (1945, table 153). 225-30.
RK2. synonym: AA; Aay: Asymmetric.
phenotype: Incomplete mediodorsal fusion and one-
origin: Spontaneous. sided reduction of tergites. When more than one
discoverer: Goldschmidt. tergite is abnormal, spiral segmentation types are
synonym: Brany P* bl. most frequent. Expression strongly dependent on
references: 1947, J. Exptl. Zool. 104: 197-221. environment. Penetrance and expressivity corre-
phenotype: Homozygotes usually lethal; rare survi- lated (Bezem and Sobels, 1953, Konixikl. Ned.
vors have short, folded wings and are sterile. Akad. Wetenschap., Proc. Ser. C 56: 48-61). In
aBay/+ is broad-angular, with occasional truncate- strains selected for penetrance of A-p, mediodorsal
like wings. In combination with svrPoi resembles fusion or asymmetrical reduction of head and
rudimentary and blistered. RK2 as lethal. thorax also occur. RK3.
8 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

*a(1)48: abnormal abdomen in chromosome 1 *A(2)51


location: 1- (not located). location: 2- (not located).
origin: Spontaneous. origin: Spontaneous.
discoverer: Zimmerman, 1948. discoverer: Zimmerman, 1951.
references: 1952, DIS 26: 69. references: 1952, DIS 26: 69.
1954, Z. Induktive Abstammungs- Vererbungslehre 1954, Z. Induktive Abstrammungs- Vererbungslehre
86: 327-72 (fig.). 86: 327-72 (fig.).
phenotype: Has no phenotype of its own but in- phenotype: None. Enhances a(2)48 and a(3)48. RK3.
creases the incidence of abdominal malformations o(3)26
in a(2)48 and a(3)48 and in progeny of such flies. location: 3-27 (to the right of se).
Viability and fertility good. RK3. origin: Spontaneous.
*a(1)50 discoverer: H. A. and N. W. Timofeeff-Ressovsky.
location: 1- (not located). synonym: a-3.
origin: Spontaneous. references: 1927, Arch. Entwicklungsmech. Organ.
discoverer: Zimmerman, 1950. 109: 70-109.
references: 1952, DIS 26: 69. Schaffer, 1935, Z. Induktive Abstammungs-
1954, Z. Induktive Abstammungs- Vererbungslehre Vererbungslehre 68: 336—60 (fig.).
86: 327-72 (fig.). phenotype: Irregular reduction of abdominal tergites,
phenotype: Irregularities in abdomen most frequently sternites, pigmentation, and bristles; more marked
involving the anterior segments. Penetrance 1 per- in females and increased by crowding and dry food
cent. Enhances maternal effects of a(2)48 and (Braun, 1938, Am. Naturalist 72: 189-92).
a(3)48. Viability and fertility good. RK3. Schaffer's data (1935) suggest irregular dominance
*o(l)Sl in heterozygote, overlapping of wild type in homo-
location: 1- (not located). zygote, and genetic modifiers. RK3.
origin: Spontaneous. *a(3)48
discoverer: Zimmerman, 1951. location: 3- (not located).
references: 1952, DIS 26: 69. origin: Spontaneous.
1954, Z. Induktive Abstammungs-Vererbungslehre discoverer: Zimmerman, 1948.
86: 327-72 (fig.). references: 1952, DIS 26: 69.
phenotype: Shows maternal effect only, with 2 per- 1954, Z. Induktive Abstammungs- Vererbungslehre
cent of progeny affected. Abnormalities more 86: 327-72 (fig.).
anterior than those of a(2)4S and a(l)50. Viability phenotype: Only a maternal effect affecting 2.5 per-
and fertility good. RK3. cent of progeny. Irregularities most frequently
*o(2)48 involve posterior segments of abdomen. Viability
location: 2- (not located), and fertility good. RK3.
origin: Spontaneous. A34: see bwV6
discoverer: Zimmerman, 1948. aa: an arista
references: 1952, DIS 26: 69. location: 3-0.
1954, Z. Induktive Abstammungs- Vererbungslehre discoverer: Bridges, 23dlO.
86: 327-72 (fig.), synonym: al~b: aristaless-b.
phenotype: Abdominal irregularities most frequently references: Morgan, Bridges, and Sturtevant, 1925,
involve anterior segments. Penetrance 7 percent. Bibliog. Genet. 2: 218.
Also shows maternal effect. Viability and fertility phenotype: Aristae bare or tufted. Wings somewhat
good. RK3. broader than wild type. Expression variable, over-
*o(2)50 laps wild type often in female and sometimes in
location: 2- (not located), male. RK3.
origin: Spontaneous. cytology: Placed between 61E2 and 62A6, on basis
discoverer: Zimmerman, 1950. of its inclusion in Df(3L)D = Dt(3L)61E2-
references: 1952, DIS 26: 69. Fl;62A4-6 from T(Y;2;3)D.
1954, Z. Inductive Abstammungs- Vererbungslehre
86: 327-72 (fig.), Aa: Altered abdomen
phenotype: None. Six percent progeny affected, i.e., location: 1- (not located).
only maternal effect. RK3. origin: X ray induced in the In(l)dl-49, y w t compo-
nent of C(1)DX, y i of Muller.
location: 2- (not located), discoverer: Cicak, 56f.
origin: Spontaneous, references: Cicak and Oster, 1957, DIS 31: 80.
discoverer: Zimmerman, 1951. phenotype: Heavy deposition of melanin in tergites
references: 1952, DIS 26: 69. of females and males. Males sterile, therefore
1954, Z. Induktive Abstammungs- Vererbungslehre homozygous females not produced. RK2A.
86: 327-72 (fig.). cytology: Probably associated with a rearrangement
phenotype: Penetrance 50 percent. Also shows in addition to ln(l)dl-49.
maternal effect. RK3. A A: see A-p
MUTATIONS

ab: abrupt cytology: Placed in region between 59E2 and 60B10


location: 2-44.0. by Bridges (1937) on basis of its being to the right
origin: Spontaneous. of In(2R)bv/Vr>»1 = In(2R)41B2-Cl;59E2-4 and to
discoverer: Bridges, 16jl6. the left of Df(2R)Px = Df(2R)60B8-10;60Dl-2.
references: Morgan, Bridges, and Sturtevant, 1925,
Bibliog. Genet. 2: 218 (fig.).
phenotype: Vein L5 usually stops after posterior
crossvein. Scutellar bristles usually fewer. Wing
effect probably ^acts during contraction period
(Waddington). Overlaps wild type. RK2.

ab: abrupt
Edith M. Wallace, unpublished.

origin: Spontaneous.
discoverer: Bridges, 23gl6.
synonym: pt: parted.
references: Morgan, Bridges, and Sturtevant, 1925, abb: abbreviated
Bibliog. Genet. 2: 232. From Bridges and Brehme, 1944, Carnegie Inst. Wash.
phenotype: Vein L5 does not reach margin. Scutellar Publ. No. 552: 11.
bristles always fewer than wild type. Hairs parted
down midline of thorax and abdomen. Supra-alar abd: abdominal
bristles sometimes absent. Coxae tend to be location: 3-27 (close to the right of se).
thickened. Males sterile and have rotated gen- origin: Spontaneous.
italia. ab/ab2 resembles ab/ab but has a stronger discoverer: Gottschewski, 1935.
bristle effect. RK2. phenotype: Abdominal bands broken and etched.
Overlaps wild type in test crosses but not in
origin: Spontaneous in In(2L)Cy + In(2R)Cy. homozygous stock. Slightly semidominant. More
discoverer: Edmonds on, 51g. extreme at 19°C. abd/a(3)26 shows slight abd
references: 1952, DIS 26: 60. effect. RK3.
phenotype: A strong allele like ab2. RK2A. abdomen rotatum: see ar
abdominal: see abd
*abt-60h: abrupt-lethal
origin: Spontaneous. *abe: abnormal eye
discoverer: Hall, 60h. location: 1-1.2.
references: 1960, Meyer, DIS 34: 52. origin: Induced by 2-chloroethyl methanesulfonate
phenotype: Homozygote rarely survives. abl"60ll/ab2 (CB. 1506).
has shortened vein L5, but no scutellar bristles discoverer: Fahmy, 1955.
missing, and there is no part down midline of references: 1959, DIS 33: 82-83.
thorax and abdomen. ab1'60fl/ab2 males are phenotype: Eyes rough; either small or deformed.
fertile. RK2. Wings slightly atypical; inner margin frequently
abb: abbreviated removed by large irregular incisions; L4 frequently
location: 2-105.5. stops well short of the wing edge. Flies slightly
discoverer: Bridges, 28d6. smaller than normal. Males about 50 percent as
references: 1937, Cytologia (Tokyo), Fujii Jub. viable as wild type and fertile. Females highly
Vol. 2: 745-55. infertile. RK2.
phenotype: Bristles smaller; especially posterior abero: see abr
scutellars. Developmental time slightly longer Abnormal: see A
than normal. Viability only slightly reduced. abnormal abdomen: see a( )
Classification difficult, especially in early eclo- Abnormal abdomen: see A
sions; improves with age of culture. Enhanced by abnormal eye: see a be
shrunken (2-2.3), making classification easy. abnormal tergites: see abt
RK3; RK2 with shr. abnormal wings: see abw
10 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

abr: abero synonym: Called ac11 by Serebrovsky.


location: 2-83. references: 1918, Genetics 3: 133—72.
origin: Spontaneous. Dubinin, 1930, Zh. Eksperim. Biol. 6: 325—46.
discoverer: Bridges, 33blO. 1933, J. Genet. 27: 443-64.
phenotype: Abdominal banding etched and irregular. phenotype: Posterior dorsocentral bristles missing,
Wing margins irregular. Eyes rough. Bristles and anteriors rarely; hairs usually fewer near posterior
hairs sparse and disarranged. abr/+ sometimes dorsocentrals; intraocellar hairs invariably fewer,
lacks anterior dorsocentrals. Viability usually typically absent. Eyes partly devoid of hairs.
poor. RK3. ac/ac or ac/+ partially suppresses h (Sturtevant).
other information: Not allelic to fr or nw. Hw/ac = Hw/+ (Sturtevant). RK1.
abrupt: see ab cytology: Placed in region 1A5-8 on basis of its in-
Abruptex: see Ax clusion in the XD3P element of T(l;3)sc260-20 _
k
abt: abnormal tergites T(l;3)lA8-B1^61Al-2 and in Dp(l;f)sc2 6 0-2 7 =
location: 1-45.6. Dp(l;t)lA8-Bl;l9F, but not being lost from
origin: Induced by 2-chloroethyl methanesulfonate Df(l)260-5 =Dt(l)lA4-5 (Sutton, 1943, Genetics 28:
(CB. 1506). 210-17).
discoverer: Fahmy, 1955. 2
references: 1959, DIS 33: 83. origin: X ray induced simultaneously with s c 3 .
phenotype: Abdomen affected to various degrees, discoverer: Dubinin, 1928.
from extreme deformation of tergites to slight references: 1929, Biol. Zentr. 49: 328-39.
abnormalities in distribution of pigment and hairs. Serebrovsky and Dubinin, 1930, J. Heredity 21:
Eyes also deformed to various degrees from gross 259-65.
alterations in shape to slight derangement of om- phenotype: Since ac2 and sc3 are for practical
matidia. Wings vary from alterations in size, out- purposes inseparable by crossing over, the effect
line, and venation to small incisions of the inner of ac2 alone cannot be assessed. The double
margin. Most-extreme effects not always posi- mutant removes all bristles except scutellars and
tively correlated, and all flies show several postdorsocentrals. ac2/ac2 or ac2/+ suppresses h
atypical characters. Males viable, fertility severly (Sturtevant). Viability of males low; females
reduced. RK3. nearly inviable. RK2.
abw: abnormal wings cytology: Salivary chromosomes normal (Schultz).
location: 1-60. : see ac3
origin: X ray induced.
discoverer: Halfer, 1963. origin: X ray induced.
phenotype: Wing size reduced; wings upturned; L5 discoverer: Dubinin, 1929.
and crossveins absent. Plexus of veins between synonym: Called ac2 by Dubinin, the earlier ac2
L3 and L4. RK1. with sc3 having been omitted from the series.
sci0. sc-'-' (Sturtevant and Schultz, 1931, Proc.
Natl. Acad. Sci. U.S. 17: 265-70).
references: 1930, Zh. Eksperim. Biol. 6: 300—24.
1932, J. Genet. 25: 163-81.
1933, J. Genet. 27: 443-64.
phenotype: Posterior and usually anterior dorsocen-
trals lacking; other bristles wild type. Hairs re-
moved from areas across rear and front edges of
thorax, through mid-dorsal area, and between
ocelli. RK2A.
cytology: Inseparable from In(l)ac3 =
In(l)lB2-3;lB14-Cl (Muller, Prokofyeva, and
Raffel, 1935, Nature 135: 253-55).
other information: Judged to be an allele of ac but
not sc; it is mutant in combination with ac but not
with sc alleles except for scs which may also
show ac variegation and sc3, now lost, which is
thought to be a sc ac double mutant (Sturtevant).
ac3: see ac*
ac: achaete *QC4
From Bridges and Brehme, 1944, Carnegie Inst. Wash. origin: X ray induced in X chromosome carrying sc.
P u b l . No. 552: 12. discoverer: Dubinin, 1929.
a c : achaete synonym: Called ac3 by Dubinin.
location: 1-0.0. references: 1930, Zh. Eksperim. Biol. 6: 300—24.
origin: Spontaneous in X chromosome carrying y. 1932, J. Genet. 26: 37-58.
discoverer: We ins tern, 16b3. 1933, J. Genet. 27: 443-64.
MUTATIONS 11

phenotype: Anterior and posterior dorsocentrals toward the anode more rapidly than those specified
removed; also thoracic hairs. A change also ap- by AdhF. As with AdhF and Adhs, three isozymes
parently induced in expression of sc; called sc13. are specified by AdhD. RK3.
The sc component also removes scutellars and cytology: Placed in region between 34E5 and 33D1,
often ocellars, postverticals, and first and second on the basis of its inclusion in Df(2L)64j =
orbital bristles. Viability low. RK2. Dt(2L)34E5-Fl;35C3-Dl (E. H. Grell).
*ac260-28
origin: X ray induced simultaneously with y260-28m AdhF; Alcohol dehydrogenase-Fast
discoverer: Sutton, 39126. origin: Naturally occurring allele.
references: 1943, Genetics 28: 210—17. discoverer: Johnson and Denniston, 1964.
cytology: Salivary chromosomes appear normal. references: 1964, Nature 204: 906—7.
Ac: see CuA Grell, Jacobson, and Murphy, 1965, Science 149:
occ: acclinal wing 80-82.
location: 1-54.5. Ursprung and Leone, 1965, J. Exptl. Zool. 160:
origin: Induced by triethylenemelamine (CB. 1246). 147-54.
discoverer: Fahmy, 1952. phenotype: Specifies isozymes of alcohol dehydro-
references: 1958, DIS 32: 67. genase that migrate Lin the system of Grell,
phenotype: Wings upheld but slope backward at 45° Jacobson, and Murphy (1965)] toward the anode
angle from abdomen. Viability and fertility good in more rapidly than the isozyme's specified by Adhs.
both sexes. RK1. Homozygote contains three electrophoretically sep-
other information: One allele each induced by arable isozymes. The one moving most rapidly to-
CB. 3007 and by CB. 3026. ward the anode is often not detected in zymograms
achaete; see ac of single adults but is nearly always detectable in
Acpfi-7A: Acid phosphatase-1-A zymograms of single larvae. The faster isozymes
location: 3-101.4. more reliably detected with use of sec-butanol than
origin: Naturally occurring allele. with ethanol as a substrate. AdhF/Adhs heterozy-
discoverer: Maclntyre, 1964. gote contains the parental isozymes plus three
references: 1966, DIS 41: 61. hybrid isozymes. Hybrid enzymes also formed in
1966, Genetics 53: 461-74. melanogaster X simulans hybrids. RK3.
phenotype: Acph-1A/Acph-1A produces acid phos-
phatase-1 enzyme that migrates slowly in starch
gel electrophoresis. Enzyme found in larva, pupa, 70 -i
and adult. RK3. - 9
Acpfi-JB 60-
origin: Naturally occurring allele.
discoverer: Maclntyre, 1964.
50- -8
references: 1966, DIS 41: 61.
1966, Genetics 53: 461-74.
phenotype: Acph-1B/Acph-1B produces more rapidly g 40 H -6
migrating acid phosphatase-1 than Acph-1A/ 5 -7
Acph-1A. Acph-1A/Acph-1B produces the two pa- o
cr ^///i/7 * ^

rental enzymes as well as a hybrid enzyme of in- ^ 30 -


termediate mobility. A hybrid enzyme is also -4
2
formed in simulans X melanogaster hybrids. RK3.
20-
ad: arco id
location: 2-60.7.
origin: Spontaneous. 10 -
discoverer: Curry, 38a2.
references: 1939, DIS 12: 45.
ORIGIN
phenotype: Wings arched, broad, and somewhat
shortened; crossveins close; scutellar groove
shallow. Legs may be slightly shorter than wild Alcohol dehydrogenase alleles
type. RK3. Adh*; b = AdhF/AdhF; c = Adh
AdhD: Alcohol dehydrogenase-D From Grell, Jacobsen, and Murphy, 2 July 1965,
location: 2-50.1 (one-tenth the distance from el be- Science 149: 80-82.
tween el and rd).
origin: Ethyl rnethanesulfonate-induced derivative of : Alcohol dehydrogenase-negative
Adh^ of Samarkand. origin: Ethyl methanesulfonate-induced derivative of
discoverer: E. H. Grell, 65k8. Adhs of Canton-S.
phenotype: Specifies isozymes of alcohol dehydro- discoverer: E. H. Grell, 66elO.
genase that migrate [in the system of Grell, phenotype: Homozygote shows no alcohol dehydro-
Jacobson, and Murphy (1965, Science 149: 80-82)] genase activity. Sensitive to alcohol, showing
12 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

evidence of intoxication within 1 hr of being adpfs: adipose-female sterile


placed on substrate containing 15 percent ethanol; origin: Spontaneous.
death invariably follows within 24 hr. Heterozygote discoverer: Counce, 1956.
with allele producing active enzyme shows evi- synonym: fs(2)adp: female sterile(2) adipose.
dence of formation of a hybrid enzyme with one ac- references: Doane, 1959, Genetics 44: 506.
tive and one mutant polypeptide subunit. RK3. 1960, J. Exptl. Zool. 145: 1-42 (fig.).
Adh»2 1961, J. Exptl. Zool. 146: 275-98.
origin: Ethyl methanesulfonate-induced derivative of phenotype: Adult fat body phenotype like adp60;
Adhs. corpus allatum hypertrophies in mated females to
discoverer: E. H. Grell, 66elO. same degree as in adp6°. Females completely
phenotype: Like Adhnl except no evidence of hybrid sterile; sterility autonomous. Eggs laid by homo-
enzyme in heterozygote with active allele. RK3. zygotes show meiotic or mitotic abnormalities, or
both, never develop beyond early cleavage stages.
origin: Ethyl methanesulfonate-induced derivative of Males 78 percent fertile. Heterozygotes fertile but
Adhs. females become sterile with age. Viability gener-
discoverer: E. H. Grell, 66f. ally good, but longevity reduced; horaozygotes
phenotype: Like Adhn2. RK3. with selective advantage under starvation; hetero-
Adh"4 zygotes superior under desiccation. Average water
origin: Ethyl methanesulfonate-induced derivative of content of well-fed adults reduced; percentage of
AdhD. lipids, as a function of dry body weight, almost
discoverer: E. H. Grell, 66g. double that of wild type. Iodine numbers show
phenotype: Like Adh^2. RK3. greater degree of saturation of mutant lipid
extracts than of wild type. RK3.
origin: Ethyl methanesulfonate-induced derivative of *ae: aeroplane
AdhD. location: 2-55.8.
discoverer: E. H. Grell, 66g. origin: Spontaneous.
phenotype: Small amount of alcohol dehydrogenase discoverer: Mohr, 26k24.
activity in homozygote but ethanol sensitive. Elec- references: Quelprud, 1931, Hereditas 15: 97-119
trophoretic migration of enzyme like that of AdhD. (fig.),
Heterozygote with fully active allele has hybrid phenotype: Wings spread, balancers drooping.
enzyme, presumably with one active and one Adhn$ Overlaps wild type. RK3.
subunit. RK3. *Ae; Aechna
Adhs: Alcohol dehydrogenase-Slow location: 3- (rearrangement).
origin: Naturally occurring allele. origin: X ray induced.
discoverer: Johnson and Denniston, 1964. discoverer: Belgovsky, 45al4.
references: 1954, Nature 204: 906—7. references: 1946, DIS 20: 63.
Grell, Jacobson, and Murphy, 1965, Science 149: phenotype: Wings spread at right angles to body
80-82. axis. Homozygous lethal. RK1A.
Ursprung and Leone, 1965, J. Exptl. Zool. 160: other information: Reduced crossing over in the th-e
147-54. region suggests presence of pericentric inversion.
phenotype: Specifies isozymes of alcohol dehydro- aeroplane: see ae
genase that [with the methods of Greil, Jacobson, *agl: angle winglike
and Murphy (1965)] migrate more slowly to the location: 1- (not located).
anode than those specified by AdhF. There are origin: Recovered among descendants of flies
also three isozymes in Adhs homozygote. RK3. treated with natural gas.
discoverer: Mickey, 49c7.
adp60: adipose synonym: Originally called angle wing, but this
location: 2-83.4. name preoccupied by ang.
origin: Spontaneous. references: 1950, DIS 24: 60.
discoverer: Doane, 1960. phenotype: Wing bent upward in middle. Overlaps
references: 1961, DIS 35: 78. wild type. RK3.
1963, DIS 38: 32. ah aristaless
phenotype: Adult fat body hypertrophies as cells location: 2-0.01 (to the right of net).
become distorted by enormous oil globules. Ab- origin: Spontaneous.
normal fat bodies visible through body wall of discoverer: Bridges, 17k7.
6-day-old and older adults when submerged in 95 references: Morgan, Bridges, and Sturtevant, 1925,
percent alcohol and then water. Adult corpus Bibliog. Genet. 2: 213 (fig.).
allatum of mated females hypertrophies. Females Stern and Bridges, 1926, Genetics 11: 510 (fig.).
fertile but egg hatchability reduced to 45—90 per- phenotype: Aristae strongly reduced. Postscutellars
cent, depending on residual genome; adult emer- widely separated, erect but strongly divergent.
gence lowered to 33—85 percent. Males viable and Scutellum shortened; sternopleurals irregular in
fertile. RK3. size, position, and number; wings slightly bowed
MUTATIONS 13

downward, narrowed, pointed; first longitudinal cytology: Associated with In(2LR)alM60, inferred
vein raised and thickened. RK1. from suppression of crossing over in most of 2L
cytology: Placed in 21C1-2 doublet on the basis of and some of 2R.
its inclusion in Df(2L)al = Df(2L)21B8-Cl;21C8-Dl *o/v: arista!ess-variegated
but not in Df(2L)S5 = Df(2L)21C2-3;22A3-4 (Lewis, origin: X ray induced.
1945, Genetics 30: 137-166). discoverer: E. B. Lewis, 1940.
0/2 references: 1945, Genetics 30: 137—66.
origin: Spontaneous. phenotype: alv/al variegated for al. Homozygous
discoverer: Stern, 26a. lethal. RK2A.
references: Stern and Bridges, 1926, Genetics cytology: Associated with In(2LR)alv =In(2LR)21B-
11: 511. C1.-41.
phenotype: Slightly less extreme than at, but vi- al-b: see aa
ability poorer. RK2. ala: see dyaJa
ala parvae: see dyata
* 0 /3 alarless: see air
origin: Spontaneous. Alcohol dehydrogenase: see Adh
discoverer: Bridges, 33g2.
phenotype: Aristae absent or much reduced. Thorax A/in: Aliesterase-negative
has wide bare area or groove down midline with location: 3- (not located).
divergent hairs and bristles; sternopleurals ab- origin: Spontaneous.
sent. Wings have weakened L2 vein and delta at discoverer: Ogita.
tip of L3. Female sterile. Viability about 10 per- synonym: ali: aliesteraseless.
cent of wild type. RK3. references: 1961, Botyu-Kagaku 26: 93—97.
1962, DIS 36: 103.
phenotype: Homozygotes practically unable to
hydrolyze methyl butyrate, whereas wild type
shows high activity; Alin/+ exhibits intermediate
activity. Homozygotes shown by Beckman and
Johnson to lack a normally present esterase that
migrates slowly on starch gel (their band F). RK3.
Alkaline phosphatase: see Aph
aliesteraseless: see Alin
almond: see Didr
almondex: see amx
almondex-55: see lzK
*o/o; alopecia
location: 1-38.3.
origin: Induced by 2-chloroethyl methanesulfonate
(CB. 1506).
discoverer: Fahmy, 1956.
references: 1958, DIS 32: 67.
al3: arrstaless-3 phenotype: Abdominal hairs much reduced in
Edith M. Wallace, unpublished. number; pigmentation frequently lighter and patchy.
Effect very pronounced in females reared at 25°C,
ah but overlaps wild type in both sexes when reared
origin: Spontaneous in In(2LR)bwvl. at a low temperature. Viability and fertility good
discoverer: Bridges, 33127. in males but reduced in females. RK3.
references: 1935, DIS 3: 5. alpha: see tyr-1
phenotype: Slight allele of al in some or all stocks *alr: alarless
of bwyl. RK2A. location: 3- (not located).
*a/3<5 origin: Spontaneous,
origin: X ray induced, discoverer: Steinberg, 40b.
discoverer: Glass, 36c. references: 1940, DIS 13: 51.
references: 1939, DIS 12: 47. phenotype: Outer postalar bristle always missing;
phenotype: Like al. RK1. posterior supra-alar missing in about 80 percent of
the flies. Anterior scutellars, humerals, and noto-
*a/AU0: art stales s of Meyer pleurals frequently duplicated. Never overlaps.
origin: X ray induced, Viability and fertility excellent. RK3.
discoverer: Meyer, 60f. Altered abdomen: see Aa
references: 1963, DIS 37: 50. Alu: Alula
phenotype: Homozygous lethal. May be variegated location: 2-54.9 (Muller places Alu to the left of
position effect. RK3A. pr and spindle attachment).
14 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

origin: Spontaneous. *amy: amethyst


discoverer: Bridges, 38al2. location: 2- (not located).
references: Curry, 1939, DIS 12: 45. discoverer: Bridges.
phenotype: Heterozygote has alula fused to main references: Morgan, Bridges, and Sturtevant, 1925,
wing; wings often bent, broader. May overlap wild Bibliog. Genet. 2: 218.
type, but intensified by cold and by heterozygous phenotype: Transparent light-purplish eye color.
ds, with buckling effect increased. Homozygote at RK3.
19°C shows extreme buckling owing to rotation of Amy1: Amylase-1
wing and alula. Homozygote viable and resembles location: 2-77.3 (Doane, 1963, DIS 38: 32).
heterozygote. RK2. origin: Naturally occurring allele.
*AluS6c discoverer: Kikkawa, 1957.
origin: Spontaneous (arose with ltS6c). synonym: Amy+.
discoverer: Meyer, 56c. references: Kikkawa and Abe, 1960, Annotationes
references: 1956, DIS 30: 77. Zool. Japon. 33: 14—23.
phenotype: Similar to Alu. RK2. Kikkawa, I960, Japan. J. Genet. 35: 382—87.
Kikkawa and Ogita, 1962, Japan.J. Genet. 37:
*o/w: arclike wing 394-95.
location: 2- (near 6). Kikkawa, 1963, DIS 37: 94.
discoverer: Sturtevant, 1948. 1964, Japan J. Genet. 39: 401-11 (fig.).
references: 1948, DIS 22: 55. phenotype: Specifies amylase isozyme system
phenotype: Wings evenly bent downward at tips. having, in agar gel electrophoresis, one major and
Overlaps wild type. RK2. one minor component. The major isozyme moves
am: see Didr rapidly toward the anode and occupies position 1.
The minor component migrates immediately behind
*amb: amber it to position 2. Heterozygotes of Amy alleles con-
location: 1-6.8. tain isozymes of both parents. RK3.
origin: Induced by triethylenemelamine (CB. 1246).
discoverer: Fahmy, 1950.
references: 1958, DIS 32: 67.
phenotype: Pale yellow body color; bristles very
thin and short; hairs less affected. Eyes slightly
brighter red. Males sterile. Viability 10—50 per-
cent wild type. RK2.
other information: One allele each induced by
CB, 1246, CB. 3007, CB. 1506, CB. 1414. Two
alleles induced by CB. 3034.
amb2
origin: Induced by L~p-NN-di(2-chloroethyl)amino-
phenylalanine (CB. 3025). 6
discoverer: Fahmy, 1954. 7
references: 1958, DIS 32: 67.
phenotype: Pale yellow body color; bristles slender Origin!—)
1.2 1.3 1.4 1.6 2.6 3.6 46
and only slightly shortened. Male viability and
fertility good; females viable but sterile. RK2. Amylase Alleles
amethyst: see amy Electrophoretic patterns of homozygotes for the various
alleles of Amy. Amy*'2 is on acrylamide gel and the
amx: almondex rest are on agar gel.
location: 1-27.7 [to the left of lz (Green and Green,
1956, Z. Induktive Abstammungs- Vererbungslehre -2
87: 708-21)]. origin: Naturally occurring allele.
origin: X ray induced. discoverer: Doane, 64e6.
discoverer: Ball, 32k20. phenotype: Major Ot-amylase isozymes occupy posi-
phenotype: Eyes slightly reduced, narrower below. tions 1 and 2 in acrylamide gel disc electropho-
Trident pattern stronger than in lz. Homozygous resis (corresponding to positions on agar gel). A
females highly infertile; all progeny that do occur minor component is present at position 0. (In acryl-
are daughters. Infertility does not resemble that of amide gels, minor components migrate more rapidly
lz females, since amx has no effect on the gen- to the anode than major isozymes. In agar gels
italia [Anderson, 1945, Genetics 30: 280-96 (fig.)]. they migrate less rapidly). The total a-amylase
Iz-'mmx is wild type. RK2. activity is quite high, being intermediate between
cytology: Located in 8D (region 8D4 through 8E2) Amy4-6 and Amy1-3. RK3.
by Green mnd Green (1956).
origin: Naturally occurring allele.
MUTATIONS 15

discoverer: Kikkawa. Amys: see Amy2-6


references: 1964, Japan. J. Genet. 39: 401—11 (fig.). Amywh: see Amy1-4
phenotype: Specifies major amylase isozymes that an: an con
occupy positions 1 and 3 after agar gel electropho- location: 2-44 (34-54).
resis and minor components at positions 2 and 7. discoverer: Bridges, 30e3.
RK3. phenotype: Wings and legs somewhat short. Over-
laps wild type. RK3.
origin: Naturally occurring allele. an2
discoverer: Kikkawa. discoverer: Bridges 30c25.
synonym: Amywh; Amy4. phenotype: Wings broad and short. Legs short and
references: 1963, DIS 37: 94. gnarled. Bristles on abdomen straggly; sclerites
1964, Japan.J. Genet. 39: 401-11 (fig.). etched. Eyes small and roughish. an2'/an is like
Doane, 1966, DIS 41: 93. an2. Overlaps wild type. RK3.
phenotype: Specifies major amylase isozymes that anarista: see era
occupy positions 1 and 4 and a minor component at oncon: see an
position 5 after agar gel electrophoresis. The ang: angle wing
isozyme at position 1 was originally considered to location: 2-10.5.
be minor, but Doane considers it major; most pro- origin: Spontaneous.
nounced in young flies. RK3. discoverer: Mittler and Goldberg, 48il6.
Amyl-6 references: Mittler, 1950, DIS 24: 61.
origin: Naturally occurring allele. phenotype: Wings held up from dorsal surfaces and
discoverer: Kikkawa. extended outward 15—90° from the mid-dorsal line.
references: 1964, Japan. J. Genet. 39: 401-11 (fig.). Longitudinal dorsal median muscles 5 and 6 fused
phenotype: Specifies major amylase isozymes that (Goldberg, 1954, Ph.D. Thesis, 111. Inst.
occupy positions 1 and 6 and minor components at Technol.). No increase in expressivity with
positions 2 and 7 after agar gel electrophoresis. temperature. Does not overlap wild type. RK2.
RK3. ang: see ano
Amy* & angle wing: see ang
origin: Naturally occurring allele. angle wing: see agl
discoverer: Kikkawa. angle winglike: see agl
synonym: AmyB. *ano: anomogenitals
references: Kikkawa and Abe, 1960, Annotationes location: 1-35.7.
Zool. Japon. 33: 14-23. origin: Induced by triethylenemelamine (CB. 1246).
Kikkawa, 1960, Japan.J. Genet. 35: 382-87. discoverer: Fahmy, 1952.
Kikkawa and Ogita, 1962, Japan. J. Genet. 37: synonym: Originally symbolized ang, but this
394-95. symbol was preoccupied.
Kikkawa, 1963, DIS 37: 94. references: 1958, DIS 32: 67.
Kikkawa, 1964, Japan. J. Genet. 39: 401-11 (fig.). phenotype: Many bristles on head and thorax either
phenotype: Specifies major amylase isozymes that reduced in size or absent. Thoracic and abdominal
occupy positions 2 and 6 and minor components at hairs appreciably fewer. External male genitalia
positions 3 and 7 after agar gel electrophoresis. invariably abnormal, sometimes completely absent.
RK3. Melanized exudate frequently present in furrow
Amy* 6 between mesonotum and scutellum near anterior
origin: Naturally occurring allele. scutellar bristles. Males sterile; viability less
discoverer: Kikkawa. than 10 percent wild type. RK3.
references: 1964, Japan. J. Genet. 39: 401-11 (fig.).
phenotype: Specifies major amylase isozymes that ant: antennaless
occupy positions 3 and 6 and minor components location: 2- (not located).
that occupy positions 4 and 7 after agar gel elec- origin: Spontaneous.
trophoresis. RK3. discoverer: Gordon, 1936.
Amy4: see Amy1-'* references: 1941, DIS 14: 39.
1941, Proc. Intern. Congr. Genet., 7th. p. 131.
origin: Naturally occurring allele. Gordon and Sang, 1941, Proc. Roy. Soc. (London),
discoverer: Kikkawa. Ser. B 130: 151-84 (fig.).
synonym: Amyad. Vogt, 1947, Biol. Zentr. 66: 388-95 (fig.).
references: 1963, DIS 37: 94. phenotype: Antennae missing on one or both sides.
1964, Japan.J. Genet. 39: 401-11 (fig.), Expression affected by residual genotype, nutri-
phenotype: Specifies major amylase isozymes that tional environment, and temperature. Time of ac-
occupy positions 4 and 6 and minor components at tion about 70 hours after hatching LBegg and Sang,
positions 5 and 7. RK3. 1945, J. Exptl. Biol. 21: 1-4 (fig.)]. Used in ex-
Amy*: see Amy1 periments to locate chemoreceptors IBegg and
Amyad: see Amy4-6 Hogben, 1946, Proc. Roy. Soc. (London), Ser.
GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

B 133: 1—19j and in studies of mating behavior phenotype: Arista tends to be transformed into
(Begg and Packman, 1951, Nature 168: 953). RK3. tarsus; third antennal segment hypertrophied and
deformed. Ocelli reduced in size and number. Ce-
Antp49: Antennapedia phalic capsule deformed. Head bristles reduced in
location: 3- (to the left of p; probably to the right number. Wings held at 45° angle from midline. Ex-
of st; determined for Antp50 by Hannah). pression variable. Homozygous lethal. RK3A.
origin: X ray induced. cytology: Associated with In(3R)AntpL>c =
discoverer: Piternick, 1949. In(3R)84A5-6; 92A5-6.
synonym: Antp4703.
phenotype: Antennae transformed into second legs
plus some differentiation toward first legs (Hannah-
Alava). Lethal in combination with AntpYu,
AntpB, Antp50, and Sex. Possibly lethal with Pc
but semilethal with P c 2 (Hannah-Alava). Quite
variable. Homozygous lethal. RK3A.
cytology: Probably in region 84A (or 83F), based
on Lewis's analysis of AntpB and AntpYu. Ant49
associated with small cytological abnormality of
undetermined nature in 83EF-84AB (Hannah-Alava).
AntpSO
origin: X ray induced.
discoverer: Piternick, 1950.
synonym: Antp4715.
phenotype: Variable transformation of entire an- AntpLC: Antennapedia of Le Calvez
tenna into a leg may occur, but effect is often From Le Calvez, 1948, Bull. Biol. France Belg. 82:
limited to slight elongation of third antennal seg- 97-113.
ment. In compounds with Pc and Pc2, the rather
well developed antennal second legs show some AntpR: Antennapedia of Rappaport
transformation into first legs (Hannah-Alava). origin: X ray induced.
Homozygous lethal and lethal in combination with discoverer: Rappaport, 1963.
AntpB, AntpYa, Antp49, and Sex (Hannah-Alava). synonym: s s ^ : spineless-Aristapedia Dominant.
RK3A. references: Falk, 1964, DIS 39: 60.
Antp4?0*: see Antp49 phenotype: Segments added to antennae, usually
Antp4?15: see Antpso distal to aristae. Claw occasionally at end of
antenna. Asymmetry pronounced. Rarely an an-
AntpB: Antennapedia of Bacon tennalike organ on sternopleura. Variable expres-
origin: X ray induced. sion, but expressivity 100 percent in combination
discoverer: Bacon, 50g. with D. Homozygous lethal; lethal in combination
references: Lewis, 1956, DIS 30: 76. withAntpB (Von Halle). RK3A.
phenotype: Antenna partially leg-like, but less ex- cytology: Associated with In(3R)AntpR =
treme than AntpYu and may overlap wild type. En- ln(3R)83F;86C (Ben-Zeev).
hanced by Pc (and Pc2"). AntpB ssa/+ ssa has AntpYu: Antennapedia of Yu
virtually complete antennal leg including coxa, origin: X ray induced.
tibia, femur, and tarsus, but no sex comb in the discoverer: Yu, 1948.
male (B. Holloway). The antennal second leg with reference: 1949, Ph.D. Thesis Calif. Inst. Technol.
some transformation into a first leg completely Lewis, 1956, DIS 30: 76.
developed only in Pc (or Pc2) ssa/AntpB s s a phenotype: Antenna transformed into second leg
compounds. Lethal with AntpYu, Antp49, Antp50, plus some differentiation toward a first leg but
and Sex (Hannah and Stromnaes, 1955, DIS 29: with recognizable arista usually present; not like
121-23 and Hannah-Alava). RK3A. ssa, in which main effect is that the arista be-
cytology: Associated with In(3R)AntpB = comes tarsus-like. Strongly enhanced by Pc and
ln(3R)84A;85E, but apparently mutant and inversion Pc2. Pc +/+ AntpYtl has a pair of excellent an-
are separable (Hannah-Alava). tennal legs complete with tarsae and with sex
combs in the male. Lethal with AntpB, Antp49,
*AntpL-C; Antennapedia of Le Calvez Antp50, and Sex (Hannah and Stromnaes, 1955,
origin: Neutron induced. DIS 29: 121—23; Hannah-Alava). RK3A.
discoverer: Le Calvez. cytology: Associated with T(2;3)AntpYu =
synonym: Ar: Aristapedia; SS^T. T(2;3)22B;83E-F + T(2;3)38E;98A.
references: 1948, Compt. Rend. 226: 123-24. *ap: apterous
1948, Bull. Biol. France Belg. 82: 97-113 (fig.). locotion: 2-55.2.
1948, Arch. Anat. Microscop. Morphol. Exptl. 37: origin: Spontaneous.
50-72. discoverer: E. M. Wallace, 13h.
MUTATIONS 17

references: Bridges and Morgan, 1919, Carnegie references: Meyer, Edmondson, Byers, and
Inst. Wash. Publ. No. 278: 236 (fig.). Erickson, 1950, DIS 24: 59.
Metz, 1914, Am. Naturalist 48: 675-92. phenotype: Compared with ap4 and very similar.
phenotype: Wings and halteres reduced to traces. Almost lethal. RK2.
Bristles eliminated from area around wing base ap
(including posterior notopleurals, anterior, and origin: Spontaneous.
posterior supra-alars, anterior postalars); posterior discoverer: Faulhaber.
scutellars erect when present, but missing in first references: 1963, DIS 37: 48.
counts; dorsocentrals smaller and fewer; hairs on phenotype: Wings vary from clublike to straplike;
thorax sparse and irregular. Sutural furrow re- seldom exceed 30 percent of normal length; lack
duced; thorax disproportionately small. Flies veins and specific bristles. Halteres 25—50 percent
small, pale, weak, and very short-lived. Viability normal length. Postalar, scutellar, and dorsocentral
about 70 percent that of wild type, but erratic. bristles missing. Hind legs sometimes deformed.
Both sexes sterile. RK2. Few ap6/ap6 or ap4/ap6 females survive past 5
cytology: Placed in salivary region 41B-C (Schultz). days; those that do are slightly fertile, the re-
mainder are not (Butterworth and King), ap6/
origin: Spontaneous. M(2)S24 more extreme than ap6/ap6 but some fe-
discoverer: Bridges, 16j20. males fertile and thoracic chaetotaxy more nearly
synonym: ap-c. normal. RK2.
references: 1919, J. Exptl. Zool. 28: 370.
Bridges and Morgan, 1919, Carnegie Inst. Wash. origin: Spontaneous.
Publ. No. 278: 237. discoverer: Ritterhoff, 49j.
Medvedev and Bridges, 1935, Tr. Inst. Genet. references: Glass, 1951, DIS 25: 76-77.
Akad. NaukSSSR 10: 199-209. phenotype: Appears to be somewhat less extreme
phenotype: Like ap but less viable. RK2. than ap4, with which it was compared. Wings and
halteres reduced to vestiges. Bristles and hairs on
origin: Spontaneous. sides absent, including posterior notopleurals,
discoverer: Morgan, 23a. anterior and posterior supra-alars, and anterior
synonym: no-wings; later, ap~c. postalars, but dorsocentrals not reduced in size
references: 1929, Carnegie Inst. Wash. Publ. No. and number, and one or a pair of pre-anterior dor-
399: 183. socentrals may be present. Posterior scutellars
phenotype: Like ap. RK2. not erect when present. Sutural furrow normal;
thorax of normal size. No adults live longer than 3
discoverer: Medvedev, 32al5. days; larval adipose cells persist in adult, and
references: Medvedev and Bridges, 1935, Tr. Inst. adult adipose tissue fails to develop. ap49i/
Genet. Akad. Nauk SSSR 10: 199-209. M(2)S24 adult sterile, short lived, and has abnormal
Beatty, 1949, Proc. Roy. Soc. Edinburgh, adipose tissue and short wing rudiments (Butter-
B 63: 249-70 (fig.). worth and King). Both sexes sterile. RK2.
King and Sang, 1958, DIS 32: 133. other information: Interacts with apXa but not
synonym: ap-d. tested in combination with ap4 or apblt.
phenotype: Wings mostly less than 10 percent normal apSSf
length and lacking veins and specific hairs. Hal- origin: Spontaneous.
teres less than 25 percent normal length, and fre- discoverer: Thompson, 56f.
quently absent. Scutellar and dorsocentral bristles references: Burdick, 1956, DIS 30: 69.
sometimes missing (Butterworth and King). Adults phenotype: Wings club-shaped, 10—30 percent normal
become paralyzed with age and die within 4 days. length, and lack veins and certain types of hairs.
Larval adipose cells persist in imago, and adult Scutellar and dorsocentral bristles missing (Butter-
adipose tissue fails to develop. Female sterile worth and King). Rear and middle legs occasionally
with underdeveloped ovaries; nurse cell nuclei be- twisted, more frequently in female than in male.
come pycnotic after stage 7, and yolk formation is Both sexes fertile when homozygous and in combi-
never initiated (King and Burnett, 1957, Growth 21: nation with other ap alleles. ap56l/M(2)S24 have
263—80). ap4 ovaries develop normally when trans- normal complement of dorsocentral and scutellar
planted into a normal host (King and Bodenstein, bristles (Butterworth and King). RK2.
1965, Z. Naturforsch. 20b: 292-97). Male sterile,
but testes appear normal with motile sperm (King apbh; apterous-blot
and Sang, 1958). ap4/M(2)S24 adult has nearly origin: Spontaneous.
normal complement of bristles but otherwise re- discoverer: Groscurth, 31bl.
sembles ap4 homozygote (Butterworth and King). synonym: bit.
RK2. phenotype: Wings blistered, inflated, often dark be-
ap- cause of dried blood. In extreme cases, a small
origin: Ultraviolet induced. mirror image wing forms by partial twinning of
discoverer: Byers, 49f. wing in third posterior wing cell. According to
18 GENETIC VARIATIONS OF DROSOPHILA MELANOCASTER

Waddington (1939, Proc. Natl. Acad. Sci. U.S. 25: cytology: Shown by Sturtevant (1934, DIS 2: 19) to
299—307), the fundamental effect is partial be associated with T(2;3)apXa = T(2;3)41F;
twinning of wing blade, which leads to difficulties 89E8-F1 which is superimposed on In(2R)Cy and
in clearance of heraolymph after inflated stage. In(3R)P (Morgan, Bridges, and Schultz, 1936,
Much overlapping with wild type. RK3. Carnegie Inst. Wash. Year Book 35: 294; Lewis,
1951, DIS 25: 109).
*apbtt2
origin: Spontaneous.
discoverer: Whittinghill, 44h.
synonyms: bit2.
references: 1947, DIS 21: 71.
phenotype: More extreme than apblt. Wings always
shorter than normal and inflated. Strong tendency
for unequal bifurcation of wing. Wings often break
off and remain attached to pupa cases, hence a
wingless phenotype. Viability about 20 percent of o p * 0 ; apterous-Xasfa
wild type in both sexes. Males fertile but females From Bridges and Brehme, 1944, Carnegie Inst. Wash.
sterile. RK2. Publ. No. 552: 228.
ap blt3
origin: Spontaneous, ap-c: see ap2
discoverer: Semenza, 49k. ap-c: see ap3
synonym: bltS49k. ap-d: see ap4
references: Barigozzi, 1950, DIS 24: 54. Apart: see Apt
phenotype: Wings uniformly inflated, more extreme
than apbtt. Does not overlap wild type. RK2. *apb: apterblister
location: 2-44.7.
apT60: apterous of Thomas origin: Ultraviolet induced.
origin: X ray induced. discoverer: Edmondson, 49K.
discoverer: Thomas, 60g. references: Meyer, Edmondson, Byers, and
references: Meyer, 1963, DIS 37: 50. Erickson, 1950, DIS 24: 59-60.
phenotype: Wings straplike; about 30 percent normal phenotype: Wings always notched, nearly always
length. Adult survives past fifth day. apT6G/ spread and usually blistered, but expression some-
M(2)S24 female lays eggs (Butterworth and King). what variable. Homozygous imagos live less than
RK2. 24 hours, owing to intestinal constrictions that
apXo; apterous-Xasta prevent defecation. Abdomens characteristically
origin: X ray induced in In(2R)Cy; In(3R)P. (The turn dark grey before death, because of accumula-
first X-ray-induced mutation recovered in the tion of digested food products. Although not at
USSR.) same locus as ap, apb +/+ ap4 flies show slight
discoverer: Serebrovsky, 28a. notching of wings and many die within a day;
synonym: Xa. those that survive are fertile, ap5 gives a similar
references: Serebrovsky and Dubinin, 1930, J. heterozygous effect. RK2.
Heredity 21: 259-65. apexless: see apx
phenotype: Wings reduced in length to about 70 per-
cent normal; irregular in outline with a V-shaped Aph°: Alkaline phosphatase deficient
incision with apex at L2, uniformly present, giving location: 3-46.3 (Maclntyre).
wing a mittenlike shape with the thumb between origin: Spontaneous.
marginal vein and L2. Excellent dominant with no discoverer: Johnson.
overlap. Fertile and fully viable in heterozygote. references: 1966, DIS 41: 157-58.
Usually lethal in homozygous conditions, but 1966, Science 152: 361-62.
occasionally ecloses very late as pale dwarf with phenotype: Homozygous larva has no detectable
wings and balancers like vg. Waddington reports alkaline phosphatase activity. Aph°/Aphp larva
deep notch visible in tip of wing fold in prepupa has alkaline phosphatase, which migrates in starch
(1939, Proc. Natl. Acad. Sci. U.S. 25: 299-307; gel electrophoresis to same position as the band in
1940, J. Genet. 41: 75-139 (fig.)]. In homozygotes AphF homozygote. Aph°/Aphs larva has bands of
and in combination with ap4, ®p6, or M(2)S24 wings activity at the Aphs position and at a position
are straplike and 30—70 percent normal length, and slightly faster than the AphF/Aphs hybrid band.
hattere length is 25—50 percent normal; longevity RK3.
fertility like ap4/ap4 except for an occasional long- AphF: Alkaline phosphatase-Fast
lived apx*/M(2)S24 female that may be fertile origin: Naturally occurring allele.
(Schultz; Butterworth and King). apXa'apbtt has discoverer: Beckman and Johnson.
combined pbenoiypes of ap^*,-+ and apblt/apbit references: 1964, Nature 201: 321 (fig.).
(Schultr). RK1A. 1964, Genetics 49: 829-35 (fig.).
MUTATIONS 19

phenotype: AphF/AphF larvae produce an a l k a l i n e Rarely L4 and 5 are interrupted. Males viable and
phosphatase that migrates rapidly in s t a r c h gel fertile; female fertility reduced. RK3.
electrophoresis under conditions d e s c r i b e d by err; abdomen rotatum
Beckman and Johnson (1964). Alkaline phospha- location: 4- (proximal to bt; Fung and Stern, 1951,
tase produced by pupae migrates faster than larval Proc. Natl. Acad. Sci. U.S. 37: 403-4).
enzyme. No enzyme demonstrable in a d u l t s . RK3. origin: Spontaneous.
Aphs: Alkaline phosphatase-Slow discoverer: Beliajeff, 1926.
origin: Naturally occurring allele. references: 1931, Biol. Zentr. 51: 701-8 (fig.).
discoverer: Beckman and Johnson, Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
references: 1964, Nature 201: 321 (fig.). Marengo and Howland, 1942, Genetics 27: 604—11
1964, Genetics 49: 829-35 (fig-)- (fig-)-
phenotype: Alkaline phosphatase of Aph^/Aph^ phenotype: Abdomen twisted clockwise through 45°
migrates more slowly in starch gel electrophoresis to 60°. No overlapping with wild type. Male ex-
than that of AphF/AphF. AphF/Aphs larvae pro- ternal genitalia often missing. Males usually
duce a hybrid enzyme of intermediate mobility as sterile; females partially fertile. Puparia not so
well as the fast and slow forms. RK3. smooth as normal; larval segmentation remains.
opp: approximated Puparia have deep constriction near posterior end
location: 3-37.5. just anterior to spiracles. Existing chromosomes
discoverer: Curry, 34a25. marked ar also carry 1(4) and in combination with
references: 1935, DIS 3: 6. Df(4)M show counterclockwise rotation of male ab-
phenotype: Crossveins close together; veins di- domen (Hochman). RK2.
verge at greater angle than wild type; effect vis- cytology: Placed in salivary chromosome region
ible in prepupal wing [Waddington, 1940, J. Genet. 101E through 102B16, on basis of its inclusion in
41: 75—139 (fig.)J. Legs short, with four-jointed Di(4)M =Di(4)101E-F;l02B6-17.
tarsi; the penultimate joint characteristically *ar2
swollen [Waddington, 1939, Growth Suppl. 37—44 origin: Spontaneous.
(fig.)J. Thickset body. Posterior scutellars farther discoverer: Nichols-Skoog, 34el7.
apart than normal. Eyes smaller and flatter than references: Bridges, 1935, Biol. Zh. (Moscow)
normal, also bumpy. Spread wings and thickened 4: 401-20.
veins. RK1. phenotype: Abdomen twisted; male genitalia often
*app61» missing. RK2.
arS7d
origin: X ray induced.
discoverer: Puro, 61e. origin: X ray induced,
references: 1964, DIS 39: 64. discoverer: Gloor, 57d.
phenotype: Slightly more extreme than app. RK1. phenotype: Abdomen twisted counterclockwise as
apr: see wa viewed from behind. RK2.
*Apt: Apart
location: 3- (between h and p). origin: X ray induced.
origin: X ray induced. discoverer: Gloor, 57g.
discoverer: Belgovsky, 34e23. phenotype: Abdomen twisted counterclockwise as
references: 1935, DIS 3: 27. viewed from behind. RK2.
phenotype: Wings spread widely. Viability, fertility, At: see Antp1* c
and separability good. Homozygous lethal. RK2A.
cytology: Associated with In(3L)Apt — no salivary arc: see a
analysis, arch: arch
other Information: Apt/D survive; therefore not an location: 2-60.5.
allele of D. origin: Spontaneous.
apterblister: see apb discoverer: Curry, 36g3.
apterous: see ap references: 1937, DIS 7: 5.
*apx: op ex/ess phenotype: Wings curved evenly downward both
location: 1-11.3. longitudinally and transversely, sometimes shorter
origin: Induced by DL-p-NN-di(2-chloroethyl)amino- and blunter, rarely divergent. RK2.
phenylalanine (CB. 3007). arclike wing: see cr/w
discoverer: Fahmy, 1954. arcoid: see ad
references: 1959, DIS 33: 83. arctops: see at
phenotype: Slightly larger fly with large eyes con- arctus ocu/us: see at
taining various numbers of deranged ommatidia. Argentine Curly: see CuA
Wings broad and blunt and in many flies margin Arista: see Ata
removed to various degrees, from a small incision aristaless: see al
of inner margin to removal of entire inner margin, aristaless-h: see aa
costal vein, and parts of the membrane, as far as Aristapedia: see AntpLC
L3. Region from L3 to costal cell unaffected. arp-1: see ssa$P
20 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

*as: as cute
location: 3-46. origin: Spontaneous in [n(2L)Cy.
origin: Spontaneous. discoverer: E. B. Lewis.
discoverer: Bridges, 16J21. references: 1945, Genetics 30: 137-66.
references: Bridges and Morgan, 1923, Carnegie phenotype: Similar to ast, but wing veins normal.
Inst. Wash. Publ. No. 327: 170. S/ast3 hatches late; has normal wing veins and
phenotype: Front of scutellum elevated, with partial small eyes similar to but slightly larger than
obliteration of transverse furrow; deep chested. S/ast. RK1A.
Bubble in scutellum or midline of thorax; dried cytology: Same as ast?,
black exudate often at each side of scutellum, may ast*
appear at any of the sutures of head and thorax; origin: Spontaneous recombinational derivative of
black deformed lump behind cheek. Wings droop at ast/ast.
sides. Overlaps wild type. RK3. discoverer: E. B. Lewis.
*as* references: 1945, Genetics 30: 137-66.
origin: Spontaneous. phenotype: ast4/ast4 is usually wild type. S/ast4
discoverer: Bridges, 18116. has smaller eye than S/+; resembles ast/ast in
references: Bridges and Morgan, 1923, Carnegie wing phenotype. RK3.
Inst. Wash. Publ. No. 327: 172. cytology: Salivary chromosomes normal.
phenotype: Same as as. RK3. other information: Recovered as an ast4 ho single
as^9: ascute-hUngende recombinant from an al ast ho/ast female.
origin: Spontaneous,
discoverer: Franke. origin: Spontaneous nonrecombinational derivative
references: 1934, DIS 2: 9. of ast/ast.
Geottschewski, 1935, DIS 4: 15. discoverer: E. B. Lewis,
phenotype: Wings held laterally downward, ends references: 1945, Genetics 30: 137—66.
occasionally resting on legs; eyes small and phenotype: Resembles ast4. RK3.
knobby. RK2. *astr v 1: astero id- re verted
origin: X ray induced in a/ ast ho.
ascutex: see asx discoverer: E. B. Lewis, 1942.
ast: asteroid references: 1945, Genetics 30: 158.
location: 2-1.3 (0.02 unit to right of S). phenotype: Wild type in most combinations, except
origin: Spontaneous. that Df(2L)S4/astrvl slightly more extreme than
discoverer: E. B. Lewis, 38b. Df(2L)S4/+; S** and SM slightly less extreme when
synonym: Sr: Star-recessive. heterozygous with astrvl than with wild type. Ho-
references: 1938, DIS 10: 55. mozygous lethal. RK3A.
1942, Genetics 27: 153-54. cytology: Associated with T(2;3)astrvl =
1945, Genetics 30: 137-66. T(2;3)21E2-3;68C2-3;88D8-9.
1951, Cold Spring Harbor Symp. Quant. Biol. 16: *astry2
159-74 (fig.). origin: X ray induced in al ast ho.
phenotype: Eyes small and rough. Veins L2, L3, discoverer: E. B. Lewis, 1942.
L4, and L5 do not always extend to margin. Over- references: 1945, Genetics 30: 158.
laps wild type rarely. S +/+ ast has very small phenotype: Like astrvl; astrv2/SM overlaps wild
eyes with fused facets; veins L2 to L5 incomplete type. RK3A.
at tip. S ast/+ ast has slightly larger eye than cytology: Associated with ln(2L)astrv3 =
S +/+ ast. S ast/+ + resembles S +/+ +. S +/+ ast In(2L)21E2-3;31.
and ast/ast partially suppress px and net. Eyes of *asfv3
ast/E(S) rough. RK2. origin: X ray induced in net ast dp c/.
cytology: Placed in the 21E1-2 doublet on the basis discoverer: E. B. Lewis, 1942.
of its being included in the synthetic deficiency references: 1945, Genetics 30: 158.
derived by combining the Y-centric portion of phenotype: Wild type in all combinations, except
T(Y;2)21E = T(Y;2)21D4-E1 and the 2-centric por- that S/astrv3 is slightly more extreme than S/+,
tion of T(2;4)astv = T(2;4)2lE2-3;101 (E. B. Lewis, Lethal homozygous and in combination with
1945). Dt(2L)S4 = Dt(2L)21C3-4;22B2-3. RK3A.
cytology: Associated with T(2;3)astrv3 =
origin: Spontaneous in ln(2L)Cy. T(2;3)21E2-3;61C2-3.
discoverer: E. B. Lewis.
references: 1945, Genetics 30: 137-66. asty: asteroid-variegated
phenotype: Similar to ast, but wing veins normal. origin: X ray induced.
S/ast2 lethal. Heterozygote strongly enhanced by discoverer: E. B. Lewis, 1940.
E(S). RK2A. references: 1945, Genetics 30: 137—66.
cytology: Normal except for presence of In(2L)Cy = phenotype: astv/ast and astv/S more variable than
In(2L)22Dl-2;33F5-34Al. but similar to ast/ast and ast/S, respectively;
MUTATIONS 21

suppressed in X/X/Y female. Homozygous lethal. origin: X ray induced.


astv/Df(2L)S2 lethal, RK1A. discoverer: Krivshenko, 1949.
cytology: Associated with T(2;4)ast^ = T(2;4)21E2- synonym: At (symbol preoccupied).
3; 101. references: 1954, DIS 28: 74-75.
astx: asteroid from X irradiation 1955, DIS 29: 73.
origin: X ray induced simultaneously with Sx. phenotype: Lateral branches of aristae reduced,
discoverer: E. B. Lewis. especially branches extending upward from central
references: 1945, Genetics 30: 137—66. axis and situated at base of arista. Axis of arista
phenotype: astx/astx is wild type, but behaves as often abnormal. Wings have small transparent spots
a very slight ast allele in compounds with S and distally. Homozygous lethal. Heterozygous via-
Sx astx. RK3. bility and fertility comparatively high. RK2A.
other information: Separated from Sx by crossing cytology: Associated with T(2;3)Ata =
over in a Sx astx/ast4 ho female. T(2;3)40;66F-67A + T(2;3)47;81.
*osx: ascutex Attenuated: see At
location: 1-26. augenwulst: see awu
origin: Spontaneous. *aw: awry
discoverer: Bridges, 24bl4. location: 1-32 (not allelic to wy).
references: Morgan, Bridges, and Sturtevant, 1925, origin: Induced by ingested radiophosphorus.
Bibliog. Genet. 2: 218. discoverer: Bateman, 1949.
phenotype: Furrow between scutellum and thorax references: 1950, DIS 24: 54.
much shallower, scutellum inflated. Body color 1951, DIS 25: 77.
pale. Legs have blackened leaky joints. Char- phenotype: Wings upcurled, slightly wavy, convex,
acter less extreme in old dry cultures. Viability opaque, or vestigial-like. Variable; overlaps wild
60 percent wild type. RK3. type. Viability about 50 percent wild type. Not
Asy: see A-p enhanced in presence of y as is dvr (1-28.1). RK3.
Asymmettid: see A-p *aw-b: awry-b
*at: arctus oculus location: 1-38 to 39.
location: 2-60.1. origin: Induced by ingested radiophosphorus.
origin: Spontaneous. discoverer: Bateman, 1950.
discoverer: Fernandez Gianotti, 42g28. synonym: aw2.
synonym: bar eye; arctops. references: 1950, DIS 24: 54.
references: 1943, DIS 17: 48. 1951, DIS 25: 77.
1944, DIS 18: 45. phenotype: Like aw. Good expression at 25°C.
1945, Rev. Inst. Genet. Fac. Agron. Vet. Univ. Viability 10 percent that of wild type. Most males
Buenos Aires 2(14): 171-77. fail to eclose. RK3.
1948, DIS 22: 53. awry: see aw
phenotype: Eyes similar to B but with more facets. *awu: augenwulst
Classification, fertility, and viability excellent. location: 2-57.
RK1. origin: Spontaneous.
At: Attenuated discoverer: Rosin, 1951.
location: 1- (in the B region). references: Volkart, 1959, DIS 33: 100.
origin: Induced with soft X rays in phenotype: Eyes deformed; in most extreme expres-
In(l)scSiL-sc8R+dl-49, scsi scsB; associated with sion, deeply indented at middle of anterior margin,
loss of B phenotype. where invaginating integument forms a padlike
discoverer: Valencia and Valencia, 1949. swelling with bristles. Expression variable, often
references: 1949, DIS 23: 64. asymmetrical. Overlaps wild type. Heterozygote
phenotype: In A t/+ females, wings incised medially occasionally has minor effects. Good viability.
and laterally; usually have one large central RK3.
blister. At/At females have badly crumpled, Ax; Abruptex
blistered, and sometimes poorly developed wings. location: 1-3.0.
Wings of At males tend to be more like those of origin: Spontaneous.
At/+ females, although many fall somewhere be- discoverer: Nazarenko, 28a.
tween Af/-s- and At/At in phenotype. Thus there is references: 1930, Biol. Zentr. 50: 385-92 (fig.)-
evidence for only slight dosage compensation for Mohr, 1932, Proc. Intern. Congr. Genet., 6th. Vol.
At. This mutant is similar to some Beadex alleles, 1: 190-212 (fig.).
but allelism with Bx difficult to determine and has phenotype: Homozygous female and male show
not been tested for. Both males and homozygous shortened L5 vein, usually also L4, L2, and some-
females viable and fertile. RK1A. times L3. Wings shortened, arched, thin. Costal
cytology: Associated with ln(l)At ~In(l)16A4- bristles clumped and frayed; costal veins thick-
5;18C4-6;20A2-3. ened. Thorax shows midfurrow with rearranged
*Ata: Arista hair directions; hairs on thorax and head fewer,
location: Not located. with clear patches and streaks. Male genitalia
22 GENETIC VARIATIONS OF DROSOPHiLA MELANOGASTER

often rotated. Ax/+ female shows short L5 in half


of the flies and sparse hair pattern on thorax. discoverer: Nichols-Skoog, 36fl.
Lower temperature (19°C) markedly decreases references: 1937, DIS 7: 5.
expression, and higher temperature enhances it. phenotype: Like b. RK1A in aged flies.
Ax/N8 approaches wild type in all characteristics. cytology: Inseparable from T(2;3)'dp, possibly posi-
Enhanced by H so that Ax/Y; H/+ and Ax/Ax; tion effect or deficiency caused by break distal to
H/+ are nearly lethal at 26° (House, 1959, Anat. 34D. Leads to some ambiguity regarding cytolog-
Record 134: 581-82). RK2 in males. ical location of b.
cytology: A single-band duplication, presumably for
3C7 (Schultz in Morgan, Schultz, and Curry, 1941, origin: Ultraviolet induced.
Carnegie Inst. Wash. Year Book 40: 283). discoverer: Meyer, 50d.
other information: Probably a member of the Notch references: Meyer and Edmondson, 1951, DIS 25: 71.
pseudoallelic complex. phenotype: Somewhat lighter than b. RK2.

origin: Ultraviolet induced.


discoverer: Meyer, 51f.
references: Meyer and Edmondson, 1951, DIS 25: 71.
phenotype: Like b50d. RK2.
*b&: black-Dominant
origin: Spontaneous.
discoverer: Goldschmidt, 1945.
synonym: 6^.
references: 1945, Univ. Calif. (Berkeley) Publ.
Zool. 49: 504, 520.
phenotype: bP/b darker than tP/+ or e/e. Homozy-
gous lethal. RK2A.
cytology: Associated with Df(2L)bD =
Df(2L)35C;35D.
Figured in Goldschmidt (1945, p. 520).
B: Bar
location: 1-57.0.
origin: Spontaneous in a female.
discoverer: Tice, 13b.
references: 1914, Biol. Bull. 26: 221-30 (fig.).
Ax; Abruptex Morgan and Bridges, 1916, Carnegie Inst. Wash.
From Mohr, 1932, Proc. Intern. Congr. Genet., 6th. Vol. Publ. No. 237: 66 (fig.).
1: 190-212. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Genet. 2: 29-33.
phenotype: Eye restricted to narrow vertical bar of
origin: X ray induced. about 90 facets in the male and 70 facets in the
discoverer: Green, 42gl. female as contrasted with normal numbers of about
references: Oliver, 1944, DIS 18: 44. 740 for males and 780 for females [Sturtevant,
phenotype: Similar to Ax, except male lethal. RK2. 1925, Genetics 10: 117-47 (fig)]. Homozygous fe-
b: black male fully viable. B/+ female has about 360 facets
location: 2-48.5. and shows indentation terminating in horizontal
origin: Spontaneous. fissure on anterior margin of eye, producing a
discoverer: Morgan, lOj. kidney-shaped eye. B/B and B/+ completely sepa-
references: Bridges and Morgan, 1919, Carnegie rable from wild type, but in some genetic back-
Inst. Wash. Publ. No. 278: 144 (fig.). grounds B/B overlaps J5/+ slightly. Classifiable
phenotype: Black pigment on body and tarsi and in single dose in triploids by slight anterior nick
along wing veins, darkening with age. Heterozy- in eye (Schultz, 1934, DIS 1: 55); is useful in the
gote shows somewhat darker trident, but is never recognition of triploids. Eyes of female heterozy-
confused with homozygote. Puparium usually gous for a deficiency for B and a normal X are
somewhat lighter than wild type and newly emerged normal (Sutton, 1943, Genetics 28: 97-107). Log
flies not clearly distinguishable from wild type of facet number inversely proportional to tempera-
(Waddington, 1941, Proc. Zool. Soc. London, ture of development (Hersh, 1930, J. Exptl. Zool.
Ser. A 111: 173-80). Tyrosinase formed in adult 57: 283-306).
(Horowitz). RK1 in aged flies.
Nonautonomous over short distances (Sturtevant,
cytology: Salivary chromosomes apparently normal.
1932, Proc. Intern. Congr. Genet., 6th, Vol. 1:
Placed in region between 34E5 and 35D1, on basis
304—7). Facet development enhanced in organ
of its inclusion in Df(2L)64j =•- Df(2L)34E5-
culture by addition of wild type cephalic com-
F1.35C3-D1 (E. H. Grell).
plexes IKuroda and Yamaguchi, 1956, Japan J.
MUTATIONS 23

Genet. 31: 97-102 (fig.)]- Facet number can be in- discoverer: Bridges, 31al5.
creased by addition of a number of compounds to references: Dobzhansky, 1932, Genetics 17:
the medium; probably not a specific inhibition of 369-92.
effect of B (see work of Chevais, Khouvine, Kaji, phenotype: Very slight Bar; merely nick in anterior
Abd-El-Wahab, and DeMarinis). margin of eye in males (no overlap) and in homo-
Embryological studies [Chen, 1929, J. Morphol. 47: zygous females. B4/+ shows slight nick in 10 per-
135-99 (fig.); Steinberg, 1941, Genetics 26: 3 2 5 - cent of cases only. RK2(A).
46 (fig.); 1942, Genetics 27: 171-72; Power, 1942, B36b
Genetics 27: 161, DeMarinis, 1952, Genetics 37: origin: Spontaneous as BB36b in BB chromosome of
75—89 (fig.)] indicate that phenotype results from BB/ln(l)AM female.
reduced number of cells in optic disk and reduced discoverer: Bridges, 36b2.
rate of cell division in anterior part of eye. Facet phenotype: Male resembles standard B; BB36b/+
development responds strongly to environmental female has smaller eye than B/+ but larger and of
factors around 60 hr after oviposition (Luce, different shape than BB/+. Poor fertility both
Quastler, and Chase, 1951, Genetics 36: 488—99). sexes. RK2A.
Pigmented but nonfaceted part of eye shows other information: Homozygous females produce
retinulae and dioptic apparatus lacking, but rudi- wild type and extreme Bar unequal recombinants.
mentary ommatidia present, consisting of hyper-
trophied accessory cells (Wolsky and Huxley, origin: Spontaneous derivative of B in ClB.
1936, Proc. Zool. Soc. London 485-89). RK1A. discoverer: Dempster, 36d9.
cytology: Located in 16A1-2. Associated with references: 1937, DIS 8: 8.
Dp(l;l)B = Dp(l;l)15F9-16Al;16A7-Bl. phenotype: Narrow Bar resembling BB. B36d/+
other information: Since B is a tandem duplication, easy to separate with unaided eye. RK1A.
B homozygotes may give rise to a nonduplicated
chromosome (reversal to normal phenotype) and a origin: Spontaneous in B + .
triplicated chromosome (i.e., double Bar = BB) as discoverer: L. V. Morgan, 36j20.
reciprocal products of unequal crossing over references: 1937, DIS 7: 5.
(Sturtevant and Morgan, 1923, Science 57: 746— phenotype: Slight B; usually stronger than B4, but
47). From successive unequal crossovers in at- shows greater fluctuation and may overlap wild
tached X's, Rapoport (1940, Zh. Obshch. Biol. 1: type. RK3.
235-70; 194.1, DIS 15: 36-37) has been able to *B489
accumulate as many as 7 or 9 Bar regions in a origin: X ray induced in In(l)sc4.
single chromosome. Bar is the first recorded in- discoverer: Yu, 48g.
stance of position effect. Presumably results from references: 1949, DIS 23: 65.
the new band association 16A7-16A1 and can be phenotype: Eyes wider and more variable in width
reversed by rearrangements that separate these than B. Male sterile. RK2A.
bands. Also the first case of cis-trans position cytology: Associated with T(1;2)B486 = T(1;2)15F-
effect, two 16A7-16A1 associations in the same 16A1;33B superimposed on In(l)sc4 =In(l)lB3-
chromosome producing greater facet reduction than 4;19F-2OC1.
two associations in homologous chromosomes;
e.g., facet number in B/B is greater than in BB/+ origin: X ray induced.
(Sturtevant, 1925). discoverer: E. B. Lewis, 5814.
references: Ogaki, 1960, DIS 34: 97.
1960, Japan. J. Genet. 35: 282.
phenotype: At 25°C, male eyes have about five
facets fused into a vertical strip; Bsst/+ female
eyes have about 35. Higher temperature decreases
facet number. Addition of 2.5 percent lactamide to
medium increases facet number to almost 540 in
heterozygous female. Male sterile. RK1A,
cytology: Associated with T(1;3)B5S1 =
B: Bar T(1;3)16A;88F.
Left: heterozygous female. Right: hemizygous male.
From Sturtevant and Beadle, 1939. An Introduction to origin: X-ray-induced partial reversion of B'fl' in
Genetics. Sounders, p. 24. male.
B3 discoverer: Demerec, 34b.
origin: Spontaneous partial reversion of B. references: Sutton, 1943, Genetics 28: 97—107.
discoverer: Stern, 1926. phenofype: Resembles Bl. Viable. RK1A.
phenotype: Eye reduced less than in B and eye cytology: Associated with Dp(l;l)B263-28 -
surface rougher. RK1(A). Dp(l;l)15F9-16Al;16A3-4;16A6-7;16A7-Bl, which
*B4 was derived by deletion of 16A4 of leftmost region
origin: Spontaneous partial reversion of B in a male. through 16A6 of middle region of B^1 triplication.
24 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

other information: May be considered to be B1 in size and roughening of remainder. Male sterile;
derived by deletion of one of the regions in the heterozygous female fertile. Interpreted as posi-
BlB1 tandem triplication. tion effect (Dobzhansky, 1936, Biol. Rev.
*B263-34 Cambridge Phil. Soc. 11: 364-84). RK3A.
origin: X-ray-induced reversion of B*B* in male. cytology: Associated with T(l;2)Bbd =
discoverer: Demerec, 34c. T(l;2)16Al-2;48C2-3 + In(2R)41A;47A.
references: Demerec, 1934, Cold Spring Harbor
Symp. Quant, Biol. 2: 110—17.
Sutton, 1943, Genetics 28: 97-107.
phenotype: Eyes wild type. Lethal and cell lethal.
RK2A as lethal.
cytology: B263'34/+ resembles B*B'/+ (Sutton,
1943).
*B263-38
origin: X-ray-induced reversion of BtB* in male,
discoverer: Demerec, 34f.
references: Sutton, 1943, Genetics 28: 97-107. Bbd: Baroid
phenotype: Eyes wild type. Lethal. RK2A as Edith M. Wallace, unpublished.
lethal,
cytology: B263'38/+ resembles BiBl/+ (Sutton, *BDG: Bar of Dubinin and Goldat
1943). origin: X ray induced in B* chromosome in male.
*B263-47 discoverer: Dubinin and Goldat, 1936.
origin: X ray induced in B + male. references: 1936, Biol. Zh. (Moscow) 5: 881—84.
discoverer: Demerec, 38d. phenotype: Eye not described. Lethal when hemi-
references: Sutton, 1943, Genetics 28: 97-107. zygous and homozygous (seems likely that latter
phenotype: Eyes Bar-Mke but larger than Bar. Not claim inferred from former). RK2A as lethal.
lethal. RK1A. cytology: Associated with T(1;2)BD<* = T(1;2)4;15F-
cytology: Associated with ln(l)B263'47 = 16A;20;40-41.
ln(l)16A2-4;20A2-3.
B1: Bar-infrabar
+
origin: X ray induced in B male. origin: Spontaneous partial reversion of B that
discoverer: Bishop, 39i26. occurred in a male.
references: 1939, DIS 12: 61. discoverer: Sturtevant, 1923.
1940, DIS 13: 48. references: 1925, Genetics 10: 117-47 (fig.).
phenotype: Eye reduction in male and heterozygous phenotype: Eye reduction about halfway between B
female between B and B1; size constant in males, and +: B1 male has 478 facets, B^B1 has 320, and
variable in females. Homozygous females viable Bty+ has 716. Facet development inversely pro-
and fertile; show a distinct bb effect. Wings portional to temperature; effective period for
usually leathery and warped at 19°C; normal at temperature treatment is 60 percent through larval
25°. RK2A. life or about 60 hr at 25°C (Luce, 1935, J. Exptl.
cytology: Associated with Tp(l)B263'48 = Zool. 71: 125-47). RK1A.
Tp(l)3E2-3;15F9-16Al;20A2-3. cytology: Apparently no change in Dp(l;l)B in
*Q263-49 which B1 arose.
origin: X ray induced in BB male. *Bl40b
discoverer: Sutton, 41b. origin: Spontaneous in BB male.
references: 1943, Genetics 28: 97—107. discoverer: Steinberg, 40b.
phenotype: Eyes vary in male from BB to wild type, references: 1940, DIS 13: 51.
in homozygous female from BB to B/+. RK2A. phenotype: Similar to B1 both in male and B f /+
cytology: No change in the BB triplication de- female. RK1A.
tectable in salivaries. (Sutton, 1943). cytology: No change from original BB triplication
(Sutton, 1943, Genetics 28: 97-107).
origin: X-ray-induced reversion of BB in male.
discoverer: Sutton, 1940. BMl: Bar of Mullet
references: 1943, Genetics 28: 97-107. origin: X ray induced in B* chromosome.
phenotype: Eyes wild type. Viable. discoverer: Muller, 34e.
cytology: B263'51/+ resembles BB/+ (Sutton, 1943). references: 1935, DIS 3: 29.
; Bar-baroid phenotype: Weak allele of B; always has at least a
origin: X ray induced in B + male. derangement of facets on anterior margin of eye.
discoverer: Dobzhansky, 31bS. RK2A.
references: 1932, Genetics 17: 369-92. cytology: Associated with ln(l)BM1 =
phenotype: Recessive. Eye of male has slight in- In(l)16A2-5;20A3-B (Sutton, 1943, Genetics 28:
dentation of anterior margin, with some reduction 97-107).
MUTATIONS 25

Bridges and Morgan, 1919, Carnegie Inst. Wash.


origin: X ray induced in B+ chromosome. Occurred Publ. No. 278: 148 (fig.).
simultaneously with a reverse mutation of v. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
discoverer: Muller, 34e. Genet. 2: 212 (fig.). 218.
references: 1935, DIS 3: 29. Bridges, 1937, Cytologia (Tokyo), Fujii Jub. Vol.
phenotype: Weak allele of B. RK2A. 2: 745-55.
cytology: Associated with In(l)B^2 = In(l)16A2- phenotype: Wings at first inflated with hemolymph
5;20E button, 1943, Genetics 28: 97-107). to produce blisters and vesicles; venation weak,
*BPar: Bar-partial plexus-like; wings smaller, warped, discolored,
origin: X-ray-induced partial reversal of B in male. divergent. Effect caused by inadequate contrac-
discoverer: Bishop, 1940. tion of epithelium after inflated stage of pupal
references: Sutton, 1943, Genetics 28: 97—107. wing [Waddington, 1940, J. Genet. 41: 75—139
phenotype: Eyes intermediate between B and +. (fig.)]. Sensitive to temperature. RK3 above 25°C;
RK2A. RK2 at 19° or below.
cytology: B duplication unchanged (Sutton, 1943). cytology: Located between 60C5 and 60D2 based on
other information: Six independent partial rever- inclusion within Df(2R)Px = Df(2R)60B8-10;6QDl-2
sions fitting this description found by Bishop. and within Df(2R)Px2 = Df(2R)60C5-6;60D9-10
(Bridges, 1937).
* S R : Bar of Rapoport other information: May be part of a pseudoallelic
origin: X ray induced in normal chromosome. complex with 6s and Px.
discoverer: Rapoport, 1935.
synonym: Bz: Bar of Zuitin.
references: Zuitin, 1935, DIS 4: 6, 16.
1936, DIS 5: 6.
phenotype: More extreme than B. BR/+ resembles
BB/+. RK1(A).
other information: Shows normal crossing over and
reverts to wild type.
*Brev-1: Bar-reversed
origin: X ray induced in B male.
discoverer: Bishop, 1940.
references: Sutton, 1943, Genetics 28: 97—107.
phenotype: Eyes and viability normal.
cytology: B***-1/+ resembles B/+ (Sutton, 1943).

B*: Bar of Sfone


origin: X-ray-induced derivative of B.
discoverer: Stone, 1931. ba: balloon
phenotype: Extreme Bar; produces narrower eye From Bridges and Morgan, 1919, Carnegie Inst. Wash.
than B, both in males and heterozygous females. Publ. No. 278: 148.
RK1A.
cytology: Associated with T(1;4)BS = T(1;4)15F9-
16A1;16A7-B1;1O2F (Griffen, 1940, Genetics 26: origin: Spontaneous.
154-55; Lewis, 1956, DIS 30: 130). discoverer: Banerjee, 58i30.
BS31: Bar-Super inserted in chromosome 3 references: Ray-Chaudkuri, 1959, DIS 33: 99.
origin: Neutron induced in X'YS, sc w B chromo- phenotype: Wings blistered at one or two places,
some. affecting quite a broad area either on the inner
discoverer: Norby. margin or centrally; both wings usually affected.
synonym: BS2; Super-Bar. Wings generally glossy and contracted. Classifica-
references: Muller and Norby, 1949, DIS 23: 61. tion good; viability slightly reduced. RK2.
phenotype: Extreme Bar resembling Bs. RK1A.
cytology: Associated with T(l;3)Bs3i = : see bio
T(1;3)1SF9-16A1;16A7-B1;19-2O;Y;66B13-C1 bio
(Muller; Lindsley). Bag: see Bg
B*: see B*
b-l3 3gis; S ee tri *bal: bandy legged
ba: balloon location: 2- (not located),
location: 2-107.4. origin: Spontaneous.
origin: Spontaneous. discoverer: Stroher, 1958.
discoverer: Morgan, 10k. references: Mainx, 1958, DIS 32: 82.
references: Marshall and Muller, 1917, J. Exptl. phenotype: Legs extremely shortened and crippled.
Zool. 22: 457-70 (fig.). All parts of legs from femur to tarsi shortened,
26 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

broadened, and irregularly curved. Tarsal number that suggest that there is enough DNA in 66 + to
not reduced. Deformities most extreme in meta- specify approximately 130 molecules each of 28S
thoracic legs. Movement unsteady and tottering. and 18S ribosomal RNA, these authors view the 66
Manifestation increased by selection. Viability locus as highly redundant and perhaps composed of
poor, especially in males; fertility good. RK2. a very large series of tandem duplications. They
bald: see ra2 interpret bb mutations as partial deletions of the
ballet: see bit locus. They postulate that in 66 flies the rate of
balloon: see ba protein synthesis is limited by the amount of ribo-
Balloon: see Bb somal RNA and the 66 phenotype results in part
balloon wing: see bs3 because normal bristle production represents max-
band: see bn imum protein synthesis on the part of the trichogen
bandy legged: see bal cells during a particular interval in development.
Bar: see B RK2.
Bar + Bar: see BB cytology: Judged to be in 20C2 (or 20C1) by Cooper
Bar double: see BB (1959, Chromosoma 10: 535—88) based on exten-
bar eye: see at sive consideration of published cytology of base
of X chromosome. The 66 locus lies in proximal
bar-3: bar on chromosome 3 heterochromatin of X, probably proximal to and
location: 3-79.1 (not an allele of ro). very close to nucieolus organizer in heterochro-
origin: Spontaneous. matic region hB (Cooper, 1959). Ritossa, Atwood,
discoverer: Ives, 49J20. and Spiegelman (1966), on the other hand, postulate
references: 1950, DIS 24: 58. that the nucieolus organizer is the cytological
phenotype: Like B/B, without significant variation counterpart of the 66 locus. Presence of a
under standard conditions. Viability good. RK1. normal allele of 66 on Y chromosome postulated
Bat-infra double: see 2?* 5* by Burlingame and demonstrated by Stern [l927, Z.
Barlike eye: see Ble Induktive Abstammungs- Vererbungslehre 44: 187—
baroid: see Bbd 231 (fig.)]. This 66 + allele almost certainly in Ys
bat: bat (see Cooper, 1959).
location: 2-71.0. other information: 66 stocks show marked tendency
discoverer: Bridges, 22J26. to accumulate modifiers that suppress the pheno-
synonym: ext-b: extended-b, type. Outcrossing generally brings about return of
phenotype: Wings extended and bent backward. 66 phenotype. Ritossa, Atwood, and Spiegelman
RK2. (1966), however, doubt that this is the case and
*baton: baton postulate that the level of tandem redundancy of
location: 2-52. the locus is subject to frequent stepwise increases
phenotype: Abdomen elongated, with defective or decreases by unequal crossover types of events.
plates; eye resembles L4. Extremely inviable; Many laboratory stocks can be shown by crossing
most homozygotes die in larval and pupal stages, to 6b* to carry 66 alleles of unknown origin.
appearing as elongated corpses. Heterozygote
shows some eye effect. RK3.

bb: bobbed
location: 1-66.0 (Bridges).
discoverer: Sturtevant, 20b.
synonym: 66*. What is now referred to as 6b was
derived from fifth finding of bb. First allele found
was lost and is here omitted from consideration.
phenotype: Bristles of homozygous females de-
creased in both length and thickness. Tergites
etched at sides. Considerable variability of
bristle character, with some overlapping; ab-
dominal character extremely erratic. X/0 male has
phenotype similar to but more extreme than homo-
zygous female. X/Y male is wild type, owing to
presence of normal allele of 66 in Ys; X/X/Y fe-
male similarly normal in phenotype. 66/66' is
extreme 66 in phenotype. Viability variable.
Ritossa, Atwood, and Spiegelman (1966, Genetics
54: 819-34) showed that 66 contains about half as
much ribosomal RNA-complementary DNA as 66 + .
They conclude that the 66 locus is the site of ribo- an extreme bobbed
somal RNA synthesis. On the basis of calculations Edith M. Wallace, unpublished.
MUTATIONS 27

bb$: see bb and bbda is a property of the 66 locus and bb }4


bb<t*>. R K 2 .
origin: Spontaneous in attached X's. *bbG 7: bobbed of Goldschmidt
discoverer: Gabritschevsky, 1926. origin: Spontaneous.
phenotype: When first found, this was a very ex- discoverer: Goldschmidt.
treme 66 with small bristles and very scaly ab- synonym: 66 s P* *P.
domen; it gradually became a weak 6b. Enhances references: 1945, Univ. Calif. (Berkeley) Publ.
expression of gt. RK3. Zool. 49: 3 9 1 - 9 3 .
phenotype: Homozygote normal; bbG1/bbG2 shows
origin: Spontaneous, extreme bristle reduction and abdominal etching.
discoverer: Bridges, 30b24. For interaction with other 66 alleles, s e e
phenotype: bb20/bb is strong bb. bb20 is homozy- Goldschmidt, 1945, table 75. RK2 in some combi-
gous lethal. RK2. nations.
other information: Claimed by Goldschmidt to recur
origin: Spontaneous, in both X and Y chromosomes of certain lines.
discoverer: Stern, 28110.
synonym: bbx. origin: Spontaneous.
references: 1935, DIS 3: 29. discoverer: Goldschmidt.
phenotype: Like 6b. RK2. synonym: bb0 Px aP hi
.
bb': see bbds references: 1945, Univ. Calif. (Berkeley) Publ.
Zool. 49: 391-93.
origin: Thought by Stern and Ogura to be an extreme phenotype: Almost completely lethal homozygous.
66 allele normally occurring on YL in addition to Shows extreme bristle shortening and abdominal
the normal allele occurring on Y$. etching in combination with bbG3. RK2.
discoverer: Stern. other information: Claimed by Goldschmidt to recur
references: Stern and Ogura, 1931, Z. Induktive frequently in certain lines.
Abstammungs- Vererbungslehre 58: 81—121.
phenotype: Only observable evidence of existence origin: Spontaneous.
of 6 6 " i s that, when added to other 66 genotypes, discoverer: Goldschmidt.
it apparently causes them to become slightly less synonym: bt>Poi.
extreme. This could be simply a suppressing references: 1945, Univ. Calif. (Berkeley) Publ.
effect of YL rather than a dosage effect attrib- Zool. 49: 390-93.
utable to a mutant allele of 66. RK3. phenotype: Weak bobbed allele. Extreme in combi-
other information: Inviability of bb*/R(Y)L renders nation with bbG2. RK2 in some combinations*
existence of 6 unlikely. other information: Claimed by Goldschmidt to recur
bba Px SP: s e e bbG1 repeatedly in certain X- chromosomes.
bba Px SP h': see bbG2
*bbD: bobbed-Dominant origin: Spontaneous.
origin: X ray induced. discoverer: Goldschmidt.
discoverer: Lefevre, 48g28. synonym: bbP°i *?.
references: 1949, DIS 23: 58. references: 1947, J. Exptl. Zool. 104: 197-221.
phenotype: Pronounced etching of abdominal phenotype: Bristle effect irregular; no abdominal
tergites; bristles only slightly reduced. Male etching. RK3.
genitalia directed posteriorly rather than ventrally. cytology: Salivary chromosomes normal (Hannah-
Viable in both sexes; fertility of females fair, but Ala va).
of males extremely low. Homozygous females not
produced. Viability and classification good in origin: Spontaneous.
combination with 66 and bbY. RK2- discoverer: Goldschmidt.
other information: Allelism with 66 not definitely synonym: bbPoi hi
.
established. references: 1945, Univ. Calif. (Berkeley) Publ.
bbd*: bobbed-deficiency sensitive Zool. 49: 387, 390-93.
origin: The allele present in some stocks marked phenotype: Homozygous lethal. bbG5/bb produces
66. shortening of bristles and abdominal etching.
synonym: 66 . RK2 as lethal.
phenotype: Females homozygous for bb^s or hetero- bb-': bobbed-lethal
zygous for bbda and a mutant allele of 66 are 66 origin: Spontaneous.
in phenotype. bbd*/Df(l)bb females are lethal. discoverer: Bridges, 1926.
66cfs-like alleles have been reported by Stern and references: Morgan, Bridges, and Sturtevant, 1926,
Ogura (1931, Z. Induktive Abstammungs- Carnegie Inst. Wash. Year Book 25: 308-12.
Vererbungslehre 58: 81—121) and by Lindsley, Stern, 1929, Biol. Zentr. 49: 261-90.
Edington, and Von Halle (I960, Genetics 45: phenotype: Homozygous lethal. 66 J /66 is extreme
1649—70). Presumably, this difference between 66 bb, very late hatching, and invariably classifiable.
28 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

bbtybb'/Y = bbl/Y = normal. 66*/0 is lethal. males. In(l)481 = In(l)12E-F;14B induced simulta-
Shown by Ritossa, Atwood, and Spiegelman (1966, neously.
Genetics 54: 819-34) to contain approximately one- *bbOf: bobbed of Offermann
fourth as much ribosomal RNA-complementary DNA origin: X ray induced in T(1;4)A1.
as 66+. RK2A. discoverer: Offermann, 1935.
other information: Segregation from Y chromosome references: 1935, DIS 3: 27.
normal. Crossing over reduced in right end of X in phenotype: Like 6b. RK2A.
6b'/+ females.
origin: X ray induced in In(l)sc8.
L L
origin: Spontaneous in X-Y , bb (X-Y , bb-bb" discoverer: Offermann.
produced X-YL, bbl~2'bb"according to Stern and references: 1935, DIS 3: 28.
Ogura). phenotype: Like 6b. RK2A.
discoverer: Stern, 28k. bbPoi: see bb°3
references: Stern and Ogura, 1931, Z. Induktive bbP°i47: see bbG4
Abstammungs-Vererbungslehre 58: 81—121.
phenotype: Homozygous lethal. bb1'2/bb is bobbed. bbx: see
RK3A. bbY: bobbed on the Y chromosome
origin: Spontaneous.
origin: X ray induced. discoverer: Bridges, 1926.
discoverer: Lindsley, Edington, and Von Halle. synonym: Ybb.
references: 1960, Genetics 45: 1649—70, phenotype: bb/bbY male has slight 6b phenotype;
phenotype: Lethal as X/0 male and in combination usually separable in crosses, but stock 66/66*'
with In(l)sc4Lsc8R and with bbds. bb1-3a/bb is male tends to change to nearly normal phenotype.
bobbed. RK2A. bbl/btiY male is good 66; always separable and
cytology: Association with Df(l)bbt-3a inferred fertile. RK2 as 66*/66^.
from its irregular segregation from y+Y in males. other information: Chromosome with 66^ described
bbl-74 as Y66 in subsection on Y derivatives.
origin: X ray induced. *bbY-20
discoverer: Lindsley, Edington, and Von Halle, origin: Spontaneous. Arose in combination with
references: 1960, Genetics 45: 1649—70. 6620.
phenotype: Like 66'-3a, RK2A. discoverer: Bridges, 30b24.
cytology: Association with Dt(l)bb1~'''4 inferred synonym: ybb-20t
from irregular segregation from y+Y in males. phenotype: Slightly less extreme but otherwise like
. RK2.
origin: X ray induced.
discoverer: Lindsley, Edington, and Von Halle. origin: Spontaneous. Arose in combination with
references: 1960, Genetics 45: 1649—70.
phenotype: Like 66*-3a. RK2A. discoverer: Sturtevant, 31c26.
cytology: Association with Df(l)bbt~iss inferred synonym: Ybb"21<.
from irregular segregation from y+Y in males. phenotype: Like 66*\ RK2.
*bb Y -2 2
origin: X ray induced. origin: Spontaneous. Arose in combination with
discoverer: Lindsley, Edington, and Von Halle. 66".
references: 1960, Genetics 45: 1649—70. discoverer: Curry, 37118.
phenotype: Like 66'-3«. RK2A. synonym: Ybb~22.
cytology: Association with Di(l)bbim4S2 inferred phenotype: Like 66^. RK2.
from slightly irregular segregation from y + F in
male. Bb: Bubble
location: 1- (not located) or 3-48.
origin: X ray induced. origin: X ray induced.
discoverer: Lindsley, Edington, and Von Halle. discoverer: R. L. King, 32d.
references: 1960, Genetics 45: 1649—70. synonym: Balloon.
phenotype: Like bb*-3*. RK2A. phenotype: Wings of heterozygous female smaller,
cytology: Association with Dl(l)bbl-4S6 inferred trimmed, and inflated. Bubble in first posterior
from irregular segregation from y*Y in male. cell. In extreme cases and usually in males, the
wing is a small inflated sac. Sexual difference
origin: X ray induced. in expression may indicate that J56 is on the X.
discoverer: Lindsley, Edington, and Von Halle. Female fertile; male entirely sterile; therefore
references: 1960, Genetics 45: 1649-70. homozygous females not obtainable. RK3A.
phenotype: Like bb'S®. RK2A. cytology: Associated with T(l:3)Bb =
cytology: Association with Dffl)hb!"*8* inferred T(1;3)13E;84F (Morgan, Bridges, and Schultz,
from grossly abnormal segregation from y*Y in 1937, Carnegie Inst. Wash. Year Book 36: 301).
MUTATIONS 29

references: 1925, Genetics 10: 117—47 (fig.).


phenotype: Less reduction in eye than BB. Facet
numbers are 38, 46, and 200 in B'B^B^' female,
BlBl male, and BiBi/+ female, respectively.
Median ocellus lacking or strongly reduced
(Lefevre, 1941, DIS 14: 40). RK1A.
cytology: Associated with Dp(l;l)BB - Dp(l;l)15F9-
16A1;16A7-B1; a tandem triplication.
other information: B* can be recovered as recom-
binant from BiBi/+.

origin: Spontaneous, presumably through unequal


crossing over in B'4(>b/B^0b%
discoverer: Steinberg, 40b.
references: 1942, DIS 16: 53.
phenotype: More extreme reduction in eye size than
Bb: Bubble Bi40b. RK2A.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. cytology: Associated with Dp(l;l)BB = Dp(l; 1)15F9-
Publ. No. 552: 23. 16A1;16A7-B1; a tandem triplication.
BB: Bar + Bar bd: see ra2
origin: Spontaneous though unequal crossing over
in B/B (see description of B).
discoverer: Zeleny.
synonym: Bar double; Ultra-bar; double Bar.
references: 1920, J. Exptl. Zool. 30: 292-324
(fig-).
Sturtevant, 1925, Genetics 10: 117-47 (fig.)-
phenotype: Eye more reduced than in B. Facet
numbers are 25, 29, and 45 in BB/BB female, BB
male, and BB/+ female, respectively. Median
ocellus lacking or strongly reduced (Lefevre,
1941, DIS 14: 40). Optic glomerulus reduced
(Power, 1942, Genetics 27: 161). RK1A.
cytology: Associated with a tandem triplication of
the region duplicated in Dp(l;l)B = Dp(l;l)15F9-
16A1;16A7-B1 [Bridges, 1936, Science 83: 210-11
(fig.)]-
BB1: Bar + Bar-infrabar
origin: Spontaneous through unequal crossing over
in B/Bl.
discoverer: Sturtevant.
references: 1925, Genetics 10: 117-47.
phenotype: Like BB. RK1A.
cytology: Associated with Dp(l;l)BB = Dp(l;l)15F9-
16A1;16A7-B1; a tandem triplication, 6 of: Beaded
other information: B is to the left of Bl and both From Bridges and Morgan, 1923, Carnegie Inst. Wash.
types can be recovered as recombinants. Publ. No. 327: 152.
B'B: Bar-infrabar + Bar 6of; Beaded
origin: Spontaneous through unequal crossing over location: 3-93.8.
in B/Bl. discoverer: Morgan, lOe.
discoverer: Sturtevant. references: Dexter, 1914, Am. Naturalist 48:
references: 1925, Genetics 10: 117-47. 712-58 (fig.).
phenotype: Like BB1. RK1A. Bridges and Morgan, 1923, Carnegie Inst. Wash.
cytology: Associated with Dp(l;l)BB = Dp(l;l)15F9- Publ. No. 327: 37, 152 (fig.)-
16A1;16A7-B1; a tandem triplication. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
other information: Bl is to the left of B and both Genet. 2: 45.
types can be recovered as recombinants. phenotype: Wings reduced by marginal excision
B'Bi both anteriorly and posteriorly. Extremely variable
origin: Spontaneous through unequal crossing over and overlaps wild type. Expression and interac-
in B'/Bl. tion studied by Goldschmidt and Gardner (1942,
discoverer: Sturtevant, 1923. Univ. Calif. (Berkeley) Publ. Zool. 49: 103-24).
synonym: Bar-infra double. Almost entirely suppressed by fif. In combination
30 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

with many different Minutes, causes incomplete Bridges and Morgan, 1923, Carnegie Inst. Wash.
development of anal and genital imaginal discs in Publ. No. 327: 179 (fig.).
males and less frequently in females (Goldschmidt, Stark and Bridges, 1926, Genetics 11: 249-66.
1948, Proc. Natl. Acad. Sci. U.S. 34: 245-52,' Stark, 1935, DIS 4: 62.
Sturtevant, 1949, Proc. Natl. Acad. Sci. U.S. 35: phenotype: Melanotic tumors appear in larvae and
311-13). Homozygous lethal. RK2 as lethal, RK3 persist in adults. Subject to modification by
as dominant. genetic factors. Nonlethal. RK3.
other information: Bd/In(3R)C, l(3)a was the first Beaded: see Bd
described case of a balanced lethal [Muller, 1918, Beadex: see Bx
Genetics 3: 422-99 (fig.)]. Beadexoid: see Bxd
bending wings: see osbdw
origin: X ray induced. benign tumor in chromosome 3: see be-3
discoverer: Ohnishi, 49116. bent: see bt
references: 1950, DIS 24: 61. bent scutellars: see bsc
phenotype: Like extreme Bd. Variable, overlaps *ber: berrytail
wild type. Homozygous lethal. RK2 as lethal; location: 1-52.4.
RK3 as dominant. origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
; Beaded of Goldschmidt phenylalanine (CB. 3007).
origin: Found among progeny of heat-treated flies. discoverer: Fahmy, 1953.
discoverer: Goldschmidt, 1934. references: 1958, DIS 32: 67.
references: Gottschewski, 1935, DIS 4: 14, 16. phenotype: Abdomen narrow, ending in a berry-like
phenotype: Like Bd but more extreme; not over- protrusion carrying defective genitalia. Wings
lapping wild type in stock. Balancers also re- opaque, with areas of deranged hairs (some with
duced with no overlap. Partially suppressed by H. cut inner margins and interrupted or abnormally
Schultz and Curry report recurrent small or welt- positioned longitudinal veins). Anterior scutellars
like eye effect that is not well understood but is often acutely bent; eyes occasionally misshapen.
in chromosome 3 and may be an effect of BdG. Males sterile and viability about 40 percent wild
Lethal homozygous and in combination with Bd. type. RK3.
RK1. bf: brief
location: 3-95.
origin: Spontaneous. origin: Spontaneous.
discoverer: Goldschmidt. discoverer: Curry, 3813.
references: 1945, Univ. Calif. (Berkeley) Publ. references: 1939, DIS 12: 45.
Zool. 49: 520. phenotype: Fly small; bristles Minute-like. Clas-
phenotype: Like Bd but with more extreme sification perfect, viability fair. Male completely
scalloping effect. RK3. sterile, female with low fertility. RK3.
*BdP: Beaded of Piternick fig; Bag
origin: X ray induced. location: 1-51.6 (to the right of sd).
discoverer: Piternick, 1949. origin: Spontaneous.
references: Goldschmidt, 1953, J. Exptl. Zool. discoverer: Bridges, 33d22.
123: 79-114. phenotype: Heterozygous female with wings shorter
phenotype: Like Bo* but more highly penetrant. and blunter, shortened L5, extra veins or gaps
30—40 percent BdP/+ are phenotypically Bd. near anterior crossvein, and inflated bag centering
Penetrance 100 per cent when heterozygous for in first basal cell. Frequently overlaps wild type.
third chromosome inversions, e.g., In(3L)P + Lethal in male. RK2 as a lethal, RK3 as a
In(3R)P. Interactions with other genes discussed dominant.
by Goldschmidt (1953). Homozygous lethal. RK2 cytology: Probably in 13C, based on Bg-like varie-
as lethal; RK3 as dominant. gation of T(l;3)rasv = T(1;3)9E;13C;81F.
*Bd": Beaded of Wallace *Bg2
origin: Spontaneous. origin: Spontaneous.
discoverer: E. M. Wallace, 15110, discoverer: S^mme.
phenotype: Like Bd but more extreme; ends of L3 synonym: Uw: Uneven wing.
and L4 split or disturbed. Overlaps wild type. phenotype: Heterozygous females with inner wing
Least extreme recognizable phenotype is nick margin frequently nicked or uneven; longitudinal
opposite L3. Homozygous lethal. RK2 as lethal. veins sometimes shortened; one wing often shorter
bdw: see osbdw than the other. Some delay in eclosion. Overlaps
*be~3: benign tumor in chromosome 3 wild type. Viability and fertility fair. Reduces
iocotion: 3-25. size of B eye and is itself exaggerated in combi-
origin: Spontaneous. nation with B. Male lethal. RK2 as lethal; RK3
discoverer: Stark, 16k. as dominant.
references: 1919, Proc. Natl. Acad. Sci. U.S., 5:
573-80 (fig.). origin: Induced by ingested radiophosphorus.
MUTATIONS 31

discoverer: R. C. King, 49h. cytology: Associated with In(3L)Bit; breakpoints


references: Poulson and King, 1949, DIS 23: 62. not determined.
phenotype: Heterozygous female has wings with no bithorax: see bx
crossveins, L5 shortened. Wings asymmetric in Bithoraxlike: see Ubx
size and blistered. L3 very thick; gaps in L3 and bithoraxoid: see bxd
L4. Extra veinlets; veins may fork at wing edge. Bitten: see Bit
Wings often excised terminally and along inner *bk: buckled
margin. Phenotype very variable, overlaps wild location: 1-59.8.
type. Viability 65 percent normal. Male lethal. origin: Induced by p-NN-di-(2-chloroethyl)amino-
RK3. phenylethylamine (CB. 3034).
*BgS2c discoverer: Fahmy, 1955.
origin: Thermal neutron induced. references: 1959, DIS 33: 83.
discoverer: R. C. King, 52c. phenotype: Wings slightly altered in shape and
references: 1952, DIS 26: 65. frequently divergent, with membranes warped be-
phenotype: Like Bg49h', overlaps wild type. Male tween longitudinal veins. Veins slightly thickened
lethal. RK3. at wing margins. Eye shape slightly altered.
hi: bifid Scutellar bristles frequently abnormal, either in-
location: 1-6.9. serted in base atypically, bent, or duplicated.
discoverer: Morgan, Ilk. Males viable and fertile. RK3.
references: Morgan and Bridges, 1916, Carnegie *bk2
Inst. Wash. Publ. No. 237: 28 (fig.). origin: Induced by L-l:6-dimethanesulfonyl mannitol
phenotype: Longitudinal veins fused at base of (CB. 2628).
wing into bifid stalk. L3 delta-like at tip; L4 discoverer: Fahmy, 1960.
often incomplete at tip. Wing margins often ex- references: 1964, DIS 39: 58.
cised at tip of L4. Wings spread in proportion to phenotype: Fly small. Legs shortened; posterior
their shortness. High temperature enhances and pair frequently deformed or absent. Wings
low temperature produces overlapping of wild type. shortened, abnormally shaped, with varying
Stronger in male than in female. Enhances Bx amounts of marginal vein incised. Sex combs may
alleles as well as sd, cp, and vgnP (Waletzky). be enlarged. Bristles stiff; occasionally an extra
RK1. scutellar bristle. bk2/bkl normal. RK3.
cytology: Between 4C7-8 and 4D1-2 according to *Bkd: Blackoid
Demerec, Kaufmann, Fano, Sutton, and Sansome location: 2-65 (Braun).
(1942, Carnegie Inst. Wash. Year Book 41: 191). origin: Spontaneous.
discoverer: Goldschmidt, 1938.
origin: Spontaneous. phenotype: Body color black in homozygote, dis-
discoverer: Gottschewski, 1935. tinctly darker than wild type in heterozygote.
phenotype: Like bi, but w h e r e a s males show 100 RK2.
percent e x p r e s s i o n at 25°C, females show 0—3 *bkl: buckledlike
percent. RK1 in male, RK3 in female. location: 1-59.9.
bis: bistre origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
location: 1-20.1. phenylalanine (CB. 3026).
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- discoverer: Fahmy, 1955.
phenylalanine (CB. 3007). references: 1959, DIS 33: 83.
discoverer: Fahmy, 1954. phenotype: Wings slightly divergent with membranes
references: 1958, DIS 32: 67. warped between longitudinal veins, which them-
phenotype: Very dark brown eye color; ocelli also selves are often slightly thickened. Abnormally
dark. Wings frequently unexpanded. Males sterile. shaped eyes, frequently compressed dorsoven-
Viability varies from less than 10 percent to 70 trally. Both sexes viable and fertile. RK3.
percent wild type. RK2A. other information: Probably a complementing allele
cytology: In bands 7B6 and 7; associated with of bk. One X-ray-induced allele.
Dt(l)bis = Df(l)7B5-6;7B7-8. BI: Bristle
*Bit: Bitten location: 2-54.8 (crossing over may be reduced).
location: 3- (not located; crossing over between ru origin: Spontaneous.
and th almost completely suppressed). discoverer: R. L. King, 25dll.
origin: X ray induced. references: 1927, Biol. Bull. 53: 465-68.
discoverer: Lefevre, 48g5. phenotype: Bristles one-half to two-thirds normal
references: 1949, DIS 23: 58. length, blunt, thicker, and beaded in outline.
phenotype: Inner margin of wing indented. Wings, Posterior scutellars often cross and adhere to
normally folded, appear to have had a bite taken body. Eyes somewhat larger and rougher. Prob-
out of the back. Marginal hairs present unlike N ably affects nature of bristle secretion, partic-
and ct. Flight is impeded, although little wing ularly outer layer [Lees and Waddington, 1942,
area lost. Homozygous lethal. RK1A. DIS 16: 70; Lees and Picken, 1945, Proc. Roy.
32 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

Soc. (London), Ser. B 132: 396-423 (fig.)]. Via- origin: X ray induced.
bility of heterozygote is good but erratic; homozy- discoverer: Crowell, 57i.
gotes usually lethal; survivors female-sterile with references: Meyer, 1958, DIS 33: 97.
roughish eye character. RK1 as dominant. phenotype: Eye shape indistinguishable from Bar.
*fi 13 0 Expression of Ble/+ varies; best at 26°C. Ex-
origin: Recovered among progeny of heat-treated cellent expression in homozygote at all tempera-
flies. tures. Ble/Ble in combination with JB results in
discoverer: Plough, 1930. an extremely narrow eye. RK1.
synonym: Sy30: Stubby-30. other information: If Ble represents a transposition
references: Plough and Ives, 1935, Genetics 20: of the Bar locus to chromosome 3, the flanking
42-69. loci of f+ and od+ have not been transposed.
phenotype: Like Bl. RK1. Also against transposition is absence of sexual
dimorphism that dosage compensation of B should
origin: Recovered among progeny of heat-treated produce in such a case.
flies. *bleg: black leg
discoverer: Ives 31119. location: 3- (near p).
synonym: Sy31U9. discoverer: Bridges, 16b23.
references: Plough and Ives, 1935, Genetics 20: references: Bridges and Morgan, 1923, Carnegie
42-69. Inst, Wash. Publ. No. 327: 158.
phenotype: Like Bl. Bl^t/Bl like Bl/Bl and phenotype: Legs black, body color pallid. Wings
poorly viable. Bl311 regularly homozygous lethal. flimsy. RK3.
RK1. blistered; see bs
*h!a: bladderwing Blister I ike: see Bsl
location: 1-43.2. blister/: see by
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- bio: bloated
phenylalanine (CB. 3025). location: 2-58.5.
discoverer: Fahmy, 1953. origin: Recovered among descendants of heat-
references: 1958, DIS 32: 67-68. treated flies.
phenotype: Wings grossly deformed, small, and discoverer: Ives, 33f26.
normally full of fluid. Eyes slightly abnormal in synonym: Originally referred to as ba^: balloon and
shape. Males fertile, females sterile; viability
about 50 percent wild type. RK3. references: Plough and Ives, 1934, DIS 1: 33.
black: see b 1934, DIS 2: 10.
black leg: see bleg 1935, DIS 3: 6.
Blackoid: see Bkd Bridges, Skoog, and Li, 1936, Genetics 21:
bladderwing: see bla 788-95.
Bid: Blond phenotype: Wings spread, crumpled, and vesicu-
location: 1- or 2- (associated with rearrangement). lated; wing shows irregular plexus of extra veins.
origin: Spontaneous in chromosome containing In extreme cases wings unexpanded. Occasional
In(2R)Cy. hooked or wavy bristles. Developmental studies
discoverer: Burkart, 1930. by Waddington [l939, Proc. Natl. Acad. Sci. U.S.
references: 1931, Rev. Fac. Agron. Vet. Univ. 25: 299-307 and 1940, J. Genet. 41: 75-139 (fig.)]
Buenos Aires 7: 393-491. show intervein material spongy and veins swollen,
Burkart and Stem, 1933, Z. Induktive with inadequate contraction after inflated stage of
Abstammungs- Vererbungslehre 64: 310—25 (fig.). pupal wing. Droplets of hemolymph often become
phenotype: Bristles of heterozygote are gleaming clothed with cells liberated from epithelium and
yellow at tips and for varying lengths of more remain along basal processes. Does not overlap
basal regions. Hairs not much paler and bristles wild type, but has poor viability and hatches
of abdomen only slightly affected. Larval mouth later. RK2.
parts wild type. No overlap. Viability and fertility cytology: Not included within and does not recom-
of male and heterozygous female excellent. For- bine with (0/1098) D((2R)Np = Df(2R)44Fl-
merly viable as a homozygous female but in lines 2;45El-2 (Bridges, Skoog, and Li, 1936).
now available the homozygote is lethal, presum- Blond: see Bid
ably owing to a lethal mutation closely linked to blot: see apbit
breakpoint of translocation in chromosome two. *btt: ballet
RK1A. location: 1- (not located).
cytology: Associated with T(l;2)Bld~ T(1;2)1C3- origin: X ray induced.
4,60B12-1J. discoverer: Iyengar.
other information: Bid phenotype associated with references: 1962, DIS 36: 38.
the 2RDXP element of the translocation. phenetype: Wings one-third the normal length;
*Bte: Barlike eye stretched outward and slightly upward; wing tip
location: 3-94. broadened; venation markedly altered as in fused.
MUTATIONS 33

Male viability impaired; females almost com- origin: Spontaneous.


pletely lethal. RK2. discoverer: Fabian, 1941.
bit: see apblt references: 1948, Arch. Julius Klaus-Stift.
*blu: blunt Vererbungsforsch. Sozialanthropol. Rassenhyg.
location: 3- (near ru). 23: 512-17.
origin: Spontaneous. phenotype: Eye color dark brownish red; darkens
discoverer: Wallbrunn, 46j23. with age. Malpighian tubules and testis sheaths
references: 1947, DIS 21: 71. colorless. Male fertile; female sterile. RK2.
phenotype: Wings slightly shorter and broader than *tow: bow wings
normal, giving a squared appearance. Sometimes location: 1- (not located).
difficult to classify. RK3. discoverer: Bridges, 12hl5.
*bn: band references: Morgan and Bridges, 1916, Carnegie
location: 3>72. Inst. Wash. Publ. No. 237: 46 (fig.).
origin: Spontaneous. phenotype: Wings curved downward over abdomen
discoverer: Morgan, 12g. and also sideways like bowl of a spoon. Overlaps
references: Bridges and Morgan, 1923, Carnegie wild type. RK3.
Inst. Wash.. Publ. No. 327: 79 (fig.). bow-legged: see bwl
Morgan, Bridges, and Sturtevant, 1925, Bibliog. bowed: see bod
Genet. 2: 215 (fig.), 218. bp: see bulbP
phenotype: Trident pattern and scutellum darker,
with dark transverse band across anterior portion
of mesonotum. Thorax vacuolated; hairs on thorax
sparse and directed medially, in bowed lines.
RK2.
DO: bordeaux
location: 1-12.5.
discoverer: Nazarenko.
phenotype: Eye color dark wine; not completely
separable from wild type. Red pigment 67 percent br: broad
wild-type level; brown pigment normal (Nolte, Edith M. Wallace, unpublished.
1955, J. Genet. 53: 1—10). Transplantation indi-
cates bo may be nonautonomous (Ephrussi and br: broad
Beadle, 1937, Genetics 22: 65-75). Larval Mal- location: 1-0.6.
pighian tubules bright yellow (Beadle, 1937, discoverer: Bridges, 15i26.
Genetics 22: 587-611). RK3. references: Morgan, Bridges, and Sturtevant, 1925,
bobbed: see bb Bibliog. Genet. 2: 145, 220 (fig.).
bobbed on the Y chromosome: see bbY phenotype: Wings somewhat broader than and about
bod: bowed 80 percent of length of normal, with round full tip;
location: 3-48.3. crossveins closer together. Shape difference vis-
origin: Spontaneous. ible in middle prepupal stage immediately after
discoverer: Nichols-Skoog, 35b20. eversion; probably an influence on cell division
references: 1937, DIS 7: 6. [Waddington, 1939, Proc. Natl. Acad. Sci. U.S.
phenotype: Wings bowed downward over abdomen, 25: 299-307; 1940, J. Genet. 41: 75-139 (fig.)].
curvature along both axes; curvature occasionally RK1.
reversed. Wings somewhat smaller than wild type. cytology: Located between 1C5 and 2C10 (Demerec,
Whole fly smaller and humpy; eyes slightly bulged. Kaufmann, Fano, Sutton, and Sansome, 1942,
Overlaps wild type slightly. Viability 75 percent Carnegie Inst. Wash. Year Book 41: 191).
wild type. RK3. br3
*bord: bordered origin: Spontaneous.
location: 1-70. discoverer: Bridges, 31el.
origin: Spontaneous. references: 1935, DIS 3: 7.
discoverer: Bridges, 1916. phenotype: Wings like br but more often arc-like
references: Morgan, Bridges, and Sturtevant, 1925, and crumpled. RK2.
Bibliog. Genet. 2: 220.
phenotype: Wings smaller and slightly extended; origin: Induced by DNA.
venation ragged and veins bordered by darker discoverer: Fahmy, 1959.
bands. Viability poor; classification unreliable. phenetype: Extreme expression of or"*? phenotype.
RK3. brS9!/br and bcS9i/ brutl have mutant phenotype.
bordeaux: see bo RK1.
bordered: see bord *brD: broad-Dominant
*bos: bordosteril origin: Spontaneous,
location: 3-0.0. discoverer: Muller, 19h.
34 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: Muller and Altenburg, 1921, Anat. Rec. discoverer: Fahmy, 1956.
20: 213. references: 1959, DIS 33: 83.
Muller, 1935, DIS 3: 29. phenotype: Fly with broad abdomen and slightly
phenotype: brD/+ resembles br/br. or-D/orand shortened thorax and wings. Wings frequently
brD/Y are lethal. RKl. slightly divergent. Eyes small and dull red with
*Jbr/-o; broad-lethal-a reflection spots. Bristles slightly shortened and
origin: Spontaneous. lying flatter on thorax. Males and females viable
discoverer: Muller, 19h. and fertile. RK2.
synonym: ifljfer^. other information: One allele induced by CB. 3025.
references: Muller and Altenburg, 1921, Anat. Rec. brc: brachymacrochaetae
20: 213. location: 1-0.0 (no recombinants with sc among
Muller, 1935, DIS 3: 29. 6746 sons).
phenotype: brlmB/br female is phenotypically br. origin: Induced by triethylenetnelamine (CB. 1246).
br'-* male dies. RK2. discoverer: Fahmy, 1952.
references: 1958, DIS 32: 68.
origin: Spontaneous. phenotype: One or more thoracic bristles much
discoverer: Muller, 19h. reduced in size; scutellars and dorsocentrals most
synonym: l(l)brb. frequently affected. Occasional bristles dupli-
references: Muller and Altenburg, 1921, Anat. Rec. cated. Good viability and fertility in both sexes.
20: 213. RK2.
Muller, 1935, DIS 3: 29. other information: One allele each induced by
phenotype: Like 6r;"a. RK2. CB. 3025, by CB. 1246, and by X rays.
*br*h: broad-short *brd: broadened
origin: Spontaneous. location: 1-33.
discoverer: Bridges, 14g20. origin: X ray induced.
references: 1916, Genetics 1: 151. discoverer: Muller, 26127.
phenotype: Wings one-half to two-thirds normal references: 1935, DIS 3: 29.
length, often arc-like and crumpled. Crossveins phenotype: Wings expanded. Viability 20 percent
closer together than normal and slight plexus wild type. RK3.
effect. Legs gnarled. RK2. *bre: bright eye
*brui: broad-unequal wings location: 1-24.6.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- origin: Induced by L~p-NN-di-(2-chloroethyl)amino-
phenylalanine (CB. 3026). phenylalanine (CB. 3025).
discoverer: Fahmy, 1955. discoverer: Fahmy, 1953.
synonym: uq. references: 1958, DIS 32: 68.
references: 1958, DIS 32: 77. phenotype: Eye color brighter red. Wings shorter,
phenotype: Wings short and very broad, often often crumpled or waved. Abdomen disproportion-
unequal in length; more extreme at 25°C. Occa- ately large. Male viability and fertility good, but
sionally one wing blistered, or grossly deformed. females have reduced fertility. Not easily classi-
Thoracic hairs irregularly distributed and sparse. fied. RK3.
Ocasional abnormality in bristle position or size. other information: One allele induced by CB. 1540.
Eyes smaller. Viability and fertility good in both brevis: see bv
sexes, bruq/br is wild type. RKl. bri: bright
other information: One X-ray-induced allele. location: 2-54.3.
*Br: Bridged origin: Spontaneous.
location: 1- (right half; crossing over suppressed to discoverer: Nichols-Skoog, 34b23.
the right of v). references: Beadle and Ephrussi, 1937, Am.
origin: X ray induced. Naturalist 71: 91-95.
discoverer: Muller, 2713. phenotype: Eye color bright red like en2 or v2;
references: 1935, DIS 3: 29. difficult to separate from wild type. Malpighian
phenotype: Plexus-like wings, with extra cross- tubules pale yellow (Beadle, 1937, Genetics 22:
veins bridging logitudinals. L4 bent. Wings 587-611). RK3.
arched. Male lethal. RK3A. Bridged; see Br
cytology: Associated with In(l)Br. brief: see bf
Br: see Sp bright: see bri
brachymacrachaetae: see brc bright eye: see bre
bran: see aba Bristle: see Bl
Bran: see aBa Bristled: see Sp
Branchlet: see Bt broad: see br
*brb; broad abdomen broad abdomen: see brb
location: 1-52.9. broadened: see brd
origin: Induced by styrylquinoline (CB. 3086). broader wing: see brw
MUTATIONS 35
2
bronze: see sf effect owing to additive semidominance; was
bronzy: see malbz originally interpreted to indicate allelism. RK2.
brown: s e e few *bs4
brown-like1: see red origin: Spontaneous.
brunette: see Hnr2 discoverer: Goldschmidt.
*6rw: broader wing references: 1947, J. Exptl. Zool. 104: 197-221.
location: 1-39.8. phenotype: More extreme allele of bs. RK2.
origin: X ray induced. cytology: Salivary chromosomes normal (Hannah).
discoverer: Fahmy, 1956. ; s e e bs*>l
references: 1959, DIS 33: 83.
phenotype: Wings broad and rounded at the tips. origin: Spontaneous.
Males show reduced viability and are sterile. discoverer: Strangio.
RK3. phenotype: More extreme allele of 6s with variable
expression, but wing generally converted to one
*bs: blistered large blister. Fully penetrant at 20°, 25°, and
location: 2-107.3. 30°C; blistering effect most marked at 20°C. RK2.
origin: Spontaneous. cytology: Salivary chromosomes normal (Strangio).
discoverer: Bridges, I l k l 6 .
references: Bridges and Morgan, 1919, Carnegie origin: Spontaneous.
Inst. Wash. Publ. No. 278: 155 (fig.). discoverer: Mohler, 54j7.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. references: 1956, DIS 30: 78.
Genet. 2: 219 (fig.). phenotype: bsS4i /bsS4i = bss^i/bs2 more extreme
phenotype: Wings blistered, small, pointed; vena- and less variable at 25°C than bs2/bs2. Viability
tion thick and plexus-like, with branches from and good in uncrowded cultures. RK2.
parallel to L5 beyond second crossvein, where
there is a semidominant free vein effect. Eye origin: Spontaneous derivative of 6s.
color softened. Temperature sensitive. RK2 at discoverer: Goldschmidt, 38i.
19°C; RK3 at 25°C. synonym: bs3Si.
cytology: Located between 60C5 and 60D2, based references: 1945, Univ. Calif. (Berkeley) Publ.
on its inclusion within Df(2R)Px = Df(2R)60B8- Zool. 49: 408-9, 416-17.
10.60D1-2 and within Df(2R)Px2 = Df(2R)60C5- phenotype: Stronger allele than bs2 but of low
6;60D9-10 (Bridges, 1937, Cytologia (Tokyo), viability and fertility. Wing shows very extended
Fujii Jub. Vol. 2: 745-55). web near the crossvein, which reaches margin of
other information: May be part of a pseudoallelic wing; above web a blister is formed. RK2.
complex with ba and Px.
bscy; blistered-curly
origin: Spontaneous.
discoverer: King and Poulson, 461.
references: Poulson and King, 1948, DIS 22: 54.
phenotype: Heterozygotes show venation abnormal-
ities, with tiny free veins usually in the third
posterior cell and occasionally in the second in
about 50 percent of males and 90 percent of fe-
males. Less often extra veinlets project from
bs*: blistered-2 posterior crossvein. Homozygotes have bizzare
Edith M. Wallace, unpublished. networks of wing veins. Wings of freshly emerged
adults inflated with hetnolymph, producing large
blister in middle of one or both wings just pos-
discoverer: Bridges, 25k24. terior to the anterior crossvein. Blisters cover
phenotype: More extreme allele of 6s. one-fourth to one-half of total wing area. Lymph
bs2/Df(2R)Px easily separable from +/Df(2R)Px, later dries, leaving wing vesiculated and curled
especially at or below 19°C. Wing effect caused upward. Condition more pronounced in females.
by same mechanism as that of px [Waddington, Flight restricted. bscy/bs2 females are like weak
1940, J. Genet. 41: 75-139 (fig.)]. RK2. bs°y and males like bs2. Viability nearly normal.
S3 RK1.
origin: Spontaneous. bsP
discoverer: Swigert, 31 d. origin: Spontaneous derivative of bs,
synonym: balloon wing; px3id*and px*>«, discoverer: Goldschmidt.
references: Plough and Ives, 1934, DIS 1: 33. references: 1945, Univ. Calif. (Berkeley) Publ.
1935, Genetics 20: 42-69. Zooi. 49: 409-18.
phenotype: Extremely blistered wing. Classifica- phenotype: Stronger allele than 6®. Sensitive to
tion easy in most stocks, + bs3/px + shows wing genetic modification. Females show broad chi-
36 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

tinous mass of extra veins; males show extra references: 1914, J. Exptl. Zool. 17: 325-36.
veins branching from L2. RK2. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
*bsPP Genet. 2: 216 (fig.), 219.
origin: Spontaneous derivative of 6s. Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
discoverer: Goldschmidt. phenotype: Wings held out at base and bent sharply
references: 1945, Univ. Calif. (Berkeley) Publ. backward. Rear legs often lumpy at first tarsal
Zool. 49: 409-18, 433-39. joint. May have one to four "preleg" or "first
phenotype: Stronger allele than bs. RK2. ventral" bristles on ventral surface of thorax
cytology: Frequently associated with short defi- anterior to first pair of legs, in space otherwise
ciency to the right of 6a (or 6s ?) within devoid of bristles or hairs. Overlaps wild type at
Df(2R)Px2? 25°C, very much at 19°, and little if any at 29°C
other Information: Claimed by Goldschmidt fre- (Metz, 1923, Proc. Soc. Exptl. Biol. Med. 20:
quently to recur by mutation in certain lines. 305-10). RK2 at 28°C.
*bsc: bent scutellars
location: 1-1.1.
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
phenylalanine (CB. 3007).
discoverer: Fahmy, 1954.
references: 1958, DIS 32: 68.
phenotype: One or more scutellars bent on them-
selves in form of inverted V. Other bristles irreg-
ularly bent. Eyes slightly smaller. Wings slightly
abnormal in shape. Male viability about 50 percent
wild type; fertility much reduced. RK3.
other information: One allele each induced by
CB. 3025 and CB. 3026.

*Bsl: Blister I ike


location: 2-104 (but located 3.2 units from bw
which is at 104.5).
origin: X ray induced.
discoverer: Oliver, 29bl. bt: bent
references: 1939, DIS 11: 47. From Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Extra veins and blister centering in Genet. 2: 216.
region of posterior crossvein, which is usually
absent. Fluctuation in expression from thickening *btD: bent-Dominant
of veins to blister covering entire wing. Vein L5 origin: X ray induced.
may be short. Homozygous lethal. RK3. discoverer: Schultz, 33all.
cytology: Not done, but Bridges has suggested that references: Bridges, 1935, Biol. Zh. (Moscow) 4:
this is probably a Plexate deficiency. 401-20.
bsp: brown spots phenotype: When found, btD/+ showed regularly
location: 2-40.6. divergent wings with some angular bend near base.
origin: Spontaneous. Legs lumpy at low temperature. Preleg bristles
discoverer: Di Pasquale. present as in bt. Homozygous lethal. RK3 as
references: 1959, DIS 33: 128. lethal.
Di Pasquale and Zambruni, 1963, DIS 37: 73 (fig.). other information: Balanced stocks in existence to-
1966, DIS 41: 119. day show only preleg bristle character and reces-
phenotype: Spots of brown pigment appear in integ- sive lethality (Lewis).
ument of bsp/bsp females only after they have
mated. Di Pasquali and Zambruni (1963) showed *Bt: Branchlet
that copulation with any male, sterile or fertile, location: 1- (rearrangement).
triggers formation of brown spots. Courtship with- origin: Induced by P 3 2 .
out copulation ineffective; virgin females never discoverer: Bateman, 1950.
show brown spots. No phenotype in males. Pene- references: 1950, DIS 24: 54.
trance 60—80 percent; viability excellent. RK3. 1951, DIS 25: 77.
bt: bent phenotype: Heterozygous female has posteriorly di-
location: 4-1.4 [mapped in diplo-4 triploids by rected branchlet on posterior crossvein as well as
Sturtevant (1951, Proc. Natl. Acad. Sci. U.S. 37: other extra venation. Abdominal segments often
405-7)]. poorly chitinized. Male lethal. RK3A.
origin: Spontaneous. First mutant found on chromo- cytology: Associated with Dp(l;l)Bt - Dp(l;l)3B2-
some 4. Cl;6F6-7.
discoverer: MuIIer, 1914. other information: Phenotype may be Co.
MUTATIONS 37

*bu: bulging phanotype: Eye color reddish brown; brilliant


location: 1-58. orange in combination with en. Good viability.
origin: X ray induced. RK1.
discoverer: Muller, 2618. burnt orange: see Jbuo
references: 1935, DIS 3: 29. bv: krmvis
phenotype: Eyes rough and bulging. Semilethal. location: 3-102.7 (recalculated from Sturtevant,
RK3. 1956, Genetics 41: 118-23).
bv: see Hn™ discoverer: Bridges, 33e25.
bu-w61): see vs61i phenotype: Bristles uniformly short and stubby.
Bubble: see Bb Body chunky. Hatches late but viability excellent.
bubble wing: see vs^ij RK1.
buckled: see bk bw: brown
buckledlike: see bkl location: 2-104.5.
bul: bulge discoverer: Waaler, 19J15,
location: 3-43.6. references: 1921, Hereditas 2: 391-94.
origin: Spontaneous. Sturtevant and Beadle, 1939, An Introduction to
discoverer: Spencer, 36d28. Genetics, Saunders, p. 64 (fig.).
references: 1937, DIS 7: 6. phenotype: Eye color light brownish wine on emer-
Curry, 1939, DIS 12: 45. gence, darkening to garnet. Red pigments lacking;
phenotype: Eyes very large and bulging; facets ommochromes at 87 percent normal level (Nolte,
rounded, in irregular rows, and some quite large. 1954, J. Genet. 52: 111-26). Adult testes and
Wing margin heavy; end of wing somewhat squared vasa colorless. Larval Malpighian tubules pale
off to L3. RK3. yellow (Beadle, 1937, Genetics 22: 587-611).
bulbP: bulge-bumpy Produces white eyes in combination with v, en, or
origin: Spontaneous, st. Eye color autonomous when transplanted into
discoverer: E. H. Grell, 1955. wild-type host (Beadle and Ephrussi, 1936,
synonym: bp, Genetics 21: 230). RK1.
references: 1955, DIS 29: 72. cytology: Placed between 59D4 and 59E1 by Bridges
phenotype: About one-half the eye surface erupted [1937, Cytologia (Tokyo), Fujii Jub. Vol. 2: 7 4 5 -
into irregular yellowish blisters. Facets larger 55], on the basis of its exclusion from the inner
than normal in nonblistered areas. Homozygotes inversion of In(2LR)bw^l = ln(2LR)21C8-
occur with 1 percent of expected frequency. Sur- Dl;60Dl-2 + In(2LR)40F;59D4-El and its inclusion
viving homozygotes vigorous and male fertility in In(2R)bwvOe2 = ln(2R)41A-B;59D6-El. Based
high; females lay eggs abundantly, but only rarely on the study of bw rearrangements, Slatis (1955,
does an egg hatch. RK3. Genetics 40: 5-23) tentatively places bw in 59D9,
bulging: see bu 10, or 11.
other information: Separable into at least two sub-
*buo: burnt orange units by recombination with bw and bw75 about
location: 2-57.1. 0.001 units to the left of bw** and
origin: Spontaneous. (Divelbiss, 1961, Genetics 46: 861).
discoverer: T. Hinton and Kleiner, 1941.
references: Hinton, 1942, DIS 16: 48. origin: Spontaneous.
phenotype: Eye color bright orange-brown. Mal- discoverer: P. R. Sturtevant, 1921.
pighian tubules colorless in larva (Brehme and references: Morgan, Bridges, and Sturtevant, 1925,
Demerec, 1942, Growth 6: 351-56). RK2. Bibliog. Genet. 2: 220.
other information: Not an allele of en. Allelism phenotype: Darker and redder than bw. RK1.
with ltd (2-56) apparently never tested.
bur: burgundy origin: X ray induced,
location: 2-55.7. discoverer: Demerec, 28dl3.
origin: Ultraviolet induced. phenotype: Like bw2. RKl.
discoverer: Edmondson and Meyer, 49c.
references: Meyer and Edmondson, 1949, DIS 23: origin: Spontaneous,
60. discoverer: Emerson, 32dl9,
phenotype: Eye color dull, darkish brown, like pr; phenotype: Like bw*. RKl.
brilliant orange in combination with en. Classifi- bw*
cation and viability excellent. Fertility of females origin: Spontaneous.
good; of males, variable. RK1. discoverer: Mohr, 31k28.
other information: Not allelic to It, ltd, or pr. phenotype: bw*/bw* is wild type. bw4fbws is
bur2 purpleoid-like. bw4fbw like bw but darker. RK3.
origin: Spontaneous. other information: aw"* originally found in bw4/bw$
discoverer: Hall, 60h. combination and called purpleoid-like. bw4 and
references: Meyer, I960, DIS 34: 52. bw-5 separated by Bridges.
38 GENETIC VARIATIONS OF DROSOPH/LA MELANOGASTER

phenotype: Homozygote not clearly distinguishable


origin: Spontaneous. from wild type. Also wild type in combination
discoverer: Mohr, 31k28. with bw24 and bw7s. Produces a phenotype inter-
phenotype: bws/bw4 is purpleoid-like (see bw4); mediate between bw and wild type in combination
bws/bw is light yellowish brown; bws/+ is wild with bw and bw^l. RK3.
type; bwS/bw* is lethal. RK2A. other information: Shown to lie to right of bw and
cytology: Associated with a deficiency or an inver- bw7S by recombination; nonrecombinant reversions
sion involving 59E1 to 59F1 (Schultz). recovered from bw*9/hw81 (Divelbiss, 1961,
*bw24 Genetics 46: 861).
origin: X ray induced. bw61i: see vs61J
discoverer: Slatis, 1950.
references: 1955, Genetics 40: 5-23. origin: X ray induced.
phenotype: Wild type in combination with bwS9 and discoverer: Slatis.
bw7S, but shows intermediate phenotype in combi- references: 1955, Genetics 40: 5-23.
nation with bw and bwai. Studied only in males phenotype: Like bw. RK1.
(see cytology). RK3A. cytology: Salivary chromosomes normal.
cytology: Associated with but presumably w72
separable from T(Y;2)R24 = T(Y;2)45A;51E. origin: X ray induced.
discoverer: Slatis.
origin: Spontaneous. references: 1955, Genetics 40: 5—23.
discoverer: Clancy, 37g26. phenotype: Like bw. RK1.
references: 1938, DIS 10: 55. cytology: Salivary chromosomes normal.
phenotype: Eye color like bw but darker (like
bw2 ?). Produces yellowish pigment in combina- origin: X ray induced.
tion with v. RK1. discoverer: Slatis, 50f25.
references: 1951, DIS 25: 75.
origin: Spontaneous in In(2L)Cy 4- In(2R)Cy, 1955, Genetics 40: 5-23.
Cy en2 sp2. phenotype: Intermediate allele of bw. bw7*/bw7s
discoverer: Ives, 45a. darker than bw7* /bw; about 1 percent of bw7* /bw
references: 1945, DIS 19: 46. flies show twin spots in eye. RK1.
Ives and Scott, 1948, DIS 22: 71. other information: Located to the left of bw81 and
Ives and Evans, 1951, DIS 25: 107. bw59 by recombination (Divelbiss, 1961, Genetics
phenotype: bw4Sa/bw is brown; bW4Sa/bw4 is wild 46: 861).
type; therefore bw45* behaves like bw4. Homozy-
gote has not been tested because bw45* has not origin: X ray induced.
been separated from Cy* RK3A. discoverer: Slatis, 50hl.
other information: Probably occurred simulta- references: 1951, DIS 25: 75.
neously with or45*. 1955, Genetics 40: 5-23.
phenotype: Intermediate allele; more extreme than
bw7*. RK1.
origin: Spontaneous. other information: Located to the right of bw and
discoverer: Ives, 47j. bw7* by recombination; bwsl /bw59 produces non-
references: Ives and Scott, 1948, DIS 22: 71. recombinant reversions (Divelbiss, 1961, Genetics
Ives and Evans, 1951, DIS 25: 107. 46: 861).
phenotype: Brown in combination with In(2L)Cy + *bw+21
In(2R)Cy, Cy en2 bw4** sp2 or4**, but wild type in origin: Isoallele of bw recovered from an X-ray-
combination with bw, bw2b, bw4, or and or4*B. induced brown-Variegated mosaic.
Homozygous normal. RK3. discoverer: Slatis.
other information: Several alleles of this type with references: 1955, Genetics 40: 5-23,
varying degrees of expression found by Ives in a phenotype: Normal, but gives less pigment in
natural population from South Amherst, Mass. combination with variegating alleles than other
bw+ alleles.
origin: Spontaneous, cytology: Salivary chromosomes normal.
discoverer: Clancy, 53i. bw°: brown-amber
references: 1960, DIS 34: 48. origin: Spontaneous.
phenotype: Intermediate between few and bw37t discoverer: R. C. King, 48fl5.
both alone and in combination with v. RK1. references: Poulson and King, 1948, DIS 22: 54.
phenotype: Eye color light brownish yellow. Adult
origin: X ray induced, testes and vasa colorless. Larval Malpighian
discoverer: Slatis, 50fl6. tubules slightly paler yellow than wild type.
references: 1951, Dig 25: 75. bW/bw gives eye color slightly lighter than bw.
1955, Genetics 4Ch 5-23. RK1.
MUTATIONS 39

: brown-Auburn phenotype: Like bw except testis sheath pale


origin: X ray induced. yellow in older males. Gives some pigmentation
discoverer: Dubinin, of eyes in combination with v. RK1.
synonym: A; Pm^'. by/R3: brown-Rearranged
references: Dubinin and Heptner, 1935, J. Genet. origin: X-ray-induced derivative of bw.
30: 423-46 (fig.). discoverer: Slatis, 48kl6.
Dubinin, 1936, Biol. Zh. (Moscow) 5: 851-74. references: 1955, Genetics 40: 5-23.
phenotype: Nearly uniform brown, but with extra Y phenotype: Heterozygotes with 6w+ strongly varie-
chromosome shows strong variegation. Homozy- gated. Homozygous lethal; lethal or semilethal in
gote usually lethal. RK1A. combination with other bw rearrangements. RK1A.
cytology: Associated with In(2R)bwA = cytology: Associated with In(2LR)bwR3 =
In(2R)41;59D. In(2LR)40F;51F;55E;57E;58D8-9.
*bwR4
*bwAD: brown of A. Das origin: X-ray-induced derivative of bw.
origin: Spontaneous. discoverer: Slatis.
discoverer: Das, 63a7. references: 1955, Genetics 40: 5—23.
synonym: bw3^. phenotype: Expression in bwR4/+ varies from
references: Sarkar, 1963, DIS 38: 28. moderate to strong, depending on origin of bw.
phenotype: Eye color light brown, darkening with Homozygote brown and almost completely lethal.
age. RK1. RK1A.
cytology: Associated with T(2;3)bwR4 =
*bwCB: brown-Chester Beatty T(2;3)S9B2-3;80-81.
origin: Induced by 2-chloroethyl methanesulfonate *bwR 7 2
(CB. 1506). origin: X ray induced.
discoverer: Reddi. discoverer: Slatis.
references: 1960, DIS 34: 53. references: 1955, Genetics 40: 5-23.
phenotype: Lethal allele of bw. RK2. phenotype: Weakly mottled in bwR12/+ heterozy-
cytology: Salivary chromosomes normal (Slizynska). gote; bwR12 homozygote somewhat more extreme.
bwD: brown-Dominant Homozygote semilethal. RK1A.
origin: Spontaneous. cytology: Associated with T(2;3)bwRl2 =
discoverer: T. Hinton, 1940. T(2;3)59D;80C.
references: 1940, DIS 13: 49.
1942, DIS 16: 48.
Slatis, 1955, Genetics 40: 246-51. origin: X ray induced.
phenotype: Eye color varies with age from purple to discoverer: Slatis.
brown. Shows slight variegation in combination references: 1955, Genetics 40: 5—23.
with st (Slatis, 1955). Wings pebbled. Variegation phenotype: Variegation intermediate. Homozygous
suppressed by extra Y chromosomes (Brosseau, lethal. RK1A.
1959, DIS 33: 123). Homozygote viable and fertile. cytology: Associated with T(2;3)bwRl* -
Larval Malpighian tubules bright yellow (Brehme T(2;3)59E2-3;80.
and Demerec, 1942, Growth 6: 351-56). RK1A.
cytology: Shultz reports an extra band in 59E that origin: X ray induced.
tends to pair with a band in the homolog, sug- discoverer: Slatis.
gesting a duplication of one band from 59E. Slatis references: 1955, Genetics 40: 5—23.
(1955) reports insertion of three or four bands, phenotype: Variegation intermediate. Homozygote
probably of heterochromatic origin. Reverts to lethal. RK1A.
wild type when extra bands separated from bw cytology: Associated with T(2;3)bwRl$ -
locus (Hinton and GoodSmith, 1950, J. Exptl. T(2;3)59D;80C.
Zool. 114: 103-14).
few**58; brown of Meyer origin: X ray induced.
origin: Spontaneous. discoverer: Slatis.
discoverer: Meyer, 58k. references: 1955, Genetics 40: 5—23.
references: 1959, DIS 33: 97. phenotype: Variegation intermediate. Homozygotes
phenotype: Intermediate allele. Reddish brown lethal. RK1A.
eyes (like g) of somewhat reduced size. May cytology: Associated with ln(2)bwRla =
overlap wild type in old crowded cultures. Testis In(2)40F-41A;59E4-Fl.
sheath light yellow in young but dark in old OWR20
males. Good viability and fertility. RK2. origin: X ray induced.
bwMiS9: brown of Mischaikow discoverer: Slatis.
origin: Spontaneous. references: 1955, Genetics 40: 5—23.
discoverer: Mischaikow, 59e. phenotype: Variegation weak. Homozygote lethal.
reference*: 1959, DIS 33: 97. RK2A.
40 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

cytology: Associated with In(2LR)bwR2° = references: 1955, Genetics 40: 5—23.


In(2LR)40D;59D5-6. phenotype: Variegated in heterozygotes with bw + ;
%WR2S strongly variegated in homozygotes, and in
origin: X-ray-induced derivative of bw. heterozygotes with bw and other variegated
discoverer: Slatis. browns. Homozygotes show melanotic clots of
references: 1955, Genetics 40: 5—23. dried hemolymph. RK1A.
phenotype: Variegation not described. Homozygote cytology: Associated with In(2)bwR47 =
like bw. RK2A. In(2)40-41;59DU-El.
cytology: Associated with T(2;4)bwR25 =
T(2;4)59D;101E. origin: X ray induced.
discoverer: Slatis.
origin: X-ray-induced derivative of bw. references: 1955, Genetics 40: 5—23.
discoverer: Slatis. cytology: Analysis incomplete; one break at
references: 1955, Genetics 40: 5—23. 59D2-3.
phenotype: Variegation intermediate; brown in wR 55
combination with bw. RK1A. origin: X ray induced.
cytology: Associated with T(Y;2)bwR27 = discoverer: Slatis, 50d23.
T(Y;2)59D11-E1. references: 1955, Genetics 40: 5—23.
phenotype: bwRS5/+ and bwR55/bw usually show
origin: X ray induced. small amounts of variegated eye tissue; but
discoverer: Slatis. occasionally entire eye nearly lacks red pigment.
references: 1955, Genetics 40: 5-23. Homozygous lethal. RK1A.
phenotype: Variegation intermediate to slight. cytology: Associated with In(2LR)bwRSS =
Homozygote shows extreme variegation and In(2LR)24El-D;42E + In(2R)40F-41A;59D4-5.
reduced viability. RK1A.
cytology: Associated with In(2R)bwR32 =
In(2R)41A;59D. origin: X ray induced.
*by,R33 discoverer: Slatis.
origin: X ray induced. references: 1955, Genetics 40: 5—23.
discoverer: Slatis. phenotype: Variegation intermediate in heterozy-
references: 1955, Genetics 40: 5—23. gote, extreme in homozygote. RK1A.
cytology: Associated with Ia(2R)bwR33 — cytology: Associated with In(2)bwRS6 =
In(2R)41;59D'E. In(2)40F-41A; 59D-E.

origin: X ray induced. origin: X ray induced.


discoverer: Slatis. discoverer: Slatis.
references: 1955, Genetics 40: 5—23. references: 1955, Genetics 40: 5—23.
phenotype: Variegation intermediate to slight; phenotype: Variegation weak in heterozygote.
homozygotes show more extreme variegation and RK2A.
reduced viability. RK1A. cytology: Associated with T(Y;2)bwRS7 =
cytology: Associated with In(2)bwR35 = T(Y;2)59D5-6.
In(2)4QF-41A;59Dl 1-E1.
origin: X ray induced.
origin: X ray induced. discoverer: Slatis,
discoverer: Slatis. references: 1955, Genetics 40: 5—23.
references: 1955, Genetics 40: 5-23. phenotype: Variegation in heterozygote weak.
phenotype: Shows slight dilution of eye pigment in Homozygote lethal. RK1A.
bwR4°f+ heterozygote. Homozygote presumably cytology: Associated with T(2;3;4)bwRS8 =
lethal. RK2A. T(2;3;4)59D;6 5;101C.
cytology: Associated with Dl(2R)bwR40 -
Df{2R)59C5~6;59E2-3. origin: X ray induced.
discoverer: Slatis.
origin: X-ray-induced derivative of bw. references: 1955, Genetics 40: 5—23.
discoverer: Slatis. phenotype: Variegation in heterozygote moderate to
references: 1955, Genetics 40: 5-23. strong. Homozygote lethal. RK1A.
pfoeno*ype: Variegation of bwR4$/+ slight. cytology: Associated with In(2)bwR67 -
Homoxygote lethal. RK2A. In(2)40F'41A;59E4-Fl.
cytology: Associated with
ln(2)4QF-41A;59E3-4. origin: X ray induced.
discoverer: Slatis.
origin: X ray induced. references: 1955, Genetics 40: 5—23.
discoverer: Slatis, 5CM16. phenotype: Like bw. RK1A.
MUTATIONS 41

cytology: Associated with complex rearrangement references: 1930, J. Genet. 22: 299-334 (fig.).
with one break near 58F; mutation and rearrange- Glass, 1934, J. Genet. 28: 69-112 (fig.).
ment presumably independently induced. 1934, Am. Naturalist 68: 107-14.
phenotype: Eye color like bwvi but with browner
origin: X ray induced. ground and numerous discrete very dark granular
discoverer: Slatis. spots; more variable. Larval Malpighian tubules
references: 1955, Genetics 40: 5-23. normal (Glass, Brehme). Homozygote generally
cytology: Associated with In(2)bwR73 = lethal. RK1A.
In(2)40F-41A;59E4-Fl. cytology: Associated with T(2;3)bwV3.
wV*
origin: X ray induced. origin: X ray induced.
discoverer: Slatis, 50g26. discoverer: Patterson, 1929.
references: 1955, Genetics 40: 5-23. synonym: Rosy.
phenotype: Strongly variegated in combination with references: Glass, 1934, J. Genet. 28: 69-112 (fig.).
either bw or bw+. Homozygous lethal and lethal 1934, Am. Naturalist 68: 107-14.
or semilethal with most other brown-Variegateds. phenotype: Eye color yellow-brown ground with
RK1A. numerous dark granular spots. Homozygous lethal
cytology: Associated with in 95 percent of cases; survivors have pale rose-
ln(2)40F-41A;59F2-3. brown eye color with few darker spots, which
change to white and yellowish in combination with
: brown-Variegated st. Larval Malpighian tubules normal (Glass,
origin: X ray induced. Brehme). RK1A.
discoverer: Muller, 1929. cytology: Associated with T(2;3)bwV44
synonym: Pm; Plum.
references: 1930, J. Genet. 22: 299-334 (fig.). origin: X ray induced.
Glass, 1934, J. Genet. 28: 69-112 (fig.). discoverer: Patterson, 1929.
1934, Am. Naturalist 68: 107-14. synonym: 143a.
Bridges, 1937, Cytologia (Tokyo), Fujii Jub. references: Glass, 1934, J. Genet. 28: 69-112 (fig.).
Vol. 2: 745-55. 1934, Am. Naturalist 68: 107-14.
phenotype: Eye color like bw or pr, mottled with phenotype: Like bwv3 and bwv* but browner
darker spots that deepen in red color with age. ground color. Rare homozygous survivors. Larval
With sf or v, has pale orange ground with dark Malpighian tubules normal (Glass, Brehme).
orange spots. Extra Y chromosome, as with other RK1A.
variegated browns, suppresses brown color, giving cytology: Associated with T(2;3)bwV5w
red eye sparsely speckled or splotched with *bwV6
darker spots. Larval Malpighian tubules normal origin: X-ray-induced derivative of bw.
(Glass, Brehme). Generally lethal homozygous discoverer: Moore, 1929.
and in combination with other brown-Variegateds. synonym: A34.
Heterozygotes fully viable and fertile. RK1A. references: Glass, 1934, J. Genet. 28: 69-112 (fig.).
cytology: Associated with In(2LR)bwV1 = 1934, Am. Naturalist 68: 107-14.
In(2LR)21C8-Dl ;60Dl-2 + In(2LR)40F;59D4-El phenotype: Like bxvVS. RK1A.
(Schultz and Bridges). cytology: Associated with T(2;3)bwv6.
other information: Lost by reversion.

origin: X ray induced. discoverer: Winchester, 1932.


discoverer: Harris, 1929. phenotype: Eye color like bwvs. Homozygote
synonym: Discolored. moderately viable; eye color of pale rose-brown
references: Muller, 1930, J. Genet. 22: 299-334 ground with few spots. RK1A.
(fig.). cytology: Associated with In(2R)bwv7\
Glass, 1934, J. Genet. 28: 69-112 (fig.).
1934, Am. Naturalist 68: 107-14. origin: X ray induced,
phenotype: Eye color similar to bwvl, but less discoverer: Levy, 1932.
purplish, v, bwv2 has yellower ground color and references: 1935, DIS 3: 7.
browner spots than bwvi; st. Homozygote nearly phenotype: Like bwV^. RK1A.
always lethal. Malpighian tubules of larvae cytology: Associated with T(2;3)bwV8.
normal (Glass, Brehme). RK1A.
cytology: Not studied. Shown genetically to be origin: Radium induced.
associated witn In(2R)bwv'2. discoverer: Van Atta, 291.
synonym: Cream; Dilute-1.
origin: X ray induced, references: Hanson and Winkelman, 1929,
discoverer: Muller, 1929. J. Heredity 20: 277-86.
synonym: Tarnished. Van Atta, 1932, Genetics 17: 637-59.
42 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

phenotype: Like bwVl, RK1A. except in the presence of an extra Y.


cytology: Associated with In(2LR)bv/V29l, shows variegation for light (Schultz, 1936). Via-
*bwV30a bility excellent. Homozygous lethal except for
origin: X ray induced. rare survivors with light eye color and somatic
discoverer: Oliver, 30a. abnormalities. Larval Malpighian tubules some-
phenotype: Almost homogeneous brown eye color. what lighter than wild type but not useful in
RK3(A). classification (Brehme and Demerec, 1942, Growth
bwV30k1 6: 351-56). RK1A.
origin: X ray induced. cytology: Associated with In(2LR)bwv32g =
discoverer: Van Atta, 30kl. In(2LR)40F;59E.
synonym: Dilute-2.
references: 1932, Genetics 17: 637-59. origin: X ray induced in In(2L)Cy +In(2R)Cy.
phenotype: Like bwVl. RK1A. discoverer: Oliver, 34k22.
cytology: Associated with In(2LR)bwV30k*-. synonym: Var34k22,
bwV3Okl0 references: 1937, DIS 7: 19.
origin: X ray induced. phenotype: Eye color nearly homogeneous brown,
discoverer: Van Atta, 30kl0. slightly mottled. Larval Malpighian tubules
synonym: Dilute-3. normal (Glass, Brehme). RK1A.
references: 1932, Genetics 17: 637-59. cytology: Associated with In(2R)bwv34k =
phenotype: Eye color blotched heavily with large In(2R)41;59E, which carries as an included
patches of red and brown. Homozygous lethal. inversion In(2R)Cy = In(2R)42A2-3;58A4-Bl.
RK1A. *bwV40b
cytology: Associated with In(2R)bwV3Okio% origin: X ray induced.
*bwV30kl2 discoverer: T. Hint on, 40b.
origin: X ray induced. references: Atwood, 1942, DIS 16: 47.
discoverer: Van Atta, 30kl2. phenotype: Eye light brown with some darker
synonym: Dilute-4. variegation. Homozygous lethal. RK1A.
references: 1932, Genetics 17: 637-59. cytology: Associated with ln(2R)bwv40b =
phenotype: Eye color mostly red with a sprinkling In(2R)41A-B;59D-E.
of spots and facets of brown, appearing as dark
spots. Homozygous lethal. RK2A. origin: Gamma ray induced.
cytology: Associated with T(2;3)bwV30kl2, discoverer: Mickey, 54a6.
references: 1963, DIS 38: 29.
origin: X ray induced. phenotype: Variegated for bw. RK1A.
discoverer: Van Atta, 30kl3. cytology: Associated with In(2R)bwv54a =
synonym: Dilute-5. Jn(2R)41A-B;59D4-9.
references: 1932, Genetics 17: 637-59. *bwV54b
phenotype: Eye color mostly red with brown discoverer: Mickey, 54bl2.
spotting. Homozygous lethal. RK2A. references: 1963, DIS 38: 29.
cytology: Associated with T(2;3)bwV30kl3, phenotype: Variegated for bw. RK1A.
cytology: Associated with In(2R)bwv54t> =
origin: X ray induced. In(2R)41A;60D9-ll.
discoverer: Van Atta, 30kl8. *bwV54c
synonym: Dilute-6. origin: Neutron induced.
references: 1932, Genetics 17: 637-59. discoverer: Yanders, 54c5.
phenotype: Eye color mostly red with sprinkling of references: Mickey, 1963, DIS 38: 29.
dark (brown) facets. Homozygous lethal. RK2A. phenotype: Variegated for bw. RK1A.
cytology: Associated with T(2;3;4)bwV30kl8w cytology: Associated with In(2R)bwVS4c =
In(2R)41;59El.
origin: X ray induced.
discoverer: Dobzhansky, 32g6. origin: X ray induced in In(2LR)SMl, al2 Cy en2 sp2.
2
synonym: Pm . discoverer: E. H. Grell, 57e.
references: Schultz and Dobzhansky, 1934, references: Hochman, 1961, DIS 35: 85—86.
Genetics 19: 344-64. Welshons, 1962, Genetics 47: 743-59.
Schultz, 1936, Proc. Natl. Acad. Sci. U.S. 22: phenotype: Eyes brown and mottled. Viability of
27-33. bwvS7e/bwvl about 40 percent at 26°C and 20
phenotype: Dominant eye color like brown but percent at 23.5°; body tends to be small, wings
flecked with darker spots. Shows 18 percent divergent and often not expanded; patches of un-
normal red pigment and 88 percent normal brown pigmented microchaetae. RK2A.
pigment (Nolte, 1954, J. Genet. 52: 127-39). cytology: Salivary chromosomes not examined, but
bwv32t/bw is like bw with very few spots. likely that bwVS7e is the result of a rearrangement
t,wV32g/+ jS easily separable from wild type superimposed on SMI.
MUTATIONS 43

: brown-Variegated Dichaete linked phenotype: Like bw in young flies but much darker
origin: X ray induced. than bw in old individuals. RK1.
discoverer: Oliver, 29k24. other information: Apparently never tested for
synonym: Ic D. allelism with ca (3-100.7).
references: 1932, Z. Induktive Abstammungs- bw-l: see red
Vererbungslehre 61: 447-88. *bwl: bow-legged
phenotype: Like bwvl, but redder and more variable. location: 1-21.9.
Homozygous lethal, but not lethal in combination origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
with other brown-Variegateds. RK2A. phenylalanine (CB. 3025).
cytology: Associated with T(2;3)bwVE>% discoverer: Fahmy, 1955.
other information: Irradiated third chromosome references: 1959, DIS 33: 83.
carried D. phenotype: Poorly viable with shorter divergent
bwVDe I; brown-Variegated of Demerec wings. Bristles thinner and shorter. Legs
origin: X ray induced. shortened and either femur or tibia, or both, bow-
discoverer: Demerec, 33i28. shaped. Males sterile. RK3.
references: Bridges, 1937, Cytologia (Tokyo),
Fujii Jub. Vol. 2: 745-55.
phenotype: Eye color variegated brown and red;
extreme dominant brown with pebbled wing.
Lethal when homozygous and in combination
with bw*. RK2A.
cytology: Associated with In(2R)bwV^el -
In(2R)41 B2-Cl;59E2-4.

origin: X ray induced.


discoverer: Demerec, 33J14.
references: Bridges, 1937, Cytologia (Tokyo),
Fujii Jub. Vol. 2: 745-55.
phenotype: Eye color mosaic of brown and dark
brown patches. RK2A.
cytology: Associated with In.(2R)bwVDf>2 -
In(2R)41A-B;59D6-El.

origin: X ray induced.


discoverer: Demerec, 33jl4.
phenotype: Eye color variegated brown and red. In bx: bithorax
bwVDe3/bw5, wings fail to expand; joints and From Bridges and Morgan, 1923, Carnegie Inst. Wash.
sutures weak with melanotic clots. Homozygous Pubi. No. 327: 152.
lethal. RK2A.
cytology: Associated with T(2;3)bwvE>e3 = bx: bithorax
T(2;3)59D;81F. location: 3-58.8.
discoverer: Bridges, 15i22.
origin: X ray induced. references: Bridges and Morgan, 1923, Carnegie
discoverer: Demerec, 33k22. Inst. Wash. Publ. No. 327: 137, 152 (fig.).
phenotype: Eye color red with brown s p o t s ; Morgan, Bridges, and Sturtevant, 1925, Bibliog.
bwVDe4/bw and X/0; bwVDe4/+ a r e brown. Genet. 2: 79, 214 (fig.).
Homozygous lethal. RK2A. Lewis, 1951, Cold Spring Harbor Symp. Quant.
cytology: Associated with T(2;3)bwyDe4 - Biol. 16: 159-74 (fig.).
T(2;3)59D2~4;80. 1963, Am. Zoologist 3: 33-56 (fig.).
*bwVh brown-Variegated of Ives phenotype: Anterior half of metathorax becomes
origin: Spontaneous. mesothoracic and posterior half remains un-
discoverer: Ives, 38113. changed. This results in the appearance, between
references: 1950, DIS 24: 58. the scutellum and the first abdominal segment, of
phenotype: Like bwD. RKlA. rudimentary anterior mesothoracic elements (i.e.,
cytology: Associated with ln(2R)bwvl - mesonotum and scutellum), with the proper bristles
In(2R)41A;59D (T. Hinton). and hairs. Balancers directed ventrally, enlarged,
*bw-b: brown-b vesiculate or winglike, with typical but rudimen-
location: 3- (between 97.0 and 104.2). tary wing venation and bristles. Metathoracic
origin: X ray induced. tibia has a mesothoracic tibial bristle. Variable,
discoverer: E. L. Smith, 34f. overlapping wild type at 25° and more so at 19°C.
references: Robertson, 1935, DIS 4: 15. Dorsal metathoracic disk of mature larva is 60 per-
Smith and Robertson, 1938, Genetics 23: 167. cent larger than wild type IChen, 1929, J. Morphol.
44 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

47: 135—99 (fig.)]. Thoracic musculature studied


by El Shatoury [l956, J. Embryol. Exptl. Morphol. origin: Spontaneous.
4: 228—39 (fig.)] and Pantelouris and Waddington discoverer: Gunson.
[l955, Arch. Entwicklungsmech. Organ. 147: 539— references: 1952, DIS 26: 63.
46 (fig.)]. Bristle patterns studied by Waddington phenotype: Penetrance 0.5 percent at 16°, 1 percent
(1962, New Patterns in Genetics and Development. at 20°, and 29 percent at 25°C. RK3.
Columbia University Press). Slight expression in bxD>: see Ubx
combination with bw34e and bww (Hollander, 1937, *bxw: bithorax-W/scons/n
DIS 8: 77), and no interaction as trans heterozy- origin: Spontaneous; recovered in one third chromo-
gote with bxd or Vbx (E. B. Lewis). RK3. some of the female in whose other third chromo-
cytology: Located close to if not within the 89E1-2 some Ubx was first recovered.
doublet (Lewis, 1951). discoverer: Hollander, 1934.
other information: The leftmost member of a pseudo- references: 1937, DIS 8: 8, 77.
allelic series including from left to right, bx, Cbx, phenotype: Metathorax developed more than any
Ubx, bxd, and pbx. other bx allele and strongly bristled. Halteres
directed ventrally, large, flat, and winglike, with
veins and bristles. Mesothorax shows clear stripe
or bifida condition. Wings usually spread, often
bx Cbx Ubx bxd pbx dragging. Sternopleural and tibial and ventral
bristles of third leg resemble those of normal
0.01 0.008 0.006 0.006 second leg. bxw/bxd = +. Viability and fertility
fair. RK2.
fix; Beadex
Map of the bx region location: 1-59.4.
From Lewis, 1963, Am. Zoologist 3: 33-56. origin: Spontaneous.
discoverer: Bridges, 23a3.
references: Morgan, Bridges, and Sturtevant, 1925,
origin: Spontaneous. Bibliog. Genet. 2: 219.
discoverer: Stern, 25b2. Green, 1953, Genetics 38: 91-105 (fig.).
references: 1935, DIS 3: 29. phenotype: Male and homozygous female with
Lewis, 1951, Cold Spring Harbor Symp. Quant. Beaded-like wings; long and narrow and excised
Biol. 16: 159-74 (fig.). along both margins. Male and homozygous female
1955, Am. Naturalist 89: 73-89. fully viable. Heterozygous female less extreme
1963, Am. Zoologist 3: 33-56 (fig.). and overlaps wild type. Some venation abnor-
phenotype: Extreme expression of 6x metathoracic mality. Development studied by Goldschmidt
effect. Little variability. Viability low but can be [1935, Biol. Zentr. 55: 535-54 (fig.)]. According
maintained homozygous. Balancers enlarged in to Waddington (1940), embryological effect is same
heterozygote. Interactions with other bx pseudo- as that of vg. RK2 (RK3 as Sx/+).
alleles described by Lewis (1951, 1955, 1963). Al- cytology: Locus in salivary region 17A, B, or C,
most completely suppressed by su(Hw)2. RK3. based on limits of Dp(l;l)Bxr49k (E. B. Lewis).
Salivary chromosomes of Bx normal. (Lewis).

origin: Spontaneous.
discoverer: Schultz, 34e20.
references: 1935, DIS 4: 6.
Lewis, 1951, Cold Spring Harbor Symp. Quant.
Biol. 16: 159-74.
1955, Am. Naturalist 89: 73-89.
1963, Am. Zoologist 3: 33-56 (fig.).
phenotype: Metathoracic outpushing is a uniform,
narrow hairy band. Balancers depressed, inflated, Bxh Beadex-2
with elongated pointed tip and heavy line of costal Edith M. Wallace, unpublished.
bristles. Base of third leg shows sternopleural
bristles like those of normal second leg. Meta-
thoracic development of bx34e/bx340 > origin: Spontaneous.
bx3«»/bxw> bx34°/bx (Hollander, 1937, DIS 8: discoverer: Mohr, 24129.
77). bx^4e/Ubx has round, flat, winglike references: 1927, Nyt Mag. Natur 65: 265—74.
halteres; bx^4e i/bx/+ + = Ubx/+ (Lewis). Comple- Green, 1953, Z. Induktive Abstammungs-
ments hxd. Reasonable viability and fecundity. Verebungslehre 85: 435—49.
Highly constant expression and easy separability. phenotype: Wings of males and homozygous females
Expression increased slightly at 29°C, decreased narrowed by marginal excision. Wings often bubbly
at 15°C (Villee, 1943, Anat. Record 87: 475). RK2. and ragged. Homozygous female fully viable.
MUTATIONS 45
2
Bx /+ less extreme; overlaps wild type. Classi- other information: Bx/Dp(l;l)Bxr produces recom-
fiable in a single dose in triploids (Schultz, 1934, binants of genotype Bx+Bx and BxBx+, which are
DIS 1: 55). RK1 (RK3 as Bx2/+). more extreme than Bx. Same holds for
Bx2/Dp(l;l)Bx*.
origin: Spontaneous.
discoverer: Gershenson, 1927.
references: Gaissinovitsch and Gershenson, 1928,
Biol. Zentr. 48: 385-87 (fig.).
phenotype: Extreme allele usually without the
bubbles in the wing. Shortened L5 a constant
character (few Bx2 show this). Wings more pointed
than Bx2 and hairs at tip of wing clumped.
Scalloping visible in prepupal wing bud
[Waddington, 1940, J. Genet. 41: 75-139 (fig)],
fully separable. RK1. Bx^9k: Beadex-recessive 49k
From Green, 1953, Z. Induktive Abstammungs-
origin: Spontaneous. Vererbungslehre 85: 435-49.
discoverer. T. J. Lee, 59h.
references: 1964, DIS 39: 60.
phenotype: Like Bx. RK2. origin: Spontaneous.
discoverer: Mossige, 49k22.
*Bxc: Beadex of Catcheside synonym: Bxl2.
origin: Spontaneous. references: 1950, DIS 24: 61.
discoverer: Catcheside, 39c3. Green, 1953, Z. Induktive Abstammungs-
references: 1939, DIS 12: 49. Vererbungslehre 85: 435-49 (fig.).
phenotype: Posterior wing margin excised as far as phenotype: Slight scalloping of posterior wing
and including most of L5; end of wing notched and margin only; overlaps wild type. RK3A.
anterior margin weakly excised. Wings blistered, cytology: Associated with Dp(l;l)BxrJ*9k -
especially basally. BxC/+ is like Bx^/Y. RK1. Dp(l;l)17A;17C (E. B. Lewis).
other information: This duplication undergoes
Bx-I: Beadex of Jollos unequal crossing over readily and forms triplica-
origin: Induced by heat treatment. tions and quadruplications. Duplication is reces-
discoverer: Jollos, 1930. sive; triplication is dominant. Phenotypic interac-
synonym: Ptd: Pointedoid. tion with Bx same as for BxT.
references: 1933, Naturwissenschaften 21: 831—34.
Gottschewski, 1935, DIS 4: 7, 14, 16.
Jollos and Waletsky, 1937, DIS 8: 9.
phenotype: Wings reduced to slender strip; only
posterior cell present at tip. Femur shortened or
legs otherwise abnormal, especially third pair.
Homozygous female viable. Interacts with bi to
give more nearly normal wings. Embryology like
Bx [Goldschmidt, 1935, Biol. Zentr. 55: 535-54;
Waddington, 1940, J. Genet. 41: 75-139 (fig.)].
RK1.

Bxr: Beadex-recessive
origin: Spontaneous.
discoverer: Ives, 35k.
references: 1937, DIS 7: 6.
Green, 1952, Proc. Natl. Acad. Sci. U.S. 38:
949-53.
1953, Genetics 38: 91-105 (fig.).
phenotype: Bxr/+ is normal. Male and homozygous
female show less extreme narrowing of wings than hxd: hithoraxoid
Bx. Anterior crossvein short and thickened and From Bridges and Morgan, 1923, Carnegie Inst. Wash.
that region blistered. May overlap wild type in Publ. No. 327: 225.
old crowded cultures at 25°, more extreme at 19°C.
RK3A. bxd: bithoraxoid
cytology: Associated with Dp(t;l)Bxr ~ location: 3-58.8.
Dp(l;l)17A;17E-F (Green, 1953, determined by origin: Spontaneous.
E. B. Lewis). discoverer: Bridges, 19127.
46 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: Bridges and Morgan, 1923, Carnegie references: 1945, Univ. Calif. (Berkeley) Publ.
Inst. Wash. Publ. No. 327: 225 (fig.). Zool. 49: 507, 520.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Like a strong Bx. RK2.
Genet. 2: 214 (fig.), 219.
Lewis, 1951, Cold Spring Harbor Symp. Quant. Bxl: see Ubx
Biol. 16: 159-74 (fig.). by: blistery
1955, Am. Naturalist 89: 73-89. location: 3-48.7.
1963, Am. Zoologist 3: 33-56 (fig.). origin: Spontaneous.
phenotype: Posterior portion of metathorax becomes discoverer: Glass, 33a.
mesothoracic; anterior metathorax unaffected. references: 1934, DIS 2: 8.
Enlarged metathoracic postnotum forms two phenotype: Wings blistered in subterminal region,
wedges of tissue, devoid of hairs and bristles, and wing surface dusky and warped. Thorax
meeting at the mid-dorsal line. Balancers enlarged humpy. RK1.
into circular disks that are cupped and bent down- *by46h
ward and show venation but are bare of bristles origin: Spontaneous.
except for weak ones at rear margin. Stalk of discoverer: Ives, 46hl5.
balancer wide and flat. First abdominal segment references: 1948, DIS 22: 53.
shows thoracic modification (modification of phenotype: Like by but without thoracic effect.
anterior portion is metathoracic-like, posterior RK1.
portion is partially mesothoracic-like). Rudiments hi: see malbz
of a first pair of abdominal legs and, in certain
combinations, of partially wing-like first abdominal
halteres may occur. Interactions with other bi-
thorax pseudoalleles described by Lewis (1951,
1955, 1963). RIG.
cytology: Locus probably 89E3-4 (E. B. Lewis).

origin: X ray induced.


discoverer: E. B. Lewis.
references: 1951, Cold Spring Harbor Symp. Quant.
Biol. 16: 159-74.
phenotype: More extreme bithoraxoid phenotype
than bxd. RK2A.
cytology: Associated with Tp(3)bxdi00 -
Tp(3)66C;89B5-6;89E2'3. bx+ but not bxd included
in transposed section.

origin: X ray induced in ss bx Su(ss)2.


discoverer: E. B. Lewis,
phenotype: Like hxd*O°. RK2A.
cytology: Associated with T(3;4)89E = T(3;4)89E2-
3;101F. c; curved
From Bridges and Morgan, 1919, Carnegie Inst. Wash.
origin: X ray induced in pP bx sr ea. Pubi. No. 278: 165.
discoverer: E. B. Lewis.
phenotype: Like bxd10®. RK2A. c: curved
cytology: Associated with Tp(3)bxd11(> = Tp(3)89E2- location: 2-75.5.
3;91C7-Dl;92A2-3. origin: Spontaneous.
discoverer: Bridges, 11124.
origin: X ray induced. references: Bridges and Morgan, 1919, Carnegie
discoverer: E. B. Lewis. Inst. Wash. Publ. No. 278: 164 (fig.)-
references: 1951, Cold Spring Harbor Symp. Quant. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Biol. 16: 159-74. Genet. 2: 211 (fig.).
1963, Am. Zoologist 3: 33-56 (fig.). phenotype: Wings thin textured, divergent, uplifted
phenotype: hxdm/bxd shows infrequent develop- at base, and curved downward throughout their
ment of abdominal wing-like halteres as well as length. RK1.
abdominal legs. RK2. *C"K: Curved of Krivshenko
hxd*>: see Vbx location: 2- or 3- (rearrangement).
*Bxd: Beadnxoid origin: X ray induced,
location: 1-45. discoverer: Krivshenko, 5513.
origins Spontaneous. references: 1956, DIS 3th 74.
dlscov»r«r: Goldschmidt. synonym: CK.
MUTATIONS 47

phenotype: Wings are thin textured, slightly diver- CHL: see In(2L)NS
gent, uplifted basally, and then curved downward. CIIIRE: see In(3R)C
Homozygous lethal. RK2A. co: claret
cytology: Associated with T(2;3)C-K = location: 3-100.7.
T(2;3)52;76;81;86. origin: Spontaneous.
C( ): Crossover suppressor discoverer: Bridges, 19112.
The terminology originally used for dominant sup- references: Bridges and Morgan, 1923, Carnegie
pressors of crossing over. These effects were Inst. Wash. Publ. No. 327: 219 (fig.).
found to be rearrangements and are so treated here. phenotype: Eye color ruby. Red and brown pig-
The symbol C in this context has been dropped ments 27 and 29 percent of normal levels, respec-
except where included under synonymy. tively (Nolte, 1955, J. Genet. 53: 1-10). With en,
*c(l)a: recessive crossover suppressor for eye color is deep reddish yellow; with bw, trans-
chromosome 1 lucent brownish yellow (Mainx, 1938, Z. Induktive
location: One factor in X and probably several Abstammungs- Vererbungslehre 75: 256—76).
autosomal modifiers, Larval Malpighian tubes colorless (Beadle, 1937,
origin: Spontaneous. Genetics 22: 587—611). Eye color autonomous
discoverer: Bridges, 1916. when larval optic disk from ca is transplanted into
references: Bonnier, 1923, Hereditas 4: 81—110. wild type or v. Wild-type disk in ca not entirely
Morgan, Bridges, and Sturtevant, 1925, Bibliog. autonomous (Beadle and Ephrussi, 1936, Genetics
Genet. 2: 220. 21: 230); ca flies produce less v + substance than
phenotype: Reduces recombination between v and f wild type (Clancy, 1942, Genetics 27: 417-40).
from 23 to 15 percent and between w° and v from Slightly narrow body and pointed wing. RK1.
31 to 10 percent. c(l)a was probably the cause of cytology: Judged to be in 99C-E, based on
a secondary nondisjunction frequency of 15—30 In(3R)cav = In(3R)81F;99C-B (E. B. Lewis).
percent. RK3. co2
other information: Validity of phenotypic descrip- origin: Spontaneous in In(3R)P = In(3R)89C2-
tion seems dubious. 3; 96A18-19.
C(2)R: see In(2R)NS discoverer: Bridges, 32f22.
C(2;3): see In(2L)t phenotype: Eye color like ca. Same body and wing
C(2L)HR: see In(2L)t effect as ca. Larval Malpighian tubes colorless
C(2L)T: see ln(2L)t (Brehme and Demerec, 1942, Growth 6: 351-56).
c(3)G: recessive crossover suppressor in RK1A.
chromosome 3 of Go wen ; clarei-nondisjunctional
location: 3-57.4 (1.0 to the left of sbd2, 4.0 to the origin: X ray induced.
right of cv-c). discoverer: E. B. Lewis and Gencarella.
origin: Spontaneous, references: 1952, Genetics 37: 600—1.
discoverer: Gowen and Gowen, 1917. phenotype: Eye color like ca, but homozygous
synonym: ex (Gowen, 1928, Proc. Natl. Acad. Sci. females lay mostly inviable eggs; surviving
U.S. 14: 475-77). progeny often show results of chromosome nondis-
references: 1922, Am. Naturalist 56: 286-88. junction and loss; meiosis normal in cand/ca
1932, Proc. Intern. Congr. Genet., 6th. Vol. 2: females. Similar in action to ca of Dzosophila
69-70. simulans (Sturtevant, 1929, Z. Wiss. Zool. Abt. A
1933, J. Exptl. Zool. 65: 83-106. 135: 323—56). One experiment yielded 1373
phenotype: In homozygous females, crossing over in progeny of females homozygous for cand, and gave
entire chromosome complement reduced to a small the following results: 42 percent regular offspring,
fraction of normal. Production of triploids and in- 15.0 percent exceptional (X/X/Y and X/0), 32.0
tersexes 300—500 times normal. Nondisjunction in- percent haplo-4, 3.9 percent haplo-4 mosaics, and
creased; egg hatching very low, probably owing to 5.4 percent gynandromorphs. The remaining 0.8
aneuploid zygotes. Core structures characteristic percent included a triploid female, triploid inter-
of electron microscope preparations of normal sexes, probable superaales, and a few other kinds
oocyte nuclei absent in c(3)G/c(3)G oocytes of mosaics. Causes nondisjunction and loss of
(Meyer, 1964, Proc. Eur. Reg. Conf. Electron. major autosomes (Davis, D. G., 1963, Ph.D.
Microscop. 3rd, pp. 461—62). Somatic crossing over Thesis, Univ. Georgia). Hinton and McEarchen
normal (Le Clerc, 1946, Science 103: 553-54). In- (1963, DIS 37: 90) reported haploid-diploid
creased recombination observed in c(3)G/+ females mosaic. Crossing over between X chromosomes
(Hinton, 1962, Genetics 47: 959; 1966, Genetics normal (Davis). cand ovaries transplanted into
53: 157—64). Meiosis in males not affected. RK3. normal host behave autonomously (Roberts, 1962,
cytology: Included within Df(3R)sbd*05 - DIS 36: 112). Chromosome segregation normal in
D((3R)88F9-89Al;89B4-5 (Lewis, 1948, DIS 22: cand males. RK3.
72-73). *: claret-variegated
C2L: see ln(2L)NS origin: X ray induced.
C3: see In(3R)C discoverer: E. B. Lewis.
48 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

phenotype: cav/ca slightly variegated. Can be *Cb: Curled blistered


confused with wild type. cav/cand females pro- location: 1-13.
duce normal progeny- Homozygous lethal. RK3A. origin: Spontaneous.
cytology: Associated with In(3R)cav = discoverer: Villee, 40b.
In(3R)81F;99C-E. references: 1945, DIS 19: 47.
*cal: coal phenotype: Heterozygous or homozygous Cb give
location: 3-59.5. curled and blistered wings only in presence of
origin: Spontaneous. homozygous pxCb. RK3.
discoverer: Grout, 47120. *cbd: cluboid
references: Ives, 1948, DIS 22: 53. location: 3- (about 30 units to right or left of D).
phenotype: Black body color similar to e4. Via- origin: Spontaneous.
bility reduced slightly. RK2. discoverer: Bridges, 16il5.
canopy wing: see cpw references: Bridges and Morgan, 1923, Carnegie
Inst. Wash. Publ. No. 327: 169.
car: carnation phenotype: Wings not expanded. Dwarfish. Low
location: 1-62.5. fertility. RK3.
origin: X ray induced. *cbf: clubfoot
discoverer: Patterson, 28c20. location: 1-45.
references: 1934, DIS 1: 31. origin: X ray induced.
phenotype: Eye color dark ruby. Body shape and discoverer: Cantor, 46d20.
proportions seem rounded. With st, eye color is references: 1946, DIS 20: 64.
yellow-brown, with bw, brownish yellow to brown phenotype: Leg segments greatly shortened; ab-
(Mainx, 1938, Z. Induktive Abstammungs- normally shaped tarsi and metathoracic legs.
Vererbungslehre 75: 256—76). Maipighian tubes Wings slightly warped, wide in center, and
pale yellow in mature larva (Beadle, 1937, tapering at ends. All flies emerging show both
Genetics 22: 587—611) but hard to distinguish wing and leg effects but expression variable.
from wild type before third instar. Eye color Only about 3 percent of cbt flies eclose. RK3.
autonomous in transplant into wild-type host other information: Not tested for allelism to pi
(Beadle and Ephrussi, 1936, Genetics 21: 230). (1-47.9).
Contains 33 percent wild-type red pigment and 47 Cbx: Contrabithorax
percent brown pigment (Nolte, 1959, Heredity 13: location: 3-58.8 (to the right of bx; to the left of
233-41). RK1. Ubx).
cytology: Shown to lie in doublet 18D1-2 by defi- origin: X ray induced. Arose simultaneously with
ciency analysis (J. I. Valencia). pbx.
discoverer: Bacon, 49h.
discoverer: Nolte, 1952. references: Lewis, 1954, DIS 28: 76.
references: 1954, DIS 28: 77. 1954, Proc. Intern. Congr. Genet., 9th. 1: 100-5.
phenotype: Visually resembles car, but contains 1955, Am. Naturalist 89: 73-89.
only one-half the amount of red pigment of car, 16 1963, Am. Zoologist 3: 35—56 (fig.).
percent of wild-type red pigment, and 48 percent phenotype: Cbx/+ and Cbx/Cbx virtually indistin-
of wild-type brown pigment (1959, Heredity 13: guishable and have small wings. The posterior
233-41). RK1. portion of the mesothorax transformed into a struc-
CQr26-48 ture that resembles posterior portion of meta-
origin: Induced by mustard gas. thorax. Wings have veins LI, L2, and L3 only. In
discoverer: Sobels and Jansen, 571. extreme cases, entire mesothorax resembles meta-
references: Sobels, 1958, DIS 32: 84. thorax so that wings resemble halteres. Meta-
phenotype: Eye color darker than car and tends to thorax is wild type. Cbx acts as dominant sup-
resemble pn more than car. RK1. pressor of homozygous pbx and partial suppressor
cardinal: see ccf of bx and bxd. Interactions with other bx pseudo-
carmine: see cm alleles described by Lewis (1955, 1963). RK2.
carnation: see car cytology: Salivary chromosomes normal. Located
Cat: see spaCat close to if not within the 89E1-2 doublet (Lewis).
*cb: club other information: A member of the bithorax pseudo-
location: 1-16.5. allelic complex (see bx).
origin: Spontaneous. *cc; chlorotic
discoverer: Morgan, 13e. location: 1-0.0 (0.1 to left of sc).
references: Morgan and Bridges, 1916, Carnegie origin: Spontaneous.
Inst. Wash. Publ. No. 237: 69 (fig.). discoverer: Mohr, 19jl8.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. references: 1923, Studia Mendeliana (Brunae):
Genet. 2: 78 (fig.), 266-87.
phenotype: Wings club-like in about half the flies. phenotype: Body color greenish yellow. Flies
Sternopleural bristles absent from all flies. RK3. small. Mortality 90 percent. RK3.
MUTATIONS 49

*CCY: chlorotic-yellowish discoverer: Glass, 39a28.


origin: Spontaneous. references: 1939, DIS 12: 47.
discoverer: Morgan, 21i. phenotype: Varies from almost complete fusion of
references: 1929, Carnegie Inst. Wash. Publ. No. veins L3 and L4 to wild type. Intermediate types
399: 190. have narrowing and closing of first posterior wing
phenotype: Body color pale yellow. Viability and cell with extra veins in region of anterior cross-
fertility low. Like cc. RK3. vein and deltas at ends of L3 and L4. Expression
ccw: concave wing is better above 25°C. Homozygous lethal. RK3.
locotion: 1-23.4., cytology: Placed in salivary chromosome region
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- 101E through 102B16, based on inclusion of Ce^
phenylalanine (CB. 3025). within Dt(4)M = Df(4)101E-F;102B6-17.
discoverer: Fahmy, 1953.
references: 1958, DIS 32: 68. origin: Spontaneous.
phenotype: Wings shorter and narrower, with L3 and discoverer: Green.
L4 shifted toward each other; occasionally trun- references: 1952, DIS 26: 63.
cated. Wing membrane depressed in center into phenotype: Ocelli reduced or absent; ocellar and
slight concavity, giving slight scooped effect. scutellar bristles absent; wing veins L3 and L4
Not easily classified. RK3. converge, giving wing phenotype much like fa,
other information: One allele induced by CB. 3025. although wing phenotype variable. Homozygous
cd: cardinal lethal; lethality occurs during embryonic period
location: 3-75.7. (Hochman). RK3.
origin: Spontaneous. other information: Allelism based on phenotype and
discoverer: Johnson, 19k24. lethal interaction with Df(4)M (Hochman).
references: Bridges and Morgan, 1923, Carnegie *Ce3
Inst. Wash. Publ. No. 327: 217 (fig.). origin: X ray induced.
phenotype: Eye color yellowish vermilion, discoverer: Green, 59cll.
changing toward wild type with age. Brown pig- references: 1959, DIS 33: 94.
ment 15 percent of normal (Nolte, 1954, J. Genet. phenotype: Identical to Ce2. RK3.
52: 111—26). Ocelli white, showing no effect of *cf; cleft
age. Eye color autonomous in transplant of larval location: 1-65.6.
optic disk into wild type, ca, en, st, or v larval origin: Spontaneous.
host (Beadle and Ephrussi, 1936, Genetics 21: discoverer: Bridges, 14J28.
230). Larval Malpighian tubes bright yellow; not references: Morgan, Bridges and Sturtevant, 1925,
distinguishable from wild type. RK2. Bibliog. Genet. 2: 55 (fig.).
*cc/3 phenotype: Wings smaller and somewhat spread.
origin: Spontaneous. L3 split just beyond first crossvein; extra cross-
discoverer: Ives, 32c30. veins and branches. Gap in L4 beyond second
references: Plough and Ives, 1934, DIS 1: 34. crossvein. Males sterile.. Viability good. RK2.
1935, Genetics 20: 42-69. *Cf: Confluent
phenotype: Like cd. RK2. location: 2- (not located).
cd63 origin: Spontaneous.
origin: Spontaneous. discoverer: Bridges, 14i23.
discoverer: Clancy, 63a. references: Bridges, 1916, Genetics 1: 151.
references: 1964, DIS 39: 65. Bridges and Morgan, 1919, Carnegie Inst. Wash.
phenotype: Like cd. RK2. Publ. No. 278: 255.
*Cd: Coildex phenotype: Veins thickened and knotted, especially
location: 2-54.6 (0.1 unit to the right of pr). L2 opposite anterior crossvein and at costa. Wing
origin: X ray induced. smaller. Low fertility in heterozygote; good
discoverer: Bateman, 1954. viability. Homozygous lethal. RK2.
synonym: Coiled. other information: May have been a Plexate.
references: 1955, DIS 29: 69.
phenotype: Similar to Cy, but wing curvature more Cf-3: see Die(-3
extreme; wings opaque and greyish. Anterior eg: comb gap
margin of wing invaginated at point where LI location: 2-71.1.
meets wing margin. When expression is weakest it origin: Spontaneous.
appears only as a slight wave in the wing margin. discoverer: Bridges, 25kl6.
In 10—15 percent of the flies, wings also curve phenotype: Sex combs of male extremely large.
downward over flanks before curling upward. In y; Some distortion and shortening of legs. Wings
Cd flies, curvature reduced to a shallow spoon. show gap in vein L4 between posterior crossvein
Cd epistatic to Cy. Homozygous lethal. RK2. and margin. Wings slightly curved. Effects result
*Ce: Cell from a combination of overgrowth and irregular
location: 4- (not located). folding of imaginal rudiments during the pupal
50 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

period. Strong exaggeration in compound hotnozy- phenotype: Wings short, papery, and downcurved
gotes with genes such as d, fj, ds, and ss fl . with short, broad alulae. Males sterile. Homozy-
Double heterozygote for eg and ci often shows gap gotes short lived and balanced stock
in L4 (Waddington, 1952, J. Genet. 51: 243-58). en che bw sp/In(2L)Cy + In(2R)Cy,
Double heterozygote en cg/++has slight degree of al% Cy en2 L4 sp? has a generation time 30 per-
L4 interruption and thinning at low temperature. cent longer than normal. RK3.
Triple heterozygote en cg/++; ci/+ has L4 inter- cfte*
ruption in half the flies (House, 1961, Genetics 46: origin: Ultraviolet induced.
871). ciw interacts strongly with eg. cg/+; ciw/+ discoverer: Meyer and Edmondson, 481.
resembles ciw/ciw (House, 1953, Genetics, 38: references: 1951, DIS 25: 71.
669—70). Females sterile. Oogenesis highly irreg- phenotype: Similar to cfte in wing characteristics
ular (Beatty, 1949, Proc. Roy. Soc. Edingurgh and male sterility. RK3.
B 63: 249-70). RK2.

eft: chubby
Left: wild-type larva. Right: chubby larva.
From Dobzhansky and Duncan, 1933, Arch. Entwicks-
lungmech. Organ. 130: 109-30.
*cfte3
origin: X ray induced.
eg; comb gap discoverer: Meyer, 60g.
From Bridges and Brehme, 1944, Carnegie Inst. Wash. references: 1963, DIS 37: 50.
Publ. No. 552: 40. phenotype: Wings curved, spread, and short. Less
extreme than che. che3/che sterile in both sexes.
eft: chubby che3/che3 not tested for sterility because fs(2)B
location: 2-72.5. and ms(2)2 on same chromosome. RK2.
origin: Spontaneous. chilblained-b: see cft-£>
discoverer: Bridges, 17J26. eft/: chaetelle
references: Morgan, Bridges, and Sturtevant, 1925, location: 2-60.8.
Bibliog. Genet. 2: 222. discoverer: Bridges, 33a4.
phenotype: Adults, pupae, and larvae thickset and references: Beatty, 1949, Proc. Roy. Soc.
short. Difficult to distinguish from wild type. Edinburgh B 63: 249-70.
Chubby larvae shorter than wild type at hatching phenotype: Bristles very small. Wing venation
[Dobzhansky and Duncan, 1933, Arch. Entwick- slightly plexus-like; exaggerates px when com-
lungsmech. Organ. 130: 109-30 (fig.)]. RK3. bined with it. Body size small. Rotated genitalia
*ch-b: chilblained-b in many males. Blunt-tipped abdomen. Females
location: 1-23.8. infertile, but ovary and oocytes appear normal.
discoverer: Moriwaki, 39e22. RK2.
references: 1939, DIS 12: 50. cft/orof/c: see cc
phenotype: Tarsi conglutinated. RK3. cfto: chocolate
cftofe//e: see eft/ location: 1-5.4 (left of ec).
*che: cherub origin: X ray induced,
location: 2-62.0. discoverer: Weigle, 1955.
origin: Ultraviolet induced. references: Sturtevant, 1955, DIS 29: 75.
discoverer: Meyer, 48g. phenotype: Eye color brown with whitish highlights.
references: Meyer and Edmondson, 1951, DIS 25: 71. Paler than se, less purplish than pa. Malpighian
MUTATIONS 51

tubes of larvae and adults contain brown pigment or absent. Other gaps and scattered branch veins
like red. Larvae easily distinguished from wild in region of crossveins. At 19°C nearly all flies
type. Brown pigment of Malpighian tubes absent have a mutant phenotype; at 25°C there is slight
when cho is combined with v, en, or sr mutations, overlap with wild type; at 30°C virtually all flies
which prevent formation of brown eye pigment. are wild type. Dosage effect such that ci/0
Eye color of cho v is yellowish, but cho g cannot haplo-4's are more extreme than ci/ci diplo-4's,
be distinguished from g. Separability, viability, which are more extreme than ci/ci/ci triplo-4's.
and fertility excellent. RK1. For interactions of ci with en, H, ve and eg, see
other information: Not included in Df(l)N8 = House, 1953, Genetics 38: 199-215, 309-27;
Df(l)3B4-Cl;3D6-El. 1955, Anat. Record 122: 471; 1959, Genetics, 44:
516; 1961, Genetics, 46: 871. Expression of ci
origin: Spontaneous. sensitive to genetic background; selection pos-
discoverer: Green, 1955. sible for more and less extreme phenotypes (House
references: Sturtevant, 1955, DIS 29: 75. and Yeatts, 1962, Genetics 47: 960). Phenotypic
phenotype: Dark brownish eye; indistinguishable effect visible in prepupa by absence of the lower
from cho. RK1. longitudinal vein. RK1 at 19°C and higher rank
chocolate: see cho with higher temperatures.
*chr: chrome cytology: Placed in salivary chromosome region
location: 1- (not located). 101F2-102A5, on the basis of its inclusion in
origin: Spontaneous. Df(4)M63a = Df(4)101F2-102Al;102A2-5.
discoverer: Bridges, 13115. other information: The expression of ci + can be
references: Morgan and Bridges, 1916, Carnegie altered in direction of ci by certain chromosome
Inst. Wash. Publ. No. 237: 74. rearrangements that have one break in vicinity of
phenotype: Body color brownish yellow or tan. ci locus. Rearranged fourth chromosomes carrying
Abdominal bands clear yellow. RK3. a mutant allele of ci, R(ci), may also show altered
other information: Probably a tan allele. expression of gene (Stern and Kodani, 1955). R(ci)
chubby: see c/i and R(ci+) terminology not retained here; interac-
chy: chunky tion with ci included in descriptions of aberrations
location: 2- (between 8 and 28). involving chromosome 4.
origin: Spontaneous. ci3 6l
discoverer: Bridges, 38blO. origin: Spontaneous.
phenotype: Body short and heavy set. Wings discoverer: Curry, 361.
shorter than wild type. Difficult to classify. RK3. phenotype: Less extreme than ci. Ranges from
appearance of a plexus in L4 between crossveins
to gaps in L4 and L5 posterior to crossveins.
RK3.

c/ + 2; cubitus interruptus-wild-type isoallele


origin: On fourth chromosome carrying ey2.
discoverer: Stern and Schaeffer, 1943.
references: 1943, Proc. Natl. Acad. Sci. U.S. 29:
361-67.
ch cubitus interruptus phenotype: Homozygote wild type at 14° and 26°C.
Wings showing from no interruption (extreme left) to ci+2/Df(4JM wild type at 26°C; shows some
complete absence (extreme right) of the cubital vein. thinning and interruption of L4 at 14°C. ci+2/ci
From Stern and Kodani, 1955, Genetics 40: 343-73. wild type at 26°C; at 14°C fewer flies show
thinning or interruption of L4 than ci + C/ci.
ci: cubitus interruptus ci*2/ciW shows significantly greater amount of
location: 4-0 (most proximal mutant in 4). thinning and interruption of L4 than c / + c / c i w \
origin: Spontaneous. RK3.
discoverer: Tiniakov and Terentieva, 30b. c/ + 3
references: Terentieva, 1931, Zh. Eksperim. Biol. discoverer: Stern and Schaeffer, 1943.
7: 187-90 (fig.). referenees: 1943, Proc. Natl. Acad. Sci. U.S. 29:
Tiniakov and Terentieva, 1933, Genetics 18: 361-67.
117-20 (fig.). phenotype: Homozygote wild type at 26° and shows
Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20. some thinning of L4 at 14°C. About half of
Sturtevant, 1951, Proc. Natl. Acad. Sci. U.S. 37: ci+3/ci heterozygotes are not wild type at 25° and
405-7. about three-fourths are not at 14°C. Only a few
Stern and Kodani, 1955, Genetics 40: 343-73 (fig.). ci*3/ciw individuals overlap wild type. RK3.
phenotype: Vein L4 shows one or more gaps both C/+5
distal and proximal to posterior crossvein, gen- origin: A male of the Cockaponsett wild stock.
erally nonterminal. Anterior crossvein shortened discoverer: Hochman, 551.
52 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: 1961, Evolution 15: 239-46. synonym: It: Interntptus.


phenotype: Wild type at 25°C; at 17°C a small frac- phenotype: Homozygote is extreme ci type. Wings
tion of flies display wing vein abnormalities; how- sometimes almost twice normal width, arclike, and
ever, not involving L4. Over 80 percent of ci+5/ci virtually lack veins. Often present is a well-
flies show gaps in L4 distal to posterior cross- organized pattern of venation in which the pos-
vein. RK3. terior crossvein flows smoothly into L5. Legs
C/+C lumpy; sex combs larger than normal; antennae
origin: Canton-S wild type. enlarged; eyes smaller; and extra bristles present.
discoverer: Stern and Schaffer, 1943. Heterozygote shows gap in L4 in 80 percent of
references: 1943, Proc. Natl. Acad. Sci. U.S. 29: flies. ciw enhanced by H, en, and Cy (House,
361-67. 1953, Genetics 38: 669-70; 1959, Genetics 44:
phenotype: When homozygous, wild type at 14°, 18°, 516), Temperature effect described by House
and 25°C. ci+c/Df(4)M wild type at 26°C; very (1955, Genetics 40: 576). RK2.
few flies show thinning or interruption of L4 at cinnabar: see en
14°C. ci^C/ci wild type at 26°; some flies show ck: crinkled
thinning or interruption at 14°C. ci+c/ciw causes location: 2-53.
significant thinning or interruption of L4 at 26 C. origin: Spontaneous.
RK3. discoverer: Bridges, 30c30.
ciD: cubitus interruptus-Dorninant phenotype: Wings flimsy, crinkled, or wavy.
origin: X ray induced. Irregular stubby or wavy bristle effect. Viability
discoverer: Ruch, 32al8. poor. RK3.
references: Bridges, 1935, Biol. Zh. (Moscow), cl: clot
4: 401-20. location: 2-16.5.
phenotype: Wings show interruptions of L4 in two origin: Spontaneous.
places; proximal to and distal to, anterior cross- discoverer: Bridges, 27a3.
vein. L5 also shows distal interruption. L3 and phenotype: Eye color dark maroon to sepia-like
L5 thick. Considerable plexus effect and knotting with age; less extreme than sepia. Sixty percent
of veins. Wings broader and warped or concave more red pigment than wild type (Nolte, 1954,
upward, regularly extended and bent backward. J. Genet. 52: 127—39). Eye color autonomous
Alula fused with and in same plane as blade of when larval optic disk is transplanted into wild-
wing. Black dried blood from axillary spiracle. type host (Beadle and Ephrussi, 1936, Genetics
Slight scalloping of inner wing margin, with hairs 21: 230). Larval Malpighian tubes pale yellow,
and tufts. Direction and extent of temperature distinguishable from wild type (Brehme and
effects depends on genetic background (Scharloo). Demerec, 1942, Growth 6: 351-56). RK1.
In general, no overlapping wild type. H/+ inhibits cytology: Placed in salivary chromosome region
scalloping of ciD but greatly enhances L4 inter- 25E1 to 26C1 (E. H. Grell).
ruption (House, 1959, Genetics 44: 516). Fully c/2
dominant in triplo-4's (Sturtevant, 1936, Genetics discoverer: Terry, 1928.
21: 448). Homozygous lethal. Lethal acts in phenotype: Eye color like cl but darker. Larval
embryonic stage (Hochman). RK1. Malpighian tubes pale yellow (Brehme and
cytology: Salivary study by Bridges revealed no Demerec, 1942, Growth 6: 351-56). RK1.
chromosomal aberration. claref: see ca
other information: Not allelic, at least with respect *c/c/: cloudy
to its lethality, since ciD/Df(4)M63a survives, location: 2-96 to -101.
whereas ci/Dt(4)M63a is mutant (Hochman, 1965, origin: Gamma ray induced.
DIS 40: 60). discoverer: Wallbrunn, 61 j6.
c / 0 - 6 ; cubitus interruptus-Dominant of G/oor references: 1964, DIS 39: 59.
origin: Obtained by recombination between chromo- phenotype: Wings opaque from fluid between upper
somes with ci® and spaP°K and lower membranes; occasionally fluid forms
discoverer: Gloor. small blisters. Males sterile, females highly
references: Scharloo, 1963, DIS 38: 32. infertile. RK2.
phenotype: Less extreme than ciD. Interruption of cleft: see cf
L5 is infrequent. Usually a terminal interruption elf; see wtwdf
of L4 distal to second crossvein and a gap prox- *Cli: Clipped wings
imal to first crossvein. Wings neither spread nor location: 1- (to the left of f).
warped. No black dots present in axillary discoverer: Agol.
spiracles, but overlaps wild type at lower temper- references: 1936, DIS 5: 7.
atures. Good expressivity at 25°C (Scharloo). phenotype: Dominant wing mutant (no description
RK1. given). Viable in male and homozygous female.
ci*; cuhitus interrvptvs of Wallace RK3.
origin: Spontaneous. clip wing: see
discoverer: E. M. Wallace, 36d20. clipped: see cp
MUTATIONS 53

Clipped wings: see Cli More precisely located by Hannah-Alava in 6E,


Clipt: see Cpt probably in or near 6E6.
elm: clumpy marginals cm: see emp
location: 1-32.6.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- origin: Induced by mustard gas.
phenylalanine (CB. 3025). discoverer: Sobels, 571.
discoverer: Fahmy, 1953. references: 1958, DIS 32: 84.
references: 1958, DIS 32: 68. phenotype: Eye color more translucent and ruby-like
phenotype: Irregularly bent marginal hairs, than cm. RK1.
especially on posterior border of wings. Bristles cm ^
stiff and frequently bent or split. Viability and origin: X ray induced in R(l)2.
fertility of males good. Homozygous females discoverer: Muller, Valencia, and Valencia, 1946-53.
reduced in viability and fertility. RK2. references: Valencia, 1966, DIS 41: 58.
other information: One allele each induced by cytology: Associated with Dt(l)cmR8BH4 =Df(l)6E
CB. 1506 and CB. 3007. (J. I. Valencia).
clot: see c/
cloudy: see eld
cloven thorax: see c/v
club: see cb
clubfoot: see cbf
cluboid: see cbd
clumpy marginals: see elm

*c/v-7; cloven thorax no. 7


location: 1-0.0. Cm: Crimp
origin: X ray induced. Edith M. Wallace, unpublished.
discoverer: Muller, 19h.
references: 1935, DIS 3: 29. *Cm: Crimp
phenotype: Thorax often has long cleft; partially location: 3-43.5.
dominant. Semilethal at low temperature, viable origin: Spontaneous.
at high one. RK3. discoverer: Bridges, 28a28.
*clv-2 phenotype: Heterozygote has crimped wings ruffled
location: 1-42.0. on rear edge. Classification good in first 4 days'
origin: X ray induced. hatch, then Cm overlaps wild type progressively.
discoverer: Muller, 26111. Better at 25° than at 19°C. Homozygous lethal.
references: 1935, DIS 3: 29. RK2 as lethal; RK3 as dominant.
phenotype: Thorax has logitudinal cleft, sometimes
half thorax. One wing often reduced or like vg.
Partially dominant. Semilethal. RK3.
*cly.2S2b
origin: X ray induced.
discoverer: Bateman, 52b.
references: 1953, DIS 27: 55.
phenotype: Like clv-2. Some flies have no dorsal
thorax at all. Viability 30 percent. Penetrance 50
percent. The apparently wild-type males are
fertile; abnormal ones sterile. Completely
recessive. RK3.
other information: Allelism with clv-2 not tested.
cm: carmine
location: 1-18.9.
origin: Spontaneous.
discoverer: Mohr, 27d27. crop: crumpled
references: 1927, Z. Induktive Abstammungs- Edith M. Wallace, unpublished.
Vererbungslehre 45: 403—5.
phenotype: Eye color translucent dark ruby. With emp: crumpled
st, eye color deep orange; with brown, slightly location: 3-93.
lighter than bw alone. Larval Malpighian tubes origin: Spontaneous.
very pale yellow. RK1. discoverer: Bridges, 22d2.
cytology: Locus lies between 6A3-4 and 6F10-11 synonym: cm.
(Demerec, Kaufmann, Fano, Sutton, and Sansome, references: Bridges and Morgan, 1923, Carnegie
1942, Carnegie Inst. Wash. Year Book 41: 191). Inst. Wash, Publ. No. 327: 247.
54 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

Morgan, Bridges, and Sturtevant, 1925, Bibliog. cytology: Associated with a tandem duplication,
Genet. 2: 223. Dp(l;l)Co = Dp(l;l)3C4-5;3D6-El (Schultz, 1941,
phenotype: Wings about two-thirds normal size and DIS 14: 54-55). Result of duplication of 3C7,
greatly crumpled or blistered. Marginal hairs deficiency for which gives Notch (Morgan, Schultz,
irregularly clumped. Legs irregularly shortened and Curry, 1941, Carnegie Inst. Wash. Year Book
and gnarled. Bristles somewhat short and thick. 40: 283).
Posterior scutellars slightly divergent. Branches other information: Reversion to wild type occurs in
of aristae bent anteriorly near middle, with apices Co/Co by unequal crossing over. A member of the
parallel to main axes of aristae. Viability and Notch pseudoallelic complex.
fertility may be low. RK3.
en: cinnabar
location: 2-57.5.
origin: Spontaneous.
discoverer: Clausen, 2018.
references: 1924, J. Exptl. Zool. 38: 423-36.
phenotype: Eye color bright red, like v or st.
Ocelli colorless. Eye color darkens with age, but
ocelli remain colorless. Larval Malpighian tubes Co: Confluens
pale yellow (Beadle, 1937, Genetics 22: 587— Edith M. Wallace, unpublished.
611). Nonautonomous in development of pigment
of transplanted eye disks (Beadle and Ephrussi, Co-3A: see l(2)S3a
1936, Genetics 21: 230), en blocks conversion of Co-7: see 1(2)S7
kynurenine to 3-hydroxykynurenine, which has coal: see cal
been identified as the en "*" hormone (Butenandt, *coc: collapsed ocelli
Weidel, and Schlossberger, 1949, Z. Naturforsch. location: 1-61.5.
4b: 242-44). RK1. origin: Induced by D-l:6-dimethanesulfonyl mannitol
cytology: Proximal to 44C, based on its inclusion in (CB. 2511).
Dp(2;3)P32 = Dp(2;3)41A;42D-E;44C-D;89D7-El discoverer: Fahmy, 1960.
(E. B. Lewis). references: 1964, DIS 39: 58.
cn2 phenotype: Ocelli small and flat; deflated owing to
origin: Spontaneous in In(2R)Cy. lack of eye fluid. Anterior ocellar hairs frequently
discoverer: L. Ward, 1921. missing. Other slight alterations in body size and
references: 1923, Genetics 8: 276—300. wing shape. RK3.
phenotype: Eye color slightly brighter than normal cytology: Placed in salivary region 18A4 through
and ocelli pale. cn2/cn bright scarlet like cn/cn 18B8 on the basis of its inclusion within the defi-
and has colorless ocelli but darkens more rapidly ciency carrying the left end of In(l)y4 = In(l)lA8-
with age. Malpighian tubes of en2/en pale yellow Bl;18A3-4 and the right end of In(l)sc^ =In(l)lB2-
(Brehme and Demerec, 1942, Growth 6: 351—56). 3;18B8-9 (Norton and Valencia, 1965, DIS 40: 40).
RK2.
en*: cinnabar-sterile Co/: Coiled
origin: Spontaneous. location: 2-48.7.
discoverer: Ives, 40el8. origin: X ray induced.
phenotype: Eye color like en, but females sterile. discoverer: Carlson, 57g.
RK2, references: 1960, DIS 34: 48.
other information: Possibility that sterility factor phenotype: Coi/+ has curled wings like Cy/+ or j.
separable from en not completely eliminated. Coi/Coi viable, with strongly curled wings similar
Found in chromosome carrying In(2L)t = to Cy/+; JD/+. Coi/j shows no interaction. Excel-
In(2L)22D3-El;34A8-9. lent viability and fertility. RK1.
Co: Confluens cytology: Not included in Df(2L)64j = Df(2L)34E5-
location: 1-3.0. F1;35C3-D1 (E. H. Grell).
origin: Recovered among progeny of cold-treated Coiled: see Cd
fly. Coiledex: see Co'
discoverer: Gottschewski 34c. collapsed ocelli: see coc
references: 1935, DIS 4: 7, 14, 16. *co/n: compressed
1937, Z. Induktive Abstammungs- Vererbungslehre location: 3-48.5.
73: 131-42. origin: Spontaneous.
1937, DIS 8: 12. discoverer: Bridges, 18k27.
phenotype: Veins irregularly thickened, especially references: Bridges and Morgan, 1923, Carnegie
toward tips, which are usually deltas and fused Inst. Wash. Publ. No. 327: 193.
broadly to marginal vein. Stronger expression in phenotype: Head flattened ventraliy. Eyes small,
males than in females. Co/N8 wild type except for displaced. Vibrissae tufted. Aristae crumpled.
slightly thicker L3 vein. Co/Ax like Ax/+. RK1A. Humeral patches elevated. Wings droopy. Poor
viability and fertility. RK3.
MUTATIONS 55

*com~d: compres sed-dilapida tor corr; corrugated wing


location: 3-68.5. location: 2-36.
origin: Spontaneous. origin: Spontaneous.
discoverer: Bridges, 19c8. discoverer: Mayeda, 61g.
phenotype: Flies small, pale, weak, with defective references: 1963, DIS 38: 31.
legs and wings. RK3. phenotype: Wings wrinkled and wavy, reduced to
comb gap: see eg three-fourths normal size. Whole wing corrugated
compressed: see com at 20°C, only posterior third at 25°. Good classi-
*con: condensed fication. RK2.
location: 1-27.1.
origin: Spontaneous. Corroded eye: see Cor
discoverer: Bridges, 36dll. corrugated wing: see corr
references: 1937, DIS 7: 6. costakink: see csk
phenotype: Thorax and abdomen shortened; abdomen cp: clipped
dilated, exposing ventral skin to side view. Eyes location: 3-45.3.
slightly roughened, occasionally kidney shaped discoverer: Mainx, 34g.
and somewhat dark. Wings short, bluntly rounded, references: 1936, Z. Induktive Abstammungs-
with crossveins closer together than normal. Vererbungslehre 71: 303-4 (fig.).
Bristles shortened and somewhat fine at 19°C, Pollitzer, 1937, DIS 8: 91.
stubby at 25°C. Postscutellars semierect and phenotype: Wing margins snipped, most often along
crossed; posterior verticals shortened or missing. marginal vein. At 19°C character slighter but
Male entirely sterile. Viability 50 percent wild completely penetrant. RK1.
type. RK2.
cytology: Salivary chromosome studies (Demerec,
Kaufmann, Fano, Sutton, and Sansome, 1942,
Carnegie Inst. Wash. Year Book 41: 191) show
locus to lie between 7C4-5 and 8C1-2. Further
restricted to 7E1 through 8C2, on the basis of its
genetic location to the right of oc which is ex-
cluded from Di(l)sn = Df(l)7B2-3;7D22-El (Hinton
and Welshons, 1955, DIS 29: 125-26).
concave wing: see ccw cp; clipped
condensed: see con Edith M. Wallace, unpublished.
Confluens: see Co
Confluent: see Cf *cpl: cupola
Confluent-3: see Dlcf-3 location: 1-0.0 (no crossing over with sc in 584
contorted: see ctt males).
Contrabithorax: see Cbx origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
convex wing: see cvw phenylalanine (CB. 3025).
cop; copper discoverer: Fahmy, 1953.
location: 1-43.3. references: 1959, DIS 33: 83-84.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- phenotype: Small, inviable fly. Wings shorter and
phenylalanine (CB. 3026). curved to form canopy over abdomen with tips
discoverer: Fahmy, 1953. converging toward mid-dorsal line. Head and eyes
references: 1958, DIS 32: 68. slightly deformed. Abdominal tergites abnormal;
phenotype: Brownish red eye color. Best classifi- from irregular pigmentation to absence or gross
cation in newly emerged flies. Occasionally wings deformation of the sixth and seventh tergites.
show cutaway inner margins. Excellent viability Males sterile. RK3.
and fertility in both sexes. RK2. *Cpt: Clipt
other information: Two alleles induced by CB. 3025. location: 2-43.7.
Cor: Corroded eye origin: Spontaneous.
location: 3- (not located), discoverer: Sturtevant, 26bl8,
origin: X ray induced, phenotype: Bristles short, like those of Sb. Homo-
discoverer: Muller. zygous lethal. Male sterile. RK1.
references: 1946, DIS 20: 66. *cpw: canopy wing
phenotype: Cor/+ shows slight irregular flecking of location: 1-2.5.
eye. In combination with v, expression enhanced, origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
producing patchy diminution in color, especially phenylalanine (CB. 3025).
near posterior margin of eye, giving impression discoverer: Fahmy, 1953.
that color was washed or eaten away, especially references: 1958, DIS 32: 69.
from deeper layers; regions of surface often phenotype: Wings short and very broad; longitudinal
blackened. Homozygote not described. RK2. veins frequently do not reach wing margin and
56 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

often diverge. Eyes large and slightly rough- Cream: see Cr


Head bristles reduced in number (ocellars most Cream: see
frequently affected). Thorax broad, one or more cream underscored: see cru
bristles occasionally absent; hairs more widely creased: see cs
separated, with noticeable hairless areas. Males creeper: see rk4
sterile. Viability 40 percent wild type. RK3. Crimp: see Cm
eg; see rk4 crinkled: see ck
*cr: crisp *crip: cripple
location: 1- (not located). location: 2- (between pr and en).
discoverer: Agol. discoverer: Komai, 1924.
references: 1936, DIS 5: 7. references: 1926, Genetics 11: 280-93.
phenotype: Bristles like forked. RK2. 1927, Mem. Coll. Sci. Univ. Kyoto, Ser. B 2: 211 —
other information: Not an allele of I or sn. 57.
*Cr-2: Cream in chromosome 2 phenotype: Middle and hind legs twisted and
location: 2- (not located). shortened. Thirty percent penetrance. RK3.
origin; Spontaneous. crisp: see cr
discoverer: Bridges, 13il5. *crk: crooked setae
references: 1919, J. Exptl. Zool. 28: 337-84. location: 1-60.1.
Bridges and Morgan, 1919, Carnegie Inst. Wash. origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
Publ. No. 278: 239 (fig.). phenylalanine (CB. 3026).
phenotype: Specific dilutor of w0, we; Cr-2/Cr-2 discoverer: Fahmy, 1953.
has a pale cream eye color. we; Cr-2/+ has eye references: 1958, DIS 32: 69.
color between eosin and cream. RK3. phenotype: Bristles thin and slightly shortened;
*cr-3: cream in chromosome 3 occasional missing scutellar. Acrostichals de-
location: 3-36.5. ranged. Abdominal hairs of female frequently
origin: Spontaneous, missing; tergites occasionally abnormal. Classifi-
discoverer: E. M. Wallace, 14b27. cation difficult. Viability and fertility good. RK3.
references: Bridges, 1919, J. Exptl. Zool. 28: other information: One allele induced by CB. 3025.
337-84. *crm: cramped
Bridges and Morgan, 1923, Carnegie Inst. Wash. location: 1-1.4 (based on crm^).
Publ. No. 327: 112 (fig.). origin: Induced by P32.
phenotype: Homozygote has slightly diluted eye discoverer: Bateman.
color. Eye color of we; cr-3 cream. Larval Mal- synonym: stap: stubarista from P32.
pighian tubes of w°; cr-3 white, those of cr-3 references: 1951, DIS 25: 78.
bright yellow (Brehme and Demerec, 1942, Growth 1953, DIS 27: 55.
6: 351-56). RK3. phenotype: Antennae stumpy with shrunken, warped
*cr-a: cream-a aristae, usually lying back on head. Initially
location: Autosomal, not located. showed narrowed and scalloped wings and eyes
origin: Spontaneous. shaped like inverted pears, but these effects vari-
discoverer: Bridges, 13gl5. able. Viability low. Sterile. RK3.
references: 1916, Genetics 1: 147.
1919, J. Exptl. Zool. 28: 337-84. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
phenotype: Strong specific dilutor of we. RK3. phenylalanine (CB. 3025).
*cr-b discoverer: Fahmy, 1953.
location: 2-24. synonym: cramped-like.
origin: Spontaneous. references: 1958, DIS 32: 69.
discoverer: Bridges, 14clO. phenotype: Antennae stumpy and short, with
references: 1916, Genetics 1: 149. reduced, abnormal aristae. Wings frequently
1919, J. Exptl. Zool. 28: 337-84. abnormally held, with cuts on inner margins. Eyes
Bridges and Morgan, 1919, Carnegie Inst. Wash. pear shaped. Small extra sex combs on second
Publ. No. 278: 245 (fig.). tarsal segment. Not easily classified. Viability
phenotype: Specific dilutor of w e . RK3. and fertility slightly reduced in males, greatly
*cr-c reduced in females. RK3.
location: 2- (near S). other information: Allelism to crm inferred from po-
origin; Spontaneous. sition of ctni at 1:14 and phenotype. Seven other
discoverer: Bridges, 16gl3. alleles: 1 each induced by CB. 1540, CB 1592,
phenotype: Weak specific dilutor of we. RK3. CB. 3007, CB. 3025, CB. 3034, and 2 induced by
cramped: see crm CB. 1506.
cramped-like: see crm2 cro: see ptg3
CRB: see T(1;4)A1 crooked: see fwc
etc: see l('2)crc crooked setae: see crk
cream: see cr crossover suppressor: see c( )
MUTATIONS 57

crossveinless; see cv *csk: costakink


crowri: see ptg3 location: 1-33.0 (no crossovers with v in 526
crs: cru sterile males).
location: 2- (between px and bw). origin: Induced by triethylenemelamine (CB. 1246).
discoverer: Muller. discoverer: Fahmy, 1953.
references: 1951, DIS 25: 119. references: 1958, DIS 32: 69.
1955, DIS 29: 146. phenotype: Eyes smaller. Wings slightly reduced in
phenotype: Male sterile. RK2. size and abnormally held; costal vein frequently
cytology: Located between 58E3 and 59A2 on basis kinked near L2. Not fully penetrant. Male
of sterility in combination with Di(2R)P + viability and fertility good, but female viability
Dp(2;Y)bw+ = Di(2R)58E3-Fl;60D14-E2 + and fertility reduced to about 50 percent wild
Dp(2;Y)YL;58Fl-59A2;60D14-E2 (Muller, 1955). type. RK3.
other information: Male sterility formerly associated other information: One X-ray-induced allele.
with but separable from cru. cf: cuf
crt: crumpled tips location: 1-20.0.
location: 1-40.3 (7.3 units from v, based on 3035 origin: Spontaneous.
flies). discoverer: Bridges, 15J12.
origin: Induced by triethylenemelamine (CB. 1246). references: Morgan, Bridges, and Sturtevant, 1925,
discoverer: Fahmy, 1952. Bibliog. Genet. 2: 35, 223 (fig.).
references: 1959, DIS 33: 84. phenotype: Wings cut to points and edges scalloped.
phenotype: Wing tips frequently shriveled, pleated, Eyes smaller and somewhat kidney shaped.
or crumpled, and often turned up or down. Wings Abdominal bands warped. Antennae often
vary from completely unexpanded to wild type. deformed. RK1.
Viability and fertility good in both sexes. RK2. cytology: Placed in salivary gland chromosome
other information: Twelve other alleles: 1 each bands 7B3-4 (Demerec, Kaufmann, Fano, Sutton,
induced by X rays, CB. 1246, CB. 1522, and and Sansome, 1942, Carnegie Inst. Wash. Year
CB. 3025; 2 induced by CB. 3034; 3 each induced Book 41: 191). Hannah-Alava agrees.
by CB. 1592 and CB. 3007.
cru: cream underscored
location: 2-52.5. origin: X ray induced.
origin: Spontaneous. discoverer: Hannah, 1947.
discoverer: Bridges, 20a5. references: 1949, Proc. Intern. Congr. Genet., 8th.
phenotype: Specific dilutor of we and P. Slight pp. 588-89.
dominant but used as a recessive. Originally phenotype: Lethal. Shows cut phenotype in combi-
thought to be male sterile, but this was caused by nation with viable ct alleles; lethal in combination
a factor in 2R, crs. Larval Malpighian tubes of with lethal ct alleles. RK2A.
we; cru colorless; those of +; cru bright yellow cytology: Associated with Dl(l)ct^a^ =
(Brehme and Demerec, 1942, Growth 6: 351—56). Df(l)7B3-6;7B6-7.
RK3. *cf2a3
crumpled: see crop origin: X ray induced.
crumpled tips: see erf discoverer: Hannah, 1947.
cru sterile: see crs references: 1949, Proc. Intern. Congr. Genet., 8th.
*cs; creased pp. 588-89.
location: 1-56. phenotype: Like ct**?. RK2A.
origin: X ray induced. cytology: Associated with D((l)ct2a3 =
discoverer: K. C. Atwood, 41i. Dt(l)7B2-3;7Cl-2.
references: 1942, DIS 16: 47. *ct2cl
phenotype: Wings logitudinally creased in first origin: X ray induced.
posterior cell from distal end of L3 virtually to discoverer: Hannah, 1947.
anterior crossvein. Fertility and viability good. references: 1949, Proc. Intern. Congr. Genet., 8th.
RK1. pp. 588-89.
CS53 phenotype: Like ct2a^. RK2.
origin: X ray induced. cytology: Salivary chromosomes normal.
discoverer: Krivshenko, 53k5. *ct*
references: 1956, DIS 30: 74. discoverer: Morgan, 17a22.
phenotype: Wing longitudinally creased. Lateral references: Morgan, Bridges, and Sturtevant, 1925,
edges of wings bent slightly downward distally. Bibliog. Genet. 2: 35 (fip.).
RK1. phenotype: Extreme ct allele. Wings short, dark,
other information: According to crossover data of and crumpled as well as cut and scalloped.
M. Aronson and description, this is an allele of Abdominal bands warped. Antennae flattened and
cs, which was lost before this mutant was embedded. Aristae concave forward. Eyes smaller
discovered. and kidney shaped. Vibrissae gone. More extreme
58 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

expression in females than in males; females have phenotype: Lethal. Expression in combination with
much poorer viability. 1 female:3 males in stock. ct6 suppressed by YL. RK2A.
RK3. cytology: Associated with Tp(l)ct6al =
* c f3o2 Tp(l)7B2-Cl;19;20.
origin: X ray induced.
discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89.
phenotype: Lethal in male and homozygous female,
as well as in combination with other lethal alleles
of ct. Extreme ct phenotype in combination with
ct6. Like ctn in combination with ctn, Phenotype
reportedly suppressed by addition of Y chromo-
some material. RK2A.
cytology: Associated with In(l)ct3a2 —
In(l)7B2-Cl; 19-20.

origin: X ray induced.


discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89.
phenotype: Lethal, but unlike other lethal ct
alleles tested by Hannah, fails to survive in
combination with Dp(l;3)sn13al. RK2A.
cytology: Associated with In(l)ct3bI =
In(l)3A4-Bl;7B2-5.
ctf>: cut-6
origin: X ray induced. From Morgan, Bridges, and Sturtevant, 1925, Bibliob.
discoverer: Hannah, 1947. Genet. 2: 35.
references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89. *C/7a7
phenotype: Like cf 2 °2. RK2A. origin: X ray induced.
cytology: Associated with Df(l)ct4bi = discoverer: Hannah, 1947.
Di(l)7B2-4;7C2-4. references: 1949, Proc. Intern. Congr. Genet., 8th.
*ct4d pp. 588-89.
origin: X ray induced. phenotype: Lethal. ct7al /Dp(l;3)sn13al males
discoverer: Hannah, 1947. show reduced viability and are probably sterile.
references: 1949, Proc. Intern. Congr. Genet., 8th. RK2A.
pp. 588-89. cytology: Associated with T(l;2)ct7ai = T(1;2)7B.
phenotype: Like ctl*2. R K 2 . * c f7o2
cytology: Salivary chromosomes normal. origin: X ray induced.
discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th.
origin: Spontaneous. pp. 588-89.
discoverer: Bridges, 2Oc20. phenotype: Like c*2a2. RK2A.
references: Morgan, Bridges, and Sturtevant, 1925, cytology: Associated with Df(l)ct?a2 =
Bibliog. Genet. 2: 35 (fig.). Di(l)7A5-Bl;7C4-9.
phenotype: Wing character uniform and reliable; *cf7b2
usually lacks the abdomen, antenna, arista, and origin: X ray induced.
eye effects of ct. Vibrissae gone or displaced discoverer: Hannah, 1947.
downward to bottom of eye. Developmental study references: 1949, Proc. Intern. Congr. Genet., 8th.
by Waddington Il939, Proc. Natl. Acad. Sci. U.S. pp. 588-89.
25: 299-308; 1940, J. Genet. 41: 75-139 (fig.)] phenotype: Like cf 2fl 2. RK2.
shows wing bud narrower than wild type as early cytology: Salivary chromosomes normal.
as just after eversion of wing in early pupa. RK1. * c f 7c7
other information: Allele most used as genetic origin: X ray induced.
marker. discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th.
origin: X ray induced, pp. 588-89.
discoverer: Hannah, 1947. phenotype: Like ct3*2. RK2A.
references: 1949, Proc. Intern. Congr. Genet., 8th. cytology: Associated with T(l;2)ct7cl -
pp. 588-89. T(1;2)7B2-3;8E2-3;25C.
MUTATIONS 59

*ct7c2
origin: X ray induced. origin: X ray induced.
discoverer: Hannah, 1947. discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th. references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89. pp. 588-89.
phenotype: Like ct2a2. RK2A. phenotype: Like ct**2. RK2A.
cytology: Associated with Df(l)ct?c2 = cytology: Associated with T(l;3)ct^a =
Df(l)6Fll-7Al;7B8-Cl. T(1;3)1B; 7B2-3;8E-F;84B.
*cf9bl * c f72a7
origin: X ray induced. origin: X ray induced.
discoverer: Hannah, 1947. discoverer: DeFrank, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th. references: Hannah, 1949, Proc. Intern. Congr.
pp. 588—89. Genet., 8th. pp. 588-89.
phenotype: Similar to ct6. RK1. phenotype: Lethal in hemizygote and in combina-
cytology: Salivary chromosomes normal, but tion with other lethal ct alleles. Expression vari-
staining of 7B1-2 and 7B5 lighter and darker than able in combination with viable ct alleles.
normal, respectively. ct12al /Dp(l;3)ani3al males show low viability
other information: Induced simultaneously with an and are sterile. RK3.
independent but closely linked recessive lethal cytology: Salivary chromosomes normal.
mutation. *C/J2o2
*cf9b2 origin: X ray induced.
origin: X ray induced. discoverer: DeFrank, 1947.
discoverer: Hannah, 1947. references: Hannah, 1949, Proc. Intern. Congr.
references: 1949, Proc. Intern. Congr. Genet., 8th. Genet., 8th. pp. 588-89.
pp. 588-89. phenotype: Like ct2a2 except that lethal not
phenotype: Semilethal but semilethality not shown covered by Dp(l;3)sn13al. RK2A.
to be at ct locus; males appear rarely; sterile with cytology: Associated with In(l)ct12a2 =
small yellowish tan bodies. Margin notched from ln(l)4E2-3;7B2-4.
the costal cell around wings to base of inner * c f72c7
margin. RK2A. origin: X ray induced.
cytology: Induced simultaneously with but appar- discoverer: Hannah, 1947.
ently independently of a complex inversion and references: 1949, Proc. Intern. Congr. Genet., 8th.
translocation between X and 3R. pp. 588-89.
phenotype: Lethal, In combination with viable ct
* c f70o7 alleles, some flies show abnormal venation.
origin: X ray induced. ctl2cl/Dp(l;3)snl3al males viable but sterile.
discoverer: DeFrank, 1947. RK2A.
references: Hannah, 1949, Proc. Intern. Congr. cytology: Associated with T(l;3)ctI2cl =
Genet., 8th. pp. 588—89. T(l ;3)7B2-3;7D2-6;85.
phenotype: Like ct2a2. RK2A.
cytology: Possibly associated with Dt(l)ct1Oal = origin: X ray induced.
Df(l)7B3-4;7B6-7. discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th.
origin: X ray induced. pp. 588-89.
discoverer: Hannah, 1947. phenotype: Like ct2*2. RK2A.
references: 1949, Proc. Intern. Congr. Genet., 8th. cytology: Possibly associated with Dt(l)ct12c2 —
pp. 588-89. Df(l)7B2-3;7B6-7.
phenotype: Like ct2a2'. RK2A.
cytology: Associated with Dl(l)ct1Obl = origin: Spontaneous.
D((1)6D8-E1;7B7-C1. discoverer: Bridges, 21f7.
* c f70c7 references: Morgan, Bridges, and Sturtevant, 1925,
origin: X ray induced. Bibliog. Genet. 25: 35 (fig.).
discoverer: DeFrank, 1947. phenotype: Like ct but females usually sterile.
references: Hannah, 1949, Proc. Intern. Congr. RK2.
Genet., 8th. pp. 588-89. Cf73o7

phenotype: Males have notched but unexpended origin: X ray induced.


wings; may have abnormal antennae and vibrissae. discoverer: Hannah, 1947.
Males show very low viability, usually dying in references: 1949, Proc. Intern. Congr. Genet., 8th.
larval stage. Surviving males sterile. Lethal in pp. 588-89.
combination with other lethal ct alleles; heterozy- phenotype: Like ct2®2. RK2A.
gous females made using Dp{l;3)&n13l*l. RK2. cytology: Associated with In(l)ct*3mi ~
cytology: Salivary chromosomes normal. In(l)7B2-3; 19-20.
60 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

cytology: Associated with Df(l)ct14cl =


origin: X ray induced. Df(l)7B3-4;7B6-9.
discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th. origin: X ray induced.
pp. 588-89. discoverer: Hannah, 1947.
phenotype: Similar to ct6. RK1. references: 1949, Proc. Intern. Congr. Genet., 8th.
cytology: 7B1-2 show abnormal staining and pp. 588-89.
ectopic pairing with heterochromatic regions. phenotype: Like ct2*2. RK2A.
cytology: Associated with Df(l)ct1Sbl =
origin: X ray induced. Df(l)7B2-4;7B6-7.
discoverer: Hannah, 1947.
references: 1949, Proc. Intern. Congr. Genet., 8th. ct15B4

pp. 588-89. origin: X ray induced in In(l)scslLsc8R+dl-49.


phenotype: Like ct^e?. RK2A. discoverer: Muller, Valencia, and Valencia, 1946-53
cytology: Associated with T(l;4)ctl3bl - references: Valencia, 1966, DIS41: 58.
T(l;4)lA;7B2-3;101A-D;102. phenotype: Male viable. RK2.
*cf36b
origin: X ray induced. origin: Spontaneous.
discoverer: Hannah, 1947. discoverer: Stalker, 36b28.
references: 1949, Proc. Intern. Congr. Genet., 8th. references: Spencer, 1937, DIS 7: 20.
pp. 588-89. phenotype: Slight nick at tip of one or both wings.
phenotype: Slightly less extreme than ct6. RK1. Less than 50 percent penetrance at 19°C, 85 per-
cytology: Salivary chromosomes normal. cent penetrance at 25°, and 100 percent pene-
*cfUa2 trance at 29°. RK3.
origin: X ray induced,
discoverer: Hannah, 1947. cf43aHl

references: 1949, Proc. Intern. Congr. Genet., 8th. origin: X ray induced.
pp. 588-89. discoverer: Muller, Valencia, and Valencia, 1946-53.
phenotype: Like ct^^. RK2A. references: Valencia, 1966, DIS 41: 58.
cytology: Associated with T(l;2)ctl4a2 = phenotype: Male lethal. RK2A.
T(l;2)7B2-4;19-20;41El-2. cytology: Associated with In(l)ct43aH1 = In(l)4Bl-
*cfl4a3 4;7B4-C1 + In(l)10D5-6;20B-C.
origin: X ray induced.
discoverer: Hannah, 1947. origin: X ray induced.
references: 1949, Proc. Intern. Congr. Genet., 8th. discoverer: King and Poulson, 461.
pp. 588-89. references: Poulson and King, 1948, DIS 22: 54.
phenotype: Like ct2a2f but ct^'fa3/Dp(l;3)sn^3ai phenotype: Distal edges of wings scalloped in area
shows reduced viability. RK2A. between L3 and L4, and, occasionally, lateral
cytology: Associated with In(l)ct14a3 = surface of wing toward L5 scalloped. Abdomen,
ln(l)7B2-3;20. antenna, arista, and eye effects of ct absent.
Classification of males and females reliable.
origin: X ray induced. More extreme than ctn and more viable than ct6.
discoverer: Hannah, 1947. ct461/ct6 flies have slightly nicked wing tips
references: 1949, Proc. Intern. Congr. Genet., 8th. resembling ctn. RK1.
pp. 588-89. * c f50e
phenotype: Like cf2«2, RK2A. origin: Spontaneous.
cytology: Associated with Dt(l)ct14bl = discoverer: Bakkum, 5Oe.
Df(l)7B2-3;7C3'4. references: Mickey, 1951, DIS 25: 74.
*CfUb2 phenotype: Wings cut to points. Eyes slightly
origin: X ray induced. ovoid. Viability and fertility lowered, especially
discoverer: Hannah, 1947. in females. RK2.
references: 1949, Proc. Intern. Congr. Genet., 8th.
pp. 588-89. origin: Recovered among progeny of radiofrequency-
phenotype: Like c t ^ . RK2A. treated male,
cytology: Associated with In(l)ct14l>2 - discoverer: Mickey.
In(l)3D2-5;7B2'4. references: 1963, DIS 38: 28.
*cfUc1 phenotype: Like ct. RK1.
origin: X ray induced. *cf62f
discoverer: Hannah, 1947. origin: Spontaneous in Base.
references: 1949, Proc. Intern. Congr. Genet., 8th. discoverer: Mickey, 62f8.
pp. 588-89. references: 1963, DIS 38: 28.
phenotype: Like cf-?«2. RK2A. phenotype: Lethal in male. RK2A.
MUTATIONS 61

*cf268-l *cf268-21
origin: X ray induced. origin: X ray induced.
discoverer: Demerec, 33j. discoverer: Hoover, 35i.
phenotype: Lethal and cell lethal. RK2. phenotype: Lethal. RK2A.
cytology: Salivary chromosomes normal (Hoover). cytology: Associated with T(l;3)ct268-21 -
* c f268-2 T(l ;3)7B3-4; 7B4-5; 96 F.
origin: X ray induced. *cf268-23
discoverer: Demerec, 33k. origin: X ray induced.
phenotype: Lethal but not cell lethal. RK2. discoverer: Hoover, 35g.
cytology: Salivary chromosomes normal (Hoover). phenotype: Lethal but not cell lethal. RK2(A).
*cf268-3 cytology: Salivary chromosomes show possible
origin: X ray induced. deficiency for 7B3.
discoverer: Demerec, 33k. *cf268-24
phenotype: Lethal but not cell lethal. RK2. origin: X ray induced.
cytology: Salivary chromosomes normal (Hoover). discoverer: Hoover, 35i.
*cf268-5 phenotype: Lethal. RK2A.
origin: X ray induced. cytology: Associated with T(l;2)ct268-24 -
discoverer: Demerec, 33k. T(1;2)7B2-5;41F6-42A1.
phenotype: Lethal. RK2A. *cf268-26
cytology: Associated with T(l;3)ct268-5 = origin: X ray induced.
T(l;3)7B2-3;90C4-Dl. discoverer: Hoover, 35j.
*cf268-6 phenotype: Lethal. RK2A.
origin: X ray induced. cytology: Associated with T(l;2)ct268-26 -
discoverer: Demerec, 331. T(1;2)7B3-C1;36E.
phenotype: Lethal but not cell lethal. RK2. *cf26B-27
cytology: Salivary chromosomes normal (Hoover). origin: X ray induced.
*cf268-13 discoverer: Hoover, 35j.
origin: X ray induced. references: 1938, Z. Induktive Abstammungs-
discoverer: Demerec, 34f. Vererbungslehre 74: 420-34.
references: Hoover, 1937, Genetics 22: 634-40. phenotype: Lethal but not cell lethal. RK2A.
1938, Z. Induktive Abstammungs- Vererbungslehre cytology: Associated with In(l)ct268-27 =
74: 420-34. In(l)3D6-El;7B3-5.
phenotype: Lethal but not cell lethal. RK2A. *cf268-29
cytology: Associated with In(l)ct26S-l 3 ~ln(l)2E3- origin: X ray induced.
F1;2F2-3;7B2-3;7B4-5;19A4-5;19A6-B1. Salivary discoverer: Demerec, 38d.
bands 2F1-2, 7B3-4, and 19A5-6 missing. phenotype: Lethal but not shown that lethality at
* c f268-75 ct locus. RK2A.
origin: X ray induced. cytology: Induced simultaneously with but presum-
discoverer: Demerec, 34g. ably separable from T(l;3)lz268'29 =
phenotype: Lethal but not cell lethal. RK2. T(1;3)8D8-9;81F.
cytology: Salivary chromosomes normal (Hoover). *cf268-30
*cf268-17 origin: X ray induced.
origin: X ray induced. discoverer: Hoover, 38d.
discoverer: Demerec, 34h. phenotype: Lethal and cell lethal. RK2A.
phenotype: Lethal. RK2A. cytology: Associated with Df(l)ct268-30 =
cytology: Associated with T(l;2)ct268-l7 - Df(l)7B2-3;7C3-4.
T(l;2)7B2-5;41E2-4 (Hoover). *ct268-31
*cf268-18 origin: X ray induced.
discoverer: Demerec, 34i. discoverer: Demerec, 38d.
references: Hoover, 1938, Z. Induktive phenotype: Lethal. RK2A.
Abstammungs- Vererbungslehre 74: 420—34. cytology: Associated with T(l;3)ct2^S-3l -
phenotype: Lethal. RK2A. T(1;3)3D2-3;7B2-5;84D4-5;86B4-C1;88F (Hoover).
cytology: Associated with InCl^t269"1^ = *cf268-32
In(l)7B2-3;7B4-5;lW8-9. origin: X ray induced.
*cf268-20 discoverer: Demerec, 38e.
origin: X ray induced. phenotype: Lethal. RK2A.
discoverer: Demerec, 35g. cytology: Associated with T(l;'2)ct26S-32 =
references: Hoover, 1938, Z. Induktive T(l;2)lE-F;3D-E;7B2-5;46 (Hoover).
Abstammungs- Vererbungslehre 74: 420—34. *ct268-33
phenotype: Lethal and cell lethal. RK2A. origin: X ray induced.
cytology: Associated with In(l)ct26S"20 = discoverer: Demerec, 38e.
In(l)6Fll-7Al;7B5-6;l OBI 1-12. phenotype: Lethal. RK2A.
62 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

cytology: Associated with T(l;2)ct268~33 - cfK; cut of Krivshenko


T(1;2)7B2-5;41E (Hoover). origin: Spontaneous.
*cf268-35 discoverer: Krivshenko.
origin: X ray induced. references: 1956, DIS 30: 74.
discoverer: Demerec, 38k. phenotype: Both margins, as well as tips, of wing
phenotype: Lethal. RK2. are scalloped. Bristles of mesonotum, and
cytology: Salivary chromosomes normal (Sutton). especially scutellum, are fine as in Minutes.
*cf268-36 Bristle abnormality cannot be separated from wing
origin: X ray induced. effect by crossing over. Viability and fertility of
discoverer: Demerec, 39j. both sexes high. RK1.
phenotype: Lethal. RK2A. cytology: Salivary gland chromosomes appear
cytology: Associated with T(l;3)ct268-36 - normal.
T(1;3)7B2-C1;66F. ct": cut'notch
*cf268-37 origin: Recovered among progeny of heat-treated
origin: X ray induced. flies.
discoverer: Demerec, 39k. discoverer: Ives, 32c3.
references: 1940, Genetics 25: 618—27 (fig.). references: Plough and Ives, 1935, Genetics
Sutton, 1940, Genetics 25: 534-40. 20: 42-69.
phenotype: Lethal. RK2A. 1934, DIS 1: 31.
cytology: Associated with T(l;3)ct268-37 - phenotype: Wings notched at tips. Classification
T(l;3)5D2-3;7B2-3;80C-F. of males reliable, of females harder, but perfect
*cf268-38 at higher temperatures. Viability excellent. RK1
origin: X ray induced. in male.
discoverer: Demerec, 39k. *Cfn4
phenotype: Lethal. RK2. origin: Spontaneous.
cytology: Salivary chromosomes normal (Sutton). discoverer: Mischaikow, 58g.
*cf268-39 references: 1958, DIS 32: 83.
origin: X ray induced. phenotype: Wings notched at tips and inner margins,
discoverer: Demerec, 40a. similar to ctn. Excellent viability and fertility.
phenotype: Lethal. RK2. RK1.
cytology: Salivary chromosomes normal (Sutton). *Ctn63
*Cf268-40 origin: Spontaneous.
origin: X ray induced. discoverer: Datta, 63bll.
discoverer: Demerec, 39k. references: Sarkar, 1963, DIS 38: 28.
phenotype: Lethal. RK2A. phenotype: Wings cut to point and notched.
cytology: Associated with T(l;2;3)ct268-40 - Abdominal bands somewhat warped. Expression
T(l;2;3)7D2-3;10A5-6;21B-C;28-29;40-41;75B- more extreme in females than in males. Classifi-
C;87D;88C;92. cation, fertility, and viability excellent. RK1.
*cf268-4l *cts°: cut of Sytko
origin: X ray induced. discoverer: Sytko.
discoverer: Demerec, 391. references: Agol, 1936, DIS 5: 7.
phenotype: Lethal. RK2A. phenotype: Deeply notched wing tips. Good
cytology: Associated with T(l;2)ct268~41 = expression and viability. RK1.
T(l;2)7B2-5;37C2-3. *ctt: contorted
cf268-42 location: 1-0.3.
origin: X ray induced. origin: Induced by ethyl methanesulfonate (CB.
discoverer: Demerec, 40a. 1528).
phenotype: Lethal. RK2A. discoverer: Fahmy, 1956.
cytology: Associated with D((l)ct26S'42 = references: 1959, DIS 33: 84.
Dt(l)7A5~6;7B8-Cl. phenotype: Wings shorter than normal and abnor-
*Cfdo'vg: cut-dominigene for vestigial mally shaped; frequently curved either convexly or
discoverer: Goldschmidt. concavely. Eyes rough and slightly altered in
references: 1935, Z. Induktive Abstamrnungs- shape. Bristles thinner and straggly; orbitals
Vererbungslehre 69: 36—131 (fig.). frequently reduced or absent. Male genitalia
1935, Biol. Zentr. 55: 535-54. frequently slightly twisted and abnormal. Males
Gardner, 1942, Univ. Calif. (Berkeley) Publ. fertile and females sterile. RK2.
Zool. 49: 85-102. cu; curled
phenotype: ctdo~vif+ interacts with vg/+ to produce location: 3-50,0.
scalloped wings. RK3. origin: Spontaneous.
other information: Presumed by Goldschmidt to discoverer: Morgan, 15115.
enhance dominance of vg and thus termed a references: Morgan and Bridges, 1923, Carnegie
' 'dominigene.'' Inst. Wash. Publ. No. 327: 152 (fig.).
MUTATIONS 63

Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Curled in combination with cu; homozy-
Genet. 2: 215 (fig.), 223. gous lethal. RK2.
Whittinghill, 1937, DIS 7: 22. cytology: Salivary chromosomes normal.
phenotype: Wings curved upward throughout length *Cu: Curl
and slightly divergent. Body color dark. Post- location: 2-55.2 (located using b and en).
scutellars erect and crossed. Good nutrition of origin: Spontaneous.
larvae enhances curled character as does high discoverer: Ives, 48e3.
temperature in last day of pupal life. (Nozawa, references: 1948, DIS 22: 53.
1956, Japan J. Genet. 31: 321-26). RK1. phenotype: Distal part of wing curled as in Cy;
cytology: Shown to be in region 86D2 through 87B2 proximal part to just beyond alula maintains
by its inclusion within the synthetic deficiency lateral compression and indentation fold of
with 3R proximal derived from T(3;4)86D = unfolded marginal wing. Usually an extra cross-
T(3;4)86D2-3;101F and 3R distal derived from vein beyond posterior crossvein extending across
T(Y;3) P102 = T(Y;3)87B2-3 (Cleland). L2, L3, and L4. Cu/+ and Cu/Cu identical; both
have good viability. RK1.
*CuA: Curl-Argentine
origin: Spontaneous.
discoverer: Fernandez Gianotti.
synonym: Ac: Argentine Curly.
references: 1948, DIS 22: 53.
phenotype: Wings curled more strongly than Cy;
waxy texture. Homozygous viable; viability and
fertility good. RK2.
other information: Allelism inferred from similarity
in phenotype and genetic location (2-56.6) to Cu.
Cu-3: Curl in chromosome 3
location: 3-66.0.
origin: Spontaneous.
discoverer: Erickson and Meyer, 51c.
synonym: Cur; Curl preoccupied.
references: Meyer, 1952, DIS 26: 66.
phenotype: Heterozygote has curly wings with
parchment-like texture resembling Cy. Homo-
zygous lethal. RK2.
cu-X: curled-X
cu: curled location: 1- (not located but not allelic to ex).
From Bridges and Morgan, 1923, Carnegie Inst. Wash. origin: Spontaneous in In(l)dl-49+BM1, y sc v.
Publ. No. 327: 152. discoverer: Krivshenko, 57j29.
references: 1956, DIS 32: 80.
*cu700.69 phenotype: Males have wings that are bent upwards
origin: X ray induced. and diverge slightly. cu-X is never expressed in
discoverer: Alexander. females. It represents a mutation whose pheno-
references: Ward and Alexander, 1957, Genetics 42: typic expression is sex limited. Expressed
42-54. equally well in males with and without a Y
phenotype: Like cu. RK1. chromosome. RK2.
cytology: Associated with T(l;3)cul 00.69 - cubitus interruptus: see ci
T(1;3)6B1-C1;88A4-B1. cuh curvi
location: 2-23.4 (1.4 to the right of Sp and 0.5 to the
*cu100.384 right of lys).
origin: X ray induced. origin: Spontaneous.
discoverer: Alexander. discoverer: Nicoletti.
references: Ward and Alexander, 1957, Genetics 42: synonym: curved.
42-54. references: 1957, DIS 31: 84.
phenotype: Curled in combination with cu; homozy- phenotype: Distal half of wing curved upward.
gotes semilethal. RK2. Viability and expressivity very good. RK1.
cytology: Salivary chromosomes normal. cupola: see cpl
cur: curvold
* c u 300.2J5 location: 3-30.
origin: X ray induced. origin: Spontaneous.
discoverer: Alexander. discoverer: Bridges, 33cl4.
references: Ward and Alexander, 1957, Genetics 42: phenotype: Wings divergent and curved down.
42-54. Resembles c. Viability erratic. RK3.
64 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

Cur: see Cu-3 cv-c: crossveinless-c


Curl: see Cu location: 3-54.1 (4.7 units to the left of sb<J2 by
curled: see co C. Hinton).
Curled blistered: see Cb origin: Spontaneous.
curl ex: see ex discoverer: Stern, 25gl3.
Curly: see Cy references: 1934, DIS 1: 35, 36.
Curlyoid: see Cyd phenotype: Posterior crossvein usually absent or
curved: see c greatly reduced. Anterior crossvein usually
curved: see cui present but often detached. Eye flattened or with
Curved of Krivshenko: see C-K vertical shallow furrow. Legs weak, especially
curvi: see cut tarsal joints. Occasionally overlaps wild type.
curvoid: see cur RK2.
cuf: see cf cytology: In region 88A through 88C, based on its
inclusion in the synthetic deficiency with 3R
proximal derived from T(Y;3)PJ02 = T(Y;3)87B2-3
and 3R distal derived from T(3;4)P86 =
T(3;4)88B-C;101 (Bernstein) as well as in the
duplication from T(1;3)O5 = T(1;3)4F2-3;62B-
C;88A-C;92C-D (Lindsley and Grell, 1958, DIS 32:
136).
cv'd: crossveinless-d
location: 3-65.
cv: crossveinless origin: Appeared among progeny of ether-treated
From Weinstein, A., 1920, Proc. Natl. Acad. Sci. U.S. flies.
6: 625-39. discoverer: Duncan, 34c.
references: 1935, DIS 4: 7.
cv: crossveinless phenotype: Posterior crossvein absent or reduced
location: 1-13.7. to an oblique fragment or bar parallel to L5.
origin: Spontaneous. Anterior crossvein sometimes detached. RK2.
discoverer: Bridges, 19112. other information: Possibly an allele of cv-b.
references: 1920, Proc. Natl. Acad. Sci. U.S. 6: *cvw; convex wing
660-63. location: 1-58.2.
Weinstein, 1920, Proc. Natl. Acad. Sci. U.S. 6: origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
625-39 (fig.). phenylalanine (CB. 3026).
phenotype: Crossveins absent or traces only discoverer: Fahmy, 1953.
present. Veins L3 and L4 slightly delta at tips. references: 1958, DIS 32: 69.
Classifiable in unexpended wings. Wing effects phenotype: Wings slightly shortened and arched
due to excessive contraction in the pupal period, convexly. Variable and may overlap wild type.
obliterating the cavity which should normally Tergites in some females have serrated edges or
remain between the epithelia to form the vein are grossly deformed. Viability and fertility good
(Waddington, 1940, J. Genet. 41: 75-139). RK1. in both sexes. RK2.
cytology: Salivary-chromosome studies by Demerec cx; cur/ex
and Sutton show locus to lie from 4F1-2 through location: 1-13.6.
5D1-2 inclusive (Demerec, Kaufmann, Fano, origin: Spontaneous,
Sutton, and Sansome, 1942, Carnegie Inst. Wash. discoverer: R. L. King, 1927.
Year Book 41: 191). Region can be narrowed to phenotype: Wings bent upward for posterior two-
4F9 through 5D2 on basis of inclusion of cv + in thirds of length; anterior one-third warped and
Dp(l;l)ybl « Dp(l;l)lB2-3;4F8-9;5D4-5 margin kinked. Wings not spread. RK2.
(Lindsley). cytology: Salivary analysis by Demerec and Sutton
cv-2: crossveinless on chromosome 2 shows that locus lies from 4F1-2 through 5D1-2
location: 2-96.2. inclusive (Demerec, Kaufmann, Fano, Sutton, and
origin: Spontaneous. Sansome, 1942, Carnegie Inst. Wash. Year Book
discoverer: Nicoletti, 62j. 41: 191).
phenotype: Anterior and posterior crossveins absent. cX: see c(3)G
RK1. cx*9; curl ex-twisted genitalia
cytology: Salivary chromosomes normal. origin: Spontaneous,
*cv-b; crossveinless-b discoverer: Curry, 37cl9.
location: 3-65. phenotype: Wings always divergent, usually 45°
origin: Spontaneous. from axis. Basal one-third of wing wavy, but less
discoverer: Bridges, 24k8. so than in cx; posterior two-thirds of wing curled
phenotype: Crossveins reduced or absent. May slightly upward or downward. Genitalia of nearly
overlap wild type. RK3. all males rotated, usually 45° counterclockwise.
MUTATIONS 65

Flies dwarfish. Viability irregular. Male sterile. d: dachs


RK2. location: 2-31.0.
cx-b: see wy2 origin: Spontaneous.
CxD: see In(3LR)CxD discoverer: Morgan and Bridges, 12k22.
CxF,D: see In(3LR)DcxF references: Bridges and Morgan, 1919, Carnegie
Inst. Wash. Publ. No. 278: 216 (fig.).
Cy: Curly Morgan, Bridges, and Sturtevant, 1925, Bibliog.
location: 2-6.1 (removed from In(2L)Cy and located Genet. 2: 212 (fig.), 223.
by Tinderholt). phenotype: Tarsi four jointed instead of five
origin: Spontaneous. jointed. Legs short and held close to body. Leg
discoverer: L. Ward, 20c. effects enhanced by s s a and ss*B (Villee, 1945,
references: 1923, Genetics 8: 276-300 (fig.)- Genetics 30: 26—27). Wings smaller than wild
phenotype: Wings curled upward; rarely overlaps type, narrowed, with L2 and L3 joined near
wild type at 25 , but frequently overlaps at 19 C. anterior crossvein; distance between crossveins
Curvature caused by the unequal contraction of smaller and crossveins sometimes absent. Angle
the upper and lower epithelia during the drying between L2 and L5 greater than normal. Eyes
period following emergence from the pupa case small and rough. Posterior scutellar bristles
(Waddington, 1940, J. Genet. 41: 75-139). Clas- erect. Viability erratic. Frequently sterile. RK2.
sifiable in single dose in triploids. Usually dU see Df(2L)d
homozygous lethal, but may emerge as dwarf with
more extreme wing character. RK1A.
cytology: Ordinarily inseparable from In(2L)Cy =
In(2L)22Dl-2;33F5-34Al, although it has been
separated by Tinderholt (1961, DIS 35: 47).
other information: Cy removed from In(2L)Cy still
causes a local reduction in crossing over in the
ed-cl region (Sederoff).

D: Dichaete
From Bridges and Morgan, 1923, Carnegie Inst. Wash.
Publ. No. 327: 127.

D: Dichaete
location: 3-40.7 (40.4-41.0).
origin: Spontaneous.
discoverer: Bridges, 15a3.
references: Bridges and Morgan, 1923, Carnegie
Inst. Wash. Publ. No. 327: 127 (fig.).
phenotype: Wings extended uniformly at 45 from
body axis and elevated 30° above (occasionally
Cy: Curly sharply downcast and dragging). Alulae missing.
From L. Ward, 1923, Genetics 8: 276-300. Dorsocentrals and some other bristles reduced in
number (Sturtevant, 1918, Carnegie Inst. Wash.
Cyd: Curlyoid Publ. No. 264; Plunkett, 1926, J. Exptl. Zool.
location: 3- (rearrangement). 46: 181-244). Head often deformed or split in
discoverer: Jollos. postvertical region. Halteres turned down. Homo-
references: Curry, 1939, DIS 12: 46. zygous lethal. Nearly lethal in combination with
phenotype: Wings curled upward in heterozygote. eyD (Sobels, Kruijt, and Spronk, 1951, DIS 25:
Homozygous lethal. RK2A. 128). Partially suppressed by sc alleles that re-
other information: Associated with an inversion of move postverticals (.sc, sf*. sc6, sc7) but not by
3R; possibly In(3R)P. others (sc^, sc$) (Sturtevant). Ciassifiahle in trip-
cy/: see rA'c>' loids. RK2A.
66 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTEFl

cytology: Inseparable from Jn(3L)D = In(3L)69D3- dark eye: see dke


El;70C13-Dl (Bridges in Morgan, Bridges and dark eye1: see sf3 2e
Schultz, 1937, Carnegie Inst. Wash. Year Book dark red brown: see drb
36: 301). Darkened eye: see Dke
darkener of w/i/fe-eos/n: see dark
origin: Spontaneous; derivative of D. darker legs: see thld
discoverer: Plunkett, 24f. darky: see dar
references: 1926, J. Exptl. Zool. 46: 181-244. daughterless: see da
phenotype: Less extreme than D. Wings extended db: dark body
and lifted; alulae missing. Head effect of D location: 3-44.4.
missing. Bristles usually wild type; occasionally origin: Spontaneous.
outer verticals, upper humerals, presuturals, and discoverer: Chovnick and Talsma, 1965.
anterior postalars absent. Viability of D^/+ references: 1966, DIS 41: 58.
better than D/+. RK2A. phenotype: Body color darker than normal. Male
cytology: In(3L)D present as in D. rarely survives; dies in late pupal stage. RK2.
*DE: Dichaete-Extended DcX: see ln(3LR)CxD
origin: Spontaneous in £>/+ culture; probably a DcxF: see In(3LR)DcxF
modified D. *dd: displaced
discoverer: Sturtevant, 16fll. location: 1-24.3.
references: Bridges and Morgan, 1923, Carnegie discoverer: Bridges, 31d7.
Inst. Wash. Publ. No. 327: 165 (fig.). phenotype: Antennae sunken into shortened head;
phenotype: Wings divergent but not bent at base. eyes also deformed. Females often sterile. RK2.
Bristles and alulae normal. Overlaps wild type. cytology: Locus lies between 7C4-5 and 8C1-2
Homozygous lethal; D/D^ is lethal. RK3(A). (Demerec, Kaufmann, Fano, Sutton, and Sansome,
da: daughterless 1942, Carnegie Inst. Wash. Year Book 41: 191).
location: 2-39.3. Further restricted to 7E1 through 8C2, on the basis
origin: Spontaneous. of its genetic location to the right of oc which is
discoverer: Bell, excluded from Df(l)sn = Df(l)7B2-3;7D22-El
references: 1954, Genetics 39: 958—59. (Hintonand Welshons, 1955, DIS 29: 125-26).
1954, DIS 28: 73. *ddh displacedlike
phenotype: Homozygous da females, mated to any location: 1-27.2.
male, produce normal sons but no daughters. origin: Induced by triethylenemelamine (CB. 1246).
Otherwise, da/da individuals appear wild type. discoverer Fahmy, 1953.
Lethal action of da occurs in egg stage. Counce synonym: dd$.
finds lethal female embryos show consistent references: 1959, DIS 33: 84.
abnormalities in midgut formation. In about half phenotype: Frontal region with antennae sunken
the abnormal embryos, shortening of germ band into shortened head. Eyes deformed. Thoracic
fails and anus and posterior spiracles open on bristles stiff and slightly shortened. Wings fre-
dorsal surface behind head segments. Differentia- quently misheld. Males sterile and viability
tion of almost all other tissues surprisingly slightly reduced. RK2.
normal. RK3. other information: One X-ray-induced allele.
cfe; see dar *de: deacon
dachs: see d location: 1-56.
dachsous: see ds origin: X ray induced.
*dar: darky discoverer: Muller, 26112.
location: 1-0 (no crossovers with sc in 547 flies). references: 1935, DIS 3: 29.
origin: X ray induced. phenotype: Body and wings narrow and rectangular.
discoverer: Fahmy, 1956. Eyes slightly flattened, with oblique cast. RK3.
synonym: da; preoccupied. other information: Possibly an allele of si (1-53.5).
references: 1959, DIS 33: 84, De; Dented
phenotype: Small, heavily melanized flies. Some- location: 2- (between dp and b).
times wings curl upward. Male sterile; viability origin: X ray induced,
about 15 percent wild type; late ©closing. RK2. discoverer: Belgovsky, 36c.
*dark: darkener of white-eosin references: 1937, DIS 8: 7.
location: Autosomal. phenotype: In heterozygote most flies show one or
discoverer: Bridges, 13i23. two indentations on thorax at front. Homozygote
references: 1916, Genetics 1: 148. has two smaller, sharper dents. Wings often
1919, J. Exptl. Zool. 28: 347. raised. RK3.
phenotype: Specific partial suppressor of we. RK3. deacon: see de
dark: see dk deep orange: see dor
dark body: see db defective: see df
dark bubbly: see dkb deflected wing: see dfw
MUTATIONS 67

Deformed: see Dfd folded flat at middle. Eyes sometimes rough and
deformed antennae: see dfa bulging. Wings slightly spread. Bristles tend to
deformed eye: see dfi break; scutellars occasionally doubled. RK3.
deformed terga: see dft Detached: see Dt
deformed wings: see dwg *df: defective
degenerated spermatheca: see dg-a location: 1-32.5.
Delta: see Dl origin: Spontaneous.
delta vein: see thvd discoverer: Bridges, 1513.
delta wing: see dta phenotype: Head bristles around ocelli missing.
deltex: see c/x Viability poor. RK3.
deltoid veins: see c//v *dfa: deformed antennae
Dented: see De location: 1-13.9.
*</ep: depressed origin: Induced by 2-chloroethyl methanesulfonate
location: 1-18. (CB. 1506).
discoverer: Bridges, 13d. discoverer: Fahmy, 1956.
references: Morgan and Bridges, 1916, Carnegie references: 1959, DIS 33: 84.
Inst. Wash. Publ. No. 237: 67 (fig.). phenotype: Wings short, broad, either convex or
phenotype: Wings turned down at tips, flat from concave, and abnormally held. Eyes small, dark,
side to side. Somewhat variable but does not and rough. Bristles short, stiff, occasionally bent.
overlap wild type. RK2. Trident pattern more pigmented. Abnormal
*depl: depressedlike antennae and aristae. Males viable and fertile.
location: 1-23. Females sterile. RK2.
origin: Recovered among progeny of flies treated
with Janus green,
discoverer: Muller, 28e20.
synonym: depr: depressed-roof.
references: 1935, DIS 3: 29.
phenotype: Wings droop at sides. Flies dark and
weak; bristles fine. Viability variable, about
20 percent wild type. RK3.
depressed: see dep
depressedlike: see depl Dfd: Deformed
*der: deranged From Bridges and Morgan, 1923, Carnegie Inst. Wash.
location: 1-57.2. Publ. No. 327: 94.
origin: Induced by triethylenemelamine (CB. 1246).
discoverer: Fahmy, 1953. Dfd: Deformed
references: 1958, DIS 32: 69. location: 3-47.5.
phenotype: Thoracic hairs deranged; many point origin: Spontaneous.
toward midline. Wings usually obliquely upheld discoverer: Cattell, 13g.
and twisted, bringing inner margins together. references: Bridges and Morgan, 1923, Carnegie
Overlaps wild type. Good viability in both sexes, Inst. Wash. Publ. No. 327: 93 (fig.).
but female fertility reduced. RK3. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
Genet. 2: 223 (fig.).
phenotype: Eye reduced ventrally and anteriorly, or
furrowed. Vibrissae tufted. Extremely variable,
often overlapping wild type, but usually excellent
character with Ly or at 19°C. Classifiable in
single dose in triploids (Schultz, 1934, DIS 1: 55).
Homozygous lethal, although Muller reported an
occasional homozygote as extreme dwarf with
flimsy wings. RK2.
det: detached cytology: Salivary gland chromosomes appear
From Bridges and Brehme, 1944, Carnegie Inst. Wash. normal.
Publ. No. 552: 54. *Dfd38k
origin: Spontaneous.
det: detached discoverer: Mossige, 38k30.
location: 3-72.5. references: 1939, DIS 12: 47.
origin: Spontaneous. phenotype: Like Did. Homozygous lethal. RK2,
discoverer: Nichols-Skoog, 35k27. Dfdr; Deformed-recessive
phenotype: Posterior crossveins detached from origin: Spontaneous.
longitudinals at one or both ends and may be discoverer: Bridges, 3014.
absent. Wings occasionally folded back under or synonym: &m: almond.
68 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

phenotype: Eye small, narrow, and kidney shaped. dfw: deflected wing
Overlaps wild type in older cultures. Dfdr/Dfd location: 1-21.6.
more extreme than Dfd/+, RK2. origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
*Dfdr2 phenylalanine (CB. 3025).
origin: Spontaneous. discoverer:: Fahmy, 1955.
discoverer: Pierce. references: 1959, DIS 33: 84.
references: 1945, DIS 19: 46. phenotype: Wings slightly divergent and upheld to
phenotype: Eyes smaller and more constantly kidney various degrees, often twisted on their axes. Inner
shaped than in Dfdr. Wings thin, dull, uneven, margins frequently incised; occasionally wing
slightly spread (about 60° from body axis), and membranes separated by fluid. Eyes slightly
drooping. Body slightly smaller and lighter colored smaller. Males viable and fertile. Females
than normal. Bristles shortened and delicate. Last sterile; viability reduced. RK2.
abdominal segment of male may be rotated. Via- other information: One X-ray-induced allele.
bility low. Dtdr2/DfdT slightly more extreme than *dg-a: degenerated spermatheca
Dfdr/Dfdr. RK2. location: 3-75.5.
origin: Spontaneous.
origin: Spontaneous. discoverer: Collins, 21a.
discoverer: Hollander, 1957. references: Wexelsen, 1928, Genetics 13: 389-400
references: 1960, DIS 34: 51. (fig.).
phenotype: Kidney-shaped eye. Penetrance and phenotype: Adult females show degeneration and
expressivity variable. RK2. pigmentation of epithelial cells of spermathecae
24 hr or more after eclosion. Viability and
origin: Spontaneous. fertility good. Penetrance 100 percent. RK3.
discoverer: Kidwell. dh: see eg2
references: 1961, DIS 35: 46. *di: dimorphos
phenotype: Eyes reduced. Expression varies from location: 1- (near spindle attachment).
absence of both eyes to wild type. Penetrance origin: Spontaneous.
varies from 75 to 100 percent. Penetrance discoverer: Harnly, 32dlO.
increased by selection for reduced eye. About 5 references: 1935, J. Exptl. Zool. 72: 75-99 (fig.).
percent of Dfdr60/+; ey/+ flies exhibit deformed 1940, DIS 13: 49.
phenotype. RK2. phenotype: Specific lengthener of vg wings, espe-
Dfdri-: Deformed-recessive of Luers cially in males (di; vg female much like vg). At
discoverer: Luers. higher temperatures, eyes small and rough and
references: Vogt, 1946, Experientia 2: 313-15. wings of both sexes approach wild type. RK2 in
1947, Biol. Zentr. 66: 81-105 (fig.)- vg male.
phenotype: Like Dfdr. RK2. dibrd: see frdi
*dfi: deformed eye Dichaete: see D
location: 3- (near D). dihedral: see eg2
origin: Recovered among descendants of heat-treated *dil: specific dilutor
flies. location: 2-57.
discoverer: Ives, 32c. origin: Spontaneous,
synonym: rough III. discoverer: Bridges, 32f22.
references: Plough and Ives, 1934, DIS 1: 34. phenotype: Dilutes bw to pale yellowish brown,
1935, Genetics 20: 42-69. and we, we2 and wb* to paler grades. RK3.
phenotype: Eyes roughish, reduced, and misshapen. *dil-3: dilute in chromosome 3
Overlaps wild type. Female sterile, poorly location: 3- (not located).
viable. RK3. discoverer: Bridges, 1519.
*dft; deformed tergi references: Bridges and Morgan, 1923, Carnegie
location: 1-33.7. Inst. Wash. Publ. No. 327: 151.
origin: Induced by 2-chloroethyl methanesulfonate phenotype: Eye color like maroon, overlaps wild
(CB. 1506). type. RK3.
discoverer. Fahmy, 1956, *c///-w°: dilutor of wft/fe-opricof
references: 1959, DIS 33: 84. location: 3- (not located).
phcnotype: Small fly with small, slightly rough discoverer: Weinstein.
eyes. Wings slightly divergent or upheld, abnor- references: Morgan, Bridges, and Sturtevant, 1925,
mally shaped with occasional incision of the inner Bibliog. Genet. 2: 218.
margin. Bristles slightly thinner and shorter with phenotype: Lightens » a . RK3.
one or both postscutellars frequently absent; and dilute in chromosome 3: see dil-3
a dorsocentral occasionally missing. Abdominal dilute ocelli: see po2
segmentation deformed to various degrees; Dilute-1: see
abdominal hairs fewer and deranged. Males poorly Dilute-2: see
fertile, viability about 50 percent wild type. RK2. Dilute-3: see
MUTATIONS 69

Dilute-4: see bwV30ki2 phenotype: Eye color soft, dull, and dark, like sf.
Dilute-5: see bwV3 0ki3 sf/dke is wild type. Flies have 65 percent normal
Dilute-6: see bwV30kis red pigment and 98 percent normal brown pigment
dilutor of white-apricot: see dil-wa (Nolte, 1955, J. Genet. 53: 1-10). RK2.
diminutive: see dm Dke: Darkened eye
dimorphos: see di location: 2- (not located).
Din: Dinty origin: X ray induced.
location: Unknown; associated with a rearrange- discoverer: Hendrix. 1963.
ment. references: 1964, DIS 39: 58.
origin: X ray induced. phenotype: In heterozygotes eye facets roughened
discoverer: Braver, 55a. with black-spotted pigmentation, varying from light
references: 1955, DIS 29: 70. spotting near margin of eye to heavy pigmentation
Pollock, 1963, DIS 38: 50. covering one-half of the eye. A bleached area
phenotype: In male and heterozygous female, sometimes appears adjacent to the pigmentation.
central portion of vein L2 interrupted. Posterior Effect usually symmetrical. Homozygous lethal.
supra-alar bristles absent in 95—99 percent of RK3.
females and 97—99.5 percent of males. Anterior cytology: Salivary chromosomes appear normal
postalar bristles absent in 6—11 percent of (Peacock).
females and 2—6 percent of males. Wings dkl: see thld
divergent. Viable and fertile in male and hetero-
zygous female; homozygous lethal. RK2.
cytology: Associated with T(l;2;3)Din =
T(1;3)3C;63A + T(2;3)39D;73A.
*dis: distorted eye
location: 1-23.
origin: Recovered among progeny of natural-gas-
treated fly.
discoverer: Mickey, 49b5.
references: 1951, DIS 25: 74.
phenotype: Whole or part of eye roughened. Some-
times bristles absent or doubled. Wings may be
roughened with nicked margins and plexus veins.
Expressivity variable. RK3.
cytology: Salivary chromosomes appear normal.
Discolored: see bwv?
dishevelled: see dsh
displaced: see dd
displaced!ike: see ddl
disrupted: see dsr
distorted eye: see dis
disturbed segmentation": see dss DI: Delta
divergent: see cfv From Bridges and Morgan, 1923, Carnegie Inst. Wash.
divergent wings: see dvw Publ. No. 327: 197.
divers: see dvr
*dk: dark DI: Delta
location: 3- (not located). location: 3-66.2.
discoverer. Clausen, 20g. origin: Spontaneous.
references: Bridges and Morgan, 1923, Carnegie discoverer: Bridges, 18k30.
Inst. Wash. Publ. No. 327: 235. references: Bridges and Morgan, 1923, Carnegie
Morgan, Bridges, and Sturtevant, 1925, Bibliog. Inst, Wash. Publ. No. 327: 197-201 (fig.).
Genet. 2: 223. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Eye color maroon. Overlaps wild type. Genet. 2: 75 (fig.).
RK3. phenotype: Veins thickened and broadened into
*dkb: dark bubbly deltas at junction with margin. Eyes somewhat
location: 2- (to the left of vg). small and slightly roughened. In extreme cases,
discoverer: Bridges, 38d25. ocelli run together into a crescent that encloses
phenotype: Thorax has dark bubbly longitudinal the ocellar bristles. Hairs on thorax straggly and
streak. RK3. more numerous. Body color dark. Wings small,
dke: dark eye dark, and somewhat spread. Effects of DI neu-
location: 2-73. tralized by H. DI and several of its alleles shown
origin: Spontaneous. to interact synergistically with certain Minutes,
discoverer: Bridges, 38cll. producing extreme phenotype and drastically
70 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

lowered viability (Schultz, 1929, Genetics 14: references: 1956, DIS 30: 71.
366-419). Homozygous lethal. RK2. MacDonald, 1957, DIS 31: 84.
cytology: Located in region 91D1-92A2, inclusively, phenotype: Spreading of veins most marked for L2
based on its inclusion within the transposed sec- and L5. Posterior crossvein thickened, coalesc-
tion of Tp(3)bxdU0 = Tp(3)89E2-3;91C7-Dl;92A2-3 ing with the delta of L5. L2 slightly thickened.
(E. B. Lewis) and in Df(3R)Dl» = D((3R)91C6- Expression more extreme at higher temperature,
Dl;92A2-3 (Slizynski). Heterozygous deficiency especially in males. Spreading of veins apparent
for 91D1-92A2 produces the Dl phenotype since in the 40- to 41-hr-old pupa. Suppressed by H.
Tp(3)bKd*l°/+ is D/ + , whereas Df(3R)bxdU0/ + j s Lethal homozygous and in combination with Dl3.
Dl. RK2.
cytology: Salivary chromosomes normal.
origin: Spontaneous. 1; Delta-Barish
discoverer: Bridges, 24110. discoverer: Schultz, 1933.
phenotype: Like Dl but deltas at margin are slight; phenotype: Delta venation and eyes of a broad
longitudinal veins between anterior and posterior heterozygous Bar type. Homozygous lethal. RK2A.
crossveins and crossveins themselves are thick- cytology: Associated with In(3R)DlB =
ened. Spreading of wing slighter than in Dl. In(3R)90A;91A (Schultz).
Better viability than Dl. Homozygous lethal. *DICf-3: Delta-Confluent
RK1. origin: Spontaneous.
*DI4 discoverer: Imaizumi.
origin: Spontaneous. synonym: Cf-3.
discoverer: Bridges, 26g28. references: 1962, DIS 36: 38.
phenotype: Slight deltas at margin; posterior parts phenotype: Like Dl. Homozygote dies as late
of L2 and L3 thickened to delta at margin; L4 and embryo or early larva. RK2.
L5 slight. RK2.
other information: Gives increased crossing over origin: Induced by unspecified chemical (probably
throughout the third chromosome. mustard gas).
Dis discoverer: Auerbach.
origin: X ray induced. references: 1943, DIS 17: 49.
discoverer: R. L. King, 32d. phenotype: Homozygous lethal. RK2A.
phenotype: Resembles Dl but deltas are slight. cytology: Associated with Dl(3R)DlH =
Wings occasionally vesiculated; only slightly Df(3R)91C6-Dl;92A2-3 (Slizynski).
spread. Homozygous lethal. RK3. *D/': Delta-lethal
other information: heterozygote shows reduced origin: Spontaneous.
crossing over. discoverer: Bridges, 38clO.
*DI6 synonym: 1(3 )DL
origin: X ray induced. phenotype: Heterozygote normal; homozygote not
discoverer: Schultz, 33a5. tested; thought to die in combination with Dl12.
phenotype: Extreme Dl. Veins at basal part of RK3.
wing thicker than Dl; thickening marked along other information: Presence of a recessive lethal
entire L2, with a striking confluent delta at allele of Dl on the H chromosome inferred from
margin; L3 has basal and distal thickening and failure to observe Dli2/H progeny from cross of
marginal delta; L4 extremely broad, especially h cu H2 ca/In(3R)P, Did ca X Dl^2/? A recessive
beyond posterior crossvein ending in large delta. lethal allele of H on the Dl*2 chromosome seems
Posterior crossvein moderately thick. Wings an equally likely interpretation.
blistered and wrinkled. Veins inhibited from *DlOf: Delta-Overflow
narrowing in late pupal period [Waddington, 1940, origin: Spontaneous.
J. Genet. 41: 75—139 (fig.)]. Homozygous lethal. discoverer: Tsukamoto, 1956.
RK2. synonym: Of.
references: 1956, DIS 30: 79.
origin: X ray induced, 1957, DIS 31: 85.
discoverer: Schultz, 33a7. phenotype: Slight deltas at margins; striking con-
phenotype: Like D/*. RK3. fluent effects on longitudinal veins, especially L2
and L5, and near posterior crossvein. Eyes nearly
*DI7P: Delto-7 of Pan shin normal, but with spac**, posterior half of eye
discoverer Panshin, 1935. surface resembles spaPot. Suppressed by H.
references: 1935, Dokl. Acad. Nauk SSSR 4: 85-88. Lethal homozygous and in combination with Dl12.
phenotype: Like Dl$. RK3A. RK1.
cytology: Associated with T(3;4)Dl^p. dlv: deltoid veins
location: 1-25.9.
origin: X ray induced. origin: Induced by S-2-chloroethylcysteine
discoverer: Clark. (CB. 1592).
MUTATIONS 71

discoverer: Fahmy, 1957. dor/dor and dor/+ females accumulate more


references: 1959, DIS 33: 85. isoxanthopterine than wild-type females, but dor
pbenotype: Wings small, abnormal, with margin males contain less than wild-type males. These
occasionally incised, and frequently either diver- differences detectable in prepupal stages. Ommo-
gent or slightly upheld. Extra venation, especially chrome pigments also affected. Reciprocal trans-
at junctions between longitudinal and costal plantation experiments show that eye color is
veins, giving Delta-like formations. In extreme autonomous (Hadorn and Counce). Homozygous
cases, wings grossly deformed and blistered. dor females exhibit same sterility effects as tu
Excess melanization throughout body. Eyes dark, (Merrell, 1947); i.e., dor progeny of dor mothers
small, and slightly rough. Total body size die. dor females crossed with wild-type males
reduced. Both sexes viable and fertile. RK1. produce some dor/+ daughters. Under certain
other information: One allele induced by CB. 1592. conditions dor males sterile as result of excess
dm: diminutive accumulation of preadult fat, which mechanically
location: 1-4.6. prevents union of gonads with rudiments of rest of
discoverer: Nichols-Skoog, 33j9. genital system (Counce). Post-blastulation devel-
references: 1935, DIS 3: 10. opment of dor progeny of dor females abnormal,
phenotype: Bristles and body small and slender. leading to embryonic death (Hildreth and
Viability excellent. Females sterile. RK1. Lucchesi). Cellular degeneration begins before
cytology: Locus placed between 3C9 and 3D2 by gastrulation is ended, and by 16 hr, degeneration
Slyzinska (1938, Genetics 23: 291—99), at 3C9 by of embryo is almost complete. Some embryos die
Schultz, and at 3D1-2 by Demerec, Kaufmann, at early cleavage regardless of sex. Eggs of dor
Fano, Sutton, and Sansome (1942, Carnegie Inst. females contain less than normal amounts of yolk
Wash. Year Book 41: 191). (Counce, 1956). The double mutants dor; ry, and
dm264-S8 dor, ry2 are lethal (Luchessi, 1963, Proc. Intern.
origin: X ray induced. Congr. Genet, 11th. Vol. 1: 169-70). RK1.
discoverer: Demerec, 38d. cytology: Placed in region from 1F1 through 2A2
references: 1940, Genetics 25: 618-27. on the basis of its inclusion in Dp(l;f)1337 =
phenotype: Described only as nonvariegated. RK2A. Dp(l;f)lF4-2A3; 19-20 but not in Dp(l;f)112 =
cytology: Associated with T(1;3)N264-58 ~ Dp(l;f)lE4-Fl; 19-20 (R. F. Grell, Gersh).
T(l;3)3B2-3;3D6-7;80D-F (Sutton).
dn: doughnut origin: Spontaneous.
location: 3-50. discoverer: Hildreth, 61e.
origin: Spontaneous. references: 1963, DIS 37: 48.
discoverer: Wallbrunn. phenotype: Orange eye color. Poor viability. RK2.
references: 1942, DIS 16: 54.
Wright, 1946, DIS 20: 68. dor1: deep orange-lethal
phenotype: Eye of se dn has unpigmented spot (in origin: Spontaneous.
middle or toward posterior) at emergence from discoverer: Bridges, 15al.
puparium. Spot gradually darkens; after 2 days synonym: 1(1)7: lethal(l) 7.
eyes appear sepia. Difficult to detect with wild- references: 1916, Genetics 1: 149.
type eye color; appears as slightly lighter red phenotype: Male larvae die 90—lOOhr after
spot, which disappears after 2 days. Viability hatching. According to Russell [l940, J. Exptl.
low; many die as pupae at 25°C. Viability nearly Zool. 84: 363—79 (fig.)], a gut abnormality appears
normal at 17°C, but character not detectable. at 65 hr resulting in obliteration of gut lumen and
Both sexes highly infertile; testes about one-third loss of all food material. Stark observed me Ian otic
normal length. Spermathecae very small. External tumors [1918, J. Cancer Res. 3: 279-301 (fig.);
genitalia of both sexes often abnormal. RK3. 1919, J. Exptl. Zool. 27: 509-29 (fig.)]. Oftedal
dd: see po2 studied histology of dor1 larvae (1953, Z. Induktive
dor: deep orange Abstammungs- Vererbungslehre 85: 408—22).
location: 1-0.3 [allelic to dor1 (Redfield and Malpighian tubes nearly colorless except at base.
Schultz; Clancy)]. dor1/'dor has orange eyes like dor/dor as well as
origin: X ray induced. the sterility effects (Clancy; Redfield and
discoverer: E. D. King. Schultz). RK2.
references: Merrell, 1947, Am. Naturalist 81:
399_400. origin: X ray induced.
Counce, 1956, Z. Induktive Abstammungs- discoverer Alikhanian.
Vererbungslehre 87: 443-61 (fig.). synonym: 1(1)76.
phenotype: Eye color true orange at 25°C, red- references: Ardoshnikov, 1941, Dokl. Akad. Nauk
orange at 18°. Chromatographic studies (Counce, SSSR 30: 344-46.
1957, Experientia 13: 354) indicate pteridine phenotype: Like dor1. Incidence of dor!2/Y larvae
patterns differ from wild type. Red pigment bearing melanotic masses reduced if one parent
reduced (but increased in heterozygous females); carries a free duplication carrying dor*. RK2.
72 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

two, or all of these phenotypes. For example:


origin: Spontaneous. dp° (o) has oblique wings, but no vortices, and is
discoverer: H. W. Lewis. not lethal; dpov (ov) has oblique wings and
references: 1954, J. Exptl. Zool. 126: 235-75 (fig.). vortices, but is not lethal; and dpol,v (olv) has all
phenotype: Like dorK Survival of larvae containing three attributes. The three recessive character-
melanotic masses optimal at 25°C. dorI3/Basc fe- istics o, I, and v complement one another. Thus
male shows more rapid development and higher via- dpol/dpv, dp°v/dpl, and dp°/dplv are phenotyp-
bility than +/Basc female. RK2. ically wild type; dp°/dp°", dpol/dp°v, and
*double: double dp°/dp°lv are o; dp°v/dpv, dplv/dp°v, and
location: 1-0. dpv/dpolv are v; any two lethals are lethal in the
origin: Spontaneous. trans heterozygote.
discoverer: Bridges, 1918, cytology: Located between 24E2 and 25A2 based on
references: Morgan, Bridges, and Sturtevant, 1925, its inclusion in Df(2L)M-zB = Df(2L)24E2-
Bibliog. Genet. 2: 224. F1-25A1-2 (Morgan, Bridges, Schultz, and Curry,
phenotype: Postvertical bristles doubled. Wings 1939, Carnegie Inst. Wash. Year Book 38: 273-77).
very small. Viability somewhat low. RK3. other information: The alleles at the dp locus are
Double Bar: see BB pseudoallelic and have been positioned by Carlson
Double Inhabar: see B'Bi (1959) and Southin and Carlson (1962) (see map).
double sex: see dsx Carlson often puts the superscripts on the line and
Doublet: see Dp(l;l)BSRMG has used an apostrophe instead of dp; dp o / " may
doughnut: see dn be written, therefore, as olv or 'olv.

Im
olm
olvW
olvbm
obm o2 olv cm2 Iv h ov ovS2b olS v2
1 .—• 1 1 1

Map of the dp locus


From Southin and Carlson, 1962, Genetics 47: 1017-
26.

dp: see dp°v


dp*: see dplv?

origin: X ray induced.


discoverer: Fogel, 1949.
references: 1950, DIS 24: 57.
other information: A series of 13 alleles of diverse
phenotype. Viability data given (Fogel, 1950).
See

rfow: downy dps*1: see


From Bridges and Brehme, 1944, Carnegie Inst. Wash. dpS2b; see
Publ. No. 552: 64. *dpS8
origin: Spontaneous.
dow: downy discoverer: Fradkin, 1958.
location: 1-8.0. references: 1958, DIS 32: 79.
origin: Spontaneous. other information: A series of 14 alleles; not
discoverer: Bridges, 36c28. described or tested for viability.
phenotype: Bristles very short and slender, nearly dp61d
as small as as. Males entirely sterile; testis origin: X ray induced.
shape normal. Viability good, RK2. discoverer: Thompson, 61d.
dp; dumpy cytology: Associated with T(Y;2)dp(>1 d.
location: 2-13.0. b see dpobw
references: Carlson, 1958, Ph.D. Thesis, Indiana *c/pcm: dumpy-comma
Univ. origin: Spontaneous.
1959, Genetics 44: 347-73 (fig.1. discoverer: Bridges, 13b5.
Seuthin and Carlson, 1962, Genetics 47: 1017-26 references: Bridges and Morgan, 1919, Carnegie
Inst. Wash. Publ. No. 278: 228 (fig.).
phenorype-: Alleles of dp produce three general phenotype: Sharply outlined depressions of comma
phermtypes: oblique wings Co), vortices on thorax shape at anterior edge of thorax. Penetrant in 20
O), and lethal CO. A specific mllele may have one, percent of females but rarely in males. RK3.
MUTATIONS 73

<Jpcm2
effect in dp!^/-h; Me/+ and dpIM +/+ ta. Pheno-
origin: Spontaneous. typically normal in combination with dp°2( dpv?,
discoverer: Meyer, 53c. dp o v , and dpc™2. RK2.
references: 1955, DIS 29: 74. dpL': see
phenotype: Homozygotes nearly normal at 25°C,
with slightly oblique wings in some. Anterior dpLS: see
edge of thorax usually shows a pair of commalike
depressions; wings may occasionally have
blisters. dpcm2/dpov shows good commas in all dpi*: dumpy-lethal vortex
flies, vortices in most, and slightly oblique origin: Spontaneous.
wings. dp<>™2/dpolv is semilethal at 22°C, and discoverer: Bridges, 20c3.
survivors are similar in phenotype to dpov/dpolv. synonym: dptx: dumpy-thoraxate.
Degree of viability of dpcm2 when heterozygous phenotype: Homozygous lethal; no abnormalities
with various dp °'"-like alleles varies, but is when heterozygous witti wild type. dplv/dpov has
usually lower at low temperatures (Carlson and strong volcanolike dorsocentral mounds or pits
Falk, 1962, DIS 36: 59-61). RK3. with brown pigmentation; comma effects striking;
sternopleural bristles turned down; wings normal.
dpD: dumpy-Dominant At higher temperature (28°C), two additional pits
origin: X ray induced. anterior to dorsocentrals occasionally appear.
discoverer: E. B. Lewis, 1962. Lethal in compound with dpo1, dpo1^, and dplM%
synonym: olvD. RK2.
references: Del Campo, 1963, DIS 38: 32. dpi* *
phenotype: dpD/+ has slightly oblique wing and origin: Spontaneous within In(2L)Cy.
moderate vortex and comma effects on thorax. discoverer: Muller.
dpD/dp°v has strong wing and thorax effects, with synonym: dp2.
reduced leg and body size. dpD/dpolv is lethal, phenotype: Similar to dpIv; slight oblique wing
and dpD resembles dpo!v in several ways. effect in dplv2/dpov; comma effects occasionally
dpD/dpv has strongly enhanced thorax effects, seen in d p ^ 2 / + . R K 2 .
and dp°/dpD has reduced wings with possibly *c/p/v57e
enhanced thoracic effect. Homozygous lethal. origin: Ultraviolet induced.
RK2A. discoverer: Meyer, 51e.
cytology: Associated with T(2;3)dpD = synonym: dp^"^5Je: dumpy-thoraxate 51e.
T(2;3)25A;95B-D. references: Meyer and Edmondson, 1951, DIS 25: 72.
dpdei: s e e r f p o d e f phenotype: Similar to dplv. Occasionally
dpdr: see dpovdr dplv51 e/dpov shows blistering of wings but no
*dpG; dumpy of Goldschmidt oblique truncation. RK2.
origin: Spontaneous.
discoverer: Goldschmidt. dplvl: dumpy-lethal vortex of Ives
references: 1945, Univ. Calif. (Berkeley) Publ. origin: Spontaneous within In(2t,)Cy.
Zool. 49: 511, 520. discoverer: Ives, 39j.
other information: Several alleles; not analyzed in synonym: dptx^i dumpy-thoraxate of Ives;
detail for effect.
dph: dumpy-humpylike phenotype: Identical to dplv. RK2.
origin: Spontaneous, dpNov: see
discoverer: Edmondson, 54g.
references: 1955, DIS 28: 73.
phenotype: As homozygote, has strongly truncated
wings, four marked thoracic humplike vortices,
strong comma effect, reduced body size, and
small, weak legs. Because of weak legs, flies
have difficulty walking and become stuck in the
food. No body or leg reduction in dph/dpv2 but
vortices and commas present as in dpv2/dpolv. dp°: dumpy-oblique
dph viable with dpo'S, dp°lv, dplv, and dp1^, From Bridges and Brehme, 1944, Carnegie Inst. Wash.
RK2. Publ. No. 552: 65.
dpH: see dpoivH
dplM: dumpy-lethal of Meyer dp°: dumpy-oblique
origin: Ultraviolet induced. origin: Spontaneous.
discoverer: Meyer, 57f. discoverer: Bridges, 17i24.
references: 1958, DIS 32: 83. phenotype: Wings obliquely truncated but nearly
phenotype: Homozygous lethal and lethal in combi- full length; overlaps wild type at 25°C when
nation with dpo!S, dp 0 ' 1 ', and dplv. Slight oblique homozygous and in combination with dp o v . RK3.
74 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

dpo2 reduction in females and slight reduction in about


origin: Spontaneous. half the males. RK3.
discoverer: Laemmerts, 1926. *dp°h dumpy-oblique lethal
synonym: clip wing. origin: Spontaneous in dp°.
phenotype: Wings truncated and shortened as in discoverer: Muller, 1919.
dpov, but without vortex or comma effects. No synonym: dpL: dumpy-Lopped.
overlap with wild type as homozygote. Over- phenotype: Homozygous lethal. dpol/dpov shows
lapping frequent at 25°C in compound with dp°v. small body and legs and drastically reduced
Mutant effect more pronounced in females than in wings; thorax normal but occasional comma
males in dpo2/dpov; sexual dimorphism less ex- effects at 26°C or higher; no vortex effect. dp°l/+
treme in dp°2/dp°2. RK1. occasionally shows wing effect. RK2.
*dpoS0c dpotM; dumpy-oblique lethal of Meyer
origin: Spontaneous. origin: Ultraviolet induced.
discoverer: Sobels, 50c7. discoverer: Meyer, 51b.
synonym: dp5°c. synonym: dpLM: dumpy-Lopped of Meyer; dpTSlb:
references: Sobels, Boterenbrood, Faber, and Truncate*1*.
Oppenoorth, 1951, DIS 25: 76. references: 1952, DIS 26: 66.
phenotype: Manifests oblique wing only when Me phenotype: Homozygous lethal. Similar to dp°l but
present. Best penetrance at 25°C; less at 28°C has occasional slight vortex effect and good
and none at 16°C. Manifestation better in females comma effect in compound with dp°v. RK2.
than males. Tests of allelism with dpov have dp°IS; dumpy-oblique lethal of Schalet
been inconclusive. RK3. origin: Spontaneous.
dpoSU discoverer: Schalet, 1955.
origin: Ultraviolet induced. synonym: dpLS; dumpy-Lopped of Schalet; dpLSch.
discoverer: Byers and Meyer, 51e. references: Carlson and Schalet, 1956, DIS 30: 70.
references: Meyer and Edmondson, 1951, DIS 25: phenotype: Homozygous lethal. Heterozygote
72. dp°l$/dpov shows small body and legs and
phenotype: Slight wing effect in dp°v/dp°Sl«; no drastically reduced wings; thorax normal but
vortex effect. Wing effect in dp°Sle/dp°lv o n i y occasional comma effects at 26°C or higher; no
slightly greater. RK3. vortex effect. Lethal with dp°tv, dplv, and dp'M.
Jpobm; dumpy-oblique from bilateral mosaic RK2.
origin: X ray induced. dp°ly: dumpy-oblique lethal vortex
discoverer: Carlson, 1957. origin: Spontaneous.
references: Carlson and Southin, 1959, Genetics discoverer: Morgan, 1923.
44: 502-3. synonym: dpT: dumpy-Truncate.
phenotype: Inviable as homozygote, perhaps from references: Altenburg and Muller, 1920, Genetics
independent lethal nearby, because heterozygotes 5: 1-59 (fig.).
with dpoIv, dp°l, and dp1* are viable. Heterozy- phenotype: Homozygous lethal. dpolv/dpov has
gous dpov/dpobm has oblique wing, slightly reduced body and leg size; wings shortened to
reduced body size, and comma effects, but normal half length and obliquely truncated; thorax shows
dorsocentral region. RK2. two and sometimes four vortices; comma effect
other information: Arose in same fly as dpolvbm; strong; stemopleural bristles turned down; wings
dpolvhm/dpobm i8 virtually lethal. often elevated 45° and sometimes contain black
*dpobw: dumpy-oblique brown blisters. Body reduction also seen in compound
origin: Spontaneous, with dp°2 but not with dpv2. Lethal in compound
discoverer: Honer, 1939. with dp°ts, dptMt a n d dp/v, R K 2 .
synonym: dpbw- dtxmpy-brovm.
references: 1939, Z. Induktive Abstammungs- origin: Spontaneous.
Verebungglehre 77: SOI-15 (fig.). discoverer: Muller, 1913.
phenotype: Homoxygous females have moderately synonym: dpT2f x2.
truncated wings, but males only slightly affected. references: Altenburg and Muller, 1920, Genetics
dpQlvH/dpobw females have reduced body and leg 5: 1-59 (fig.).
size, but males almost normal; variable expression phenotype: Similar to dp°lvM. RK2.
in wing size. Some wings pointed in females. RK3. *dpotv54d
*dp»d*f: dumpy-oblique deformed origin: Ultraviolet induced,
origin: Spontaneous, discoverer: Meyer, 54d.
discoverer. Honer, 1939. synonym: dpT54dw
synonym: dp^®1; dumpy-deformed. references: 1955, DIS 29: 74.
references: 1939, Z. Induktive Abstammungs- phenotype: Like dp°'*. RK2.
Vererbungslehre 77: 501-15 (fig.), *dpolvSSb
phenofype: Males normal, female wings slightly origin: Spontaneous.
oblique. dp@de(/dp°lvH shows moderate wing discoverer: Meyer, 55b.
MUTATIONS 75

synonym: dpTSSb. c/po'vR; dumpy-oblique lethal vortex Ruffled


references: 1955, DIS 29: 74. origin: X ray induced.
phenotype; Similar to dpolv, RK2. discoverer: Schultz, 33a25.
*dpolv55c synonym: dp&t-. dumpy-Ruffled.
origin: Ultraviolet induced. phenotype: Similar to dpoly but heterozygous
discoverer: Meyer, 55c. dpolvR has slightly spread wings with uneven
synonym: dpTSSc. surface and ruffled margin. Raffling effect over-
references: 1955, DIS 29: 74. laps wild type at 25°C, RK2A.
phenotype: Like dpolv. RK2. cytology: Associated with In(2L)dpolvR =
*dpolvS7g In(2L)25A;25B3-4.
origin: Ultraviolet induced, dpoIvS; dumpy-oblique lethal vortex of Schalet
discoverer: Meyer, 57g. origin: Neutron induced.
synonym: dpr57g# discoverer: Schalet, 1955.
references: 1958, DIS 32: 78. synonym: dpTSch. dumpy-Truncate of Schalet.
phenotype: Like dpoJv. RK2. references: Carlson and Schalet, 1955, DIS 29: 71.
phenotype: Similar to dpoIv, RK2.
dpolvbm; dumpy-oblique lethal vortex from other information: Arose simultaneously with
bilateral mosaic mutation to y in Dp(l;2)sc1 9, but ed, cl, and
origin: X ray induced. dw-24F not affected; possibly a minute inversion.
discoverer: Carlson, 1957. *c/po'vSn: dumpy-oblique lethal vortex Snub
references: Carlson and Southin, 1959, Genetics origin: Spontaneous.
44: 502-3. discoverer: Bridges, 16bl7.
phenotype: Like dpotv. RK2. synonym: dpTs: dumpy-Truncate Snub.
other information: Recovered from same fly that references: Bridges and Morgan, 1919, Carnegie
produced dpobm. Inst. Wash. Publ. No. 278: 140.
*dpolvD: dumpy-oblique lethal vortex of Duncan phenotype: Similar to dp°lvM. RK2.
origin: Spontaneous. c/po/vW; dumpy-oblique lethal vortex of Williams
discoverer: Duncan, 1914. origin: Spontaneous.
synonym: dpTD: dumpy-Truncate of Duncan. discoverer: Williams, 1957.
references: 1915, Am. Naturalist 49: 575-82. synonym: dpTW: dumpy-Truncate of Williams.
phenotype: Similar to dp°'vM. RK2. references: Carlson, 1956, DIS 30: 70.
*cfpo/vH; dumpy-oblique lethal vortex of Hb'ner phenotype: Similar to dp°tv. RK2.
origin: Spontaneous, *dpou: dumpy-oblique ultraviolet
discoverer. Hb'ner, 1931. origin: Ultraviolet induced.
synonym: dpH: dumpy of Honer, discoverer: Meyer, 54g.
references: 1939, Z. Induktive Abstammungs-
references: 1954, DIS 28: 76.
Vererbungslehre 77: 501-15 (fig.). phenotype: Similar to dp°2. RK2.
1939, DIS 11: 45-46.
phenotype: Similar to dpolv. dp°IvH/+ may show
some wing truncation. RK2.
cytology: Salivary chromosomes normal.
*t/po/vM; dumpy-oblique lethal vortex of Morgan
origin: Spontaneous,
discoverer: Morgan, lOh.
synonym: dpTO. dumpy-Truncate zero.
references: Bridges and Morgan, 1919, Carnegie
Inst. Wash. Publ. No. 278: 136 (fig.).
Altenburg and Muller, 1920, Genetics 5: 1—59
(fig.).
phenotype: dp°lvM/+ showed oblique truncated
wings in up to 90 percent of flies in maximally
selected lines. RK2.
other information: Altenburg and Muller established
that temperature and modifier genes affect a major
gene at 2-13.0. Later tests with dpo*, dpy, dp°
and other alleles established similarity to dp°lv.
*e/po'vP; dumpy-oblique lethal vortex of Plough
dpov: dumpy-oblique vortex
origin: Recovered among progeny of temperature-
Edith M. Wallace, unpublished.
shocked parents.
discoverer: Plough, 40cl5.
synonym: dpTP', dumpy-Truncate of Plough.
t/pov; dumpy-oblique vortex
origin: Spontaneous.
phenotype: Like dpotr, RK2.
discoverer: Morgan, 18jl6.
76 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

synonym: dp. dpy: dumpy-vortex


references: Morgan, 1929, Carnegie Inst. Wash. origin: Spontaneous.
Publ. No. 399: 196. discoverer: Bridges, 1919.
phenotype: Wings reduced one-third, with oblique references: Bridges and Mohr, 1919, Genetics 4:
truncation; angle between veins L2 and L3 283-306 (fig.).
widened and intercrossvein distances shortened; Bridges and Morgan, 1923, Carnegie Inst. Wash.
marginal vein intact; two vortices with whorls of Publ. No. 327: 168 (fig.).
hairs in dorcentral region; dorsocentral bristles phenotype: Homozygote normal; expresses vortices
medially displaced; vortex expression in most only when e(dpv) on the third chromosome is also
flies at 26°C; comma effect in anterior thorax also homozygous. Heterozygotes with dpv2, dp°v,
present in most flies at 26°C. At lower tempera- dplv, and dpolv have good vortices without e(dpv).
tures, wing effect persists but thorax appears RK3.
normal. Body and leg size normal except at
temperatures above 28°C, when they are somewhat origin: Spontaneous.
reduced. RK1. discoverer: Mohr, 20a30.
dpov51f references: 1923, Studia Mendeliana (Brunae)
origin: Spontaneous, pp. 266-87.
discoverer: Mossige, 1951. 1929, Z. Induktive Abstammungs- Vererbungslehre
synonym: dpsli, 50: 113-200.
phenotype: Like dp°*. RK1. phenotype: Thorax shows two vortices in dorso-
Jpev52b central region, frequently pitted with brown
origin: Spontaneous. pigment. Whorls of hairs and medially displaced
discoverer: Edmonds on, 1952. dorsocentral bristles in almost all flies at 26°C.
synonym: dp^^b^ Females more extreme than males. No wing
references: 1952, DIS 26: 60. effect, no body or leg reduction in any compound.
phenotype: Like dp°*. RK1. Commas absent in dpv2/dpv2 and dpv2/dpov, but
present in dpvl/dp1* and dpv2/dp°lv. RK2.
dpovdr; dumpy-oblique vortex drumlins ; dumpy-vortex of Muller
origin: Spontaneous in dpov. origin: Spontaneous.
discoverer: R. C. King, 48b26. discoverer: Muller, 1920.
synonym: dp*; dumpy-dtumlins. synonym: dpvl; dumpy-volcano.
references: Poulson and King, 1948, DIS 22: 54. phenotype: Homozygote has vortices, but less well
phenotype: Has a more pronounced vortex effect developed and less numerous than in dpy2.
than dp°y but otherwise similar. Viability slightly Overlaps wild type. RK2.
reduced. RK1. dp**: dumpy-vortex of Vfaddington
other information: Probably a modifier in or near origin: Spontaneous.
the dumpy locus. discoverer: Waddington.
synonym: dpvo.
dpovN; dumpy-oblique vortex of Novitski phenotype: Similar to dpv2. RK2.
origin: X ray induced.
discoverer: Novitski. dpwh dumpy-warped
v
•ynonym: dpN° : dumpy of Novitski. origin: X ray induced.
phenotype: Like dpov. RK1. discoverer: Schalet, 1955.
dp^h see references: Carlson and Schalet, 1955, DIS 29: 71.
dpT; see Carlson, 1958, DIS 32: 117-18.
dpT0: see phenotype: Heterozygote of dpov/dpwl has variable
dpT2; see and asymmetrical expression of vortex and oblique
dpTsib: s e e wing effects. Mutant phenotype completely sup-
dpTS4d: pressed by an extra Y chromosome. Homozygous
dpT55b; lethal but viable in compound with lethal dp
see alleles. dpwl/dptvl have wrinkled opaque wings.
see dpotvsrg RK2A.
see d cytology: Associated with T(2;3)dpwl (breakpoints
not determined).
other information: Appears to be a variegated
dpi position effect.
c/pw2
see
dpTS: see origin: X ray induced.
discoverer: Schalet, 55k.
references: Carlson and Schalet, 1956, DIS 30: 71.
dp**: see c Carlson, 1958, DIS 32: 117-18.
dp**': see phenotype: Similar to dp*!. RK2A.
MUTATIONS 77

cytology: Associated with T(Y;2)dp*2 (break appearance; constant expression. Lethal homozy-
points not determined), gous but survives in combination with DrWe (E. B.
Dp: see Dr Lewis). RK1.
*dpy: dumpoidy other information: DrJAio/DrWe yields rare wild-type
locotion: 3- (right arm near 90). recombinants (Lewis).
origin: Spontaneous. DrWe: Drop-Wec/ge
discoverer: Villee, 40a. origin: Spontaneous.
phenotype: Wings obliquely truncated and reduced discoverer Muller.
in length; marginal vein intact. No vortices or references: 1965, DIS 40: 36.
whorls of bristles on thorax. Suppressed by Cy phenotype: Eyes are small vertical wedges with
and Gla; made dominant by Me, Overlaps wild points downward; much smaller than homozygous
type slightly. RK2. 23. Lethal homozygous and in combination with Dr,
d<j: see Ld(i but DtWe/DiMio survives (E. B. Lewis). RK1.
*dr: droopy other information: Dr^e/Dr^io yields rare wild-type
location: 2-71.2 (to the left of Lobe). recombinants (Lewis).
origin: Ultraviolet induced. drb: dark red brown
discoverer: Meyer and Edmondson, 49c. location: 3-47.7 (may be rearrangement; st—p
references: 1949, DIS 23: 60. crossing over 50 percent of normal),
phenotype: Wings spread wide apart and droop origin: Spontaneous.
downward, like c, often crumple and drag in the discoverer: Rosin, 48b.
food. Alula broad and short. Viability at hatching references: 1951, DIS 25: 75.
fair; females tend to die before males. Penetrance phenotype: Eye color dark red-brown at 18° and
100 percent. Fertility good. RK2. dark red at 28°C. drb/+ darker than wild type at
other information: Not a lie lie to c, 18°, but not at 28°C. RK2(A).
dt: see drw droop wings: see drp
Dr: Drop droopy: see dr
location: 3-99.2. droopy wing: see drw
origin: X ray induced. Drop: see Dr
discoverer: Krivshenko, 54c25. *drp: droop wings
synonym: Dp. location: 1- (rearrangement),
references: 1954, DIS 28: 75. origin: Spontaneous,
phenotype: Heterozygote has 1—10 eye facets, discoverer: Ives, 48f.
which appear dark red. Viability of heterozygote references: 1949, DIS 23: 58.
excellent. Homozygous lethal. RK1. cytology: Associated with In(l)drp - ln(l)12B;20B.
cytology: No visible rearrangement in salivary *drw: droopy wing
chromosomes, location: 1-52.3.
other information: Recombination between Dr*1*0 and origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
DrWe indicates that Dr is a pseudoallelic locus phenylalanine (CB. 3025).
(E. B. Lewis). discoverer: Fahmy, 1953.
*DrA: Drop of Abrahamson synonym: Symbol originally dr, which was
origin: X ray induced. preoccupied.
discoverer: Abrahamson, 60d28. references: 1958, DIS 32: 69.
synonym: DpA. phenotype: Small fly with drooping wings. Chitin
references: Abrahamson and Siegel, 1960, DIS 34: of abdomen irregularly ridged and pigmented.
48. Hairs deranged. Males infertile; viability 10 per-
phenotype: Facet number of heterozygote averages cent wild type. RK3.
four. Homozygote viable and usually lacks eye ds: dachsous
facets. RK1. location: 2-0.3.
*DrL: Drop of Lewis origin: Spontaneous,
origin: X ray induced, discoverer: Bridges, 17kl2.
discoverer. E. B. Lewis. references: Stern and Bridges, 1926, Genetics 11:
phenotype: Like Dr. RK1A. 511.
cytology: Associated with T(2;3)DrL = Mohr, 1929, Z. Induktive Abstammungs-
T(2;3)44;89F-90A + In(3R)89C;95D-96Bl, which is Vererbungslehre 50: 113-200 (fig.),
probably independent of the mutation. phenotype: Wings shorter, blunter, and broader,
>: Drop-Microp/ifhalmia with crossveins uniformly very close together.
origin: Mustard gas induced, Abdomen and legs chunky. Slight dominance of
discoverer: Sobels, 57J22. close crossveins. Strong interaction with d, ij,
synonym: Mlo. and eg; double homozygotes often have excessive
references: 1958, DIS 32: 84. growth of thoracic parts and sometimes conversion
phenotype: Dr^io/+ show extreme reduction in eye of one organ into another [e.g., twinning of wings
size; facets coalesce to give a shiny, dark red or antennae or conversion of eyes into antennae,
78 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

Waddington, 1943, J. Genet., 45: 44-50 (fig.)]. cytology: Associated with In(2LR)bwvl =
Tarsal shortening enhanced by homozygous s s a and In(2LR)21C8-Dl;60Dl-2 + ln(2LR)40F;59D4-El.
ssaB (Villee, 1945, Genetics 30: 26-27). RK1. other information: ds33k has not been separated
cytology: Analysis by E. B. Lewis (1945, Genetics from the recessive lethal factor of In(2LR)bwVl,
30: 137-66) indicates that ds is located in 21D1-2 and so has not been observed in homozygous
or possibly slightly to the left in the last band of condition.
21C.
origin: Spontaneous.
discoverer: Waddington, 38k.
references: Curry, 1939, DIS 12: 45.
phenotype: ds38k/ds has close crossveins; fly
short and thick bodied. Homozygote probably like
dsd. RK2.

origin: Spontaneous,
discoverer: T. Hinton and Bliven, 41b.
references: Hinton, 1942, DIS 16: 48.
phenotype: Wings shorter and extremely blunt.
Less extreme and more varied in male. Viability
and fertility good. RK2.
cytology: Salivary chromosomes normal.

origin: Ultraviolet induced.


discoverer: Meyer and Byers, 51a.
references: Meyer and Edmonds on, 1951, DIS 25: 72.
phenotype: Like dsd. RK3.
*dsd: dachsous-dachsoid
origin: Spontaneous.
discoverer: Sturtevant, 17b9.
ds: dachsous references: Bridges and Morgan, 1919, Carnegie
Edith M. Wallace, unpublished. tost. Wash. Publ. No. 278: 294 (fig.).
phenotype: Wings broader but half normal length,
almost round and widely spread; crossveins
origin: Spontaneous. broken, very close together. Hairs erect on costal
discoverer: Bridges, 25d2. vein near base. Body small and foreshortened.
references: Stern and Bridges, 1926, Genetics 11: Legs stumpy. Viability about 40 percent wild
513. type. RK3.
phenotype: Crossveins closer together, but only *Jsd53h
slight shortening of legs, wings, and abdomen. origin: Ultraviolet induced.
Fully viable and fertile. RK1. discoverer: Meyer, 53h.
s3 references: 1953, DIS 27: 58.
origin: Spontaneous, phenotype: Wings rounded; crossveins close. Legs
discoverer: Bridges, 25k5. have thickened coxae. Viability low; females
references: Stern and Bridges, 1926, Genetics 11: sterile. RK3.
513. ds*: dachsous-Wide
phenotype: Crossveins extremely close, but wings, origin: Spontaneous.
legs, and abdomen shortened only slightly. Via- discoverer Bridges, 29d24.
bility low (40 percent wild type), and females synonym: da2*d.
completely sterile. Males fertile. Emergence references: 1935, DIS 3: 10.
delayed. RK3. phenotype: Heterozygote has uniformly widely
spaced posterior scute liar bristles; wings a bit
see ds* short and blunt; crossveins close together. Via-
ds33k bility and separability excellent. Homozygote
origin: X ray induced at same time as resembles ds rf and dtsd3 with viability 40 percent
ln(2LR}bwVl. wild type. RK1 as heterozygous dominant; RK3
discoverer: Bridges, 33k28. as homozygote.
references: 1935, DIS 4: 7. dsh: dishevelled
j*h#n©ty|H>: dt$33k/+ has phenotype like dsw/+; location: 1-33.5.
s cute liar bristles wide apart, wings shorter and origin: Induced by methyl methanesulfonate
wider than normal, and crossveins close together. (CB. 1540).
d@33k/dsW n^g extreme phenotype like d«d. discoverer: Fahmy, 1956.
RK3A. references: 1959, DIS 33: 85.
MUTATIONS 79

phenotype: Thoracic hairs deranged. One or more has sex combs; however, on basitarsus of forelegs
hairs abnormally curved. Wings usually divergent of each, bristles of last transverse row are
and blistered. Eyes ellipsoid, with some deranged enlarged and rotated toward area comb would
ommatidia. Males viable and fertile; females occupy if present. Internally, the X/X intersex
sterile. RK2. usually has male and female reproductive parts
other information: One allele induced by CB. 3026. with varying degrees of completeness. Well-
developed ovaries present in some. Frequently a
single gonad is attached to both male and female
systems. The X/Y intersex usually has a predom-
inantly male internal reproductive system, but
occasionally both a male and a female system are
present. In all cases, gonads are poorly de-
veloped. RK2.
cytology: Salivary chromosomes normal.

origin: Spontaneous.
discoverer: Puro, 601.
dsr: disrupted synonym: Originally called ix-3: intersex on
From Bridges and Brehme, 1944, Carnegie Inst. Wash. chromosome 3.
Publ. No. 552: 7 1 . references: 1964, DIS 39: 64.
phenotype: Like dsx, X/X and X/Y flies have
dsr: disrupted external characteristics of both sexes and are
location: 2-90. similar except for a slight difference in size. No
origin: Spontaneous. sex combs. Last abdominal segments form rather
discoverer: Curry, 38a28. well-developed but rotated male-like terminalia
phenotype: Wings have plexus of extra and doubled with female-like structures of the seventh and
veins at anterior and posterior cross veins and at eighth tergites. Rudimentary gonads usually a
L3 and L4. L3 and L4 spread wide apart. Wing mass of undifferentiated tissue. RK2.
slightly wider and warped. At 25°, overlaps wild cytology: Salivary chromosomes normal.
type; at 19°C, no overlap but viability reduced to other information: Not allelic to tra and no inter-
60 percent wild type. RK3. action with ix in doubly heterozygous compounds.
*dss: disturbed segmentation *Dt: Detached
location: 1-27.3. location: 2-10.
origin: Spontaneous. discoverer: Bridges, 17ell.
discoverer: Fahmy, 1954. references: Morgan, Bridges, and Sturtevant, 1925,
references: 1959, DIS 33: 85. Bibliog. Genet. 2: 224.
phenotype: Extremely abnormal abdomen with phenotype: Vein L2 fails to reach margin in 60
segmentation grossly deformed, very few hairs, percent of flies. Homozygote not known. RK3.
and disturbed pigmentation. Occasionally some other information: Bridges considered this a
bristles shortened. Eyes reduced in size and possible effect of 5 or requiring S as an enhancer
sometimes abnormal in shape. Males fertile; as it was found in a S stock and apparently was
viability about 10 percent wild type. Females never separated from S.
sterile. RK3. *dta: delta wing
dsx: double sex location: 1- (rearrangement).
location: 3-48.1 (to the right of pP). origin: Induced by triethylenemelamine (CB. 1246).
origin: Recovered among progeny of flies fed discoverer: Fahmy, 1953.
tritiated thymidine. references: 1958, DIS 32: 69.
discoverer: Hildreth and Lucchesi, 62c. phenotype: Wings widely outspread, frequently
synonym: Originally called ix62c: intersex-62c, drooping in homozygous female. Viability good
but name and symbol preoccupied. but female sterile. RK2A.
references: 1963, Proc. Intern. Congr. Genet., 11th, cytology: Associated with ln(l)dta =
Vol. 1: 171. In(l)6B2-3;15E7-F2.
Hildreth, 1965, Genetics 51: 659-78 (fig.)- dtv: see thv**
phenotype: Chromosomal females (X/X) and males *du; dunk el
(X/Y) transformed into intersexes. The two types location: 3-47.
are similar. Pigmentation of tergites similar to origin: Spontaneous,
wild-type males. Like females, they have seventh discoverer: Hadorn, 49el5.
tergite with seventh spiracle at its base. Ventral references: Hadorn and Fritz, 1950, Arch. Julius
part of eighth abdominal segment has protuberance Klaus-Stift. Vererbungsforach. Sozialanthropol.
similar to female gonopod; ninth segment bears Rassenhyg. 25: 504-8.
claspers like males. Anal plates situated verti- phenotype: Body color dark, sootylike. Wings
cally as in males. Neither X/X nor X/Y ixttersex blistered. Viability almost normal at 25°, greatly
80 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

reduced at 18°C. Males fertile; females sterile. references: 1936, Biol. Zh. (Moscow) 5: 537—40
Ovaries and eggs normal size and morphology. (fig.).
Insemination of females normal (motile sperm in 1937, DIS 8: 81.
spermathecae and receptaculum). Either eggs from phenotype: Has shorter, darker wings; post-
du females not fertilized or zygotes die before scutellars bowed in; body size small; sterility
blastoderm formation, du ovaries behave auton- high; semilethal. In combination with yellow-
omously as implants in normal hosts, and wild- bodied y alleles, gives strongly curled wings with
type ovaries are fertile in du hosts. RK2 at 25°C. slight outward twist. With f, gives crumpled
other information: Not an allele of by or cu. wings. With sc, almost lethal. RK3.
cytology: Salivary chromosome studies by Demerec
dumpoidy: see dpy and Sutton show locus to lie at right of 8D8-9
dumpy: see dp (Demerec, Kaufmann, Fano, Sutton, and Sansome,
dunkel: see du 1942, Carnegie Inst. Wash. Year Book 41: 191).
dusky: see dy
dusky body: see dyb origin: Spontaneous.
discoverer: Curry, 37kl7.
phenotype: Practically wild type. With y2, wings
tightly curled; with y, wings spirally curled. RK2
with y.
*dvrs: divers-subliminal
origin: Spontaneous.
discoverer: Muller.
synonym: dvri*).
references: 1946, DIS 20: 67.
phenotype: Wild type either alone, heterozygous to
dvr-2, or in combination with y. y dvra/y dvt%, on
the other hand, has wings distinctly curly or
wavy, usually as in typical Cy, but other effects
noted in dvr flies not evident. RK3.
*dvw: divergent wings
location: 1-13.3.
dv: divergent origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
From Bridges and Morgan, 1923, Carnegie Inst. Wash. phenylalanine (CB. 3026).
Publ. No. 327: 182. discoverer: Fahmy, 1953.
references: 1959, DIS 33: 85.
*dv: divergent phenotype: Sex-limited character. Males late
location: 3-20.0, hatching; wings divergent, occasionally upheld,
origin: Spontaneous, with inner margins frequently cut away to various
discoverer: Bridges, 17fl3. degrees. Bristles short and stiff. Homozygous
references: Bridges and Morgan, 1923, Carnegie females normal. RK1 in males.
Inst. Wash. Publ. No. 327: 182 (fig.). *dw: dwarf
Morgan, Bridges, and Sturtevant, 1925, Bibliog. location: 3-50.
Genet. 2: 58 (fig.). origin: Spontaneous.
Mohr, 1937, DIS 8: 12. discoverer: Bridges, 13kl2.
phenotype: Wings spread, smaller and have slight references: Bridges and Morgan, 1923, Carnegie
venation disturbances. Both sexes rather in- Inst. Wash. Publ. No. 327: 101.
fertile. dv/Dt(3L)Vn progeny of homozygous dv Morgan, Bridges, and Sturtevant, 1925, Bibliog.
mothers practically lethal although the same geno- Genet. 2: 58 (fig.).
type from other crosses survives (Mohr and phenotype: Body weight 76 percent that of hetero-
Mossige, 1943, Avhandl. Norske Videnskaps-Akad. zygous sibs. Females usually sterile (3 of 63
Oslo, I. Mat.-Natur. Kl. No. 7: 1-51). RK2. gave a few offspring). RK3.
cytology: Salivary chromosome locus placed be- dw-24F: dwarf in salivary chromosome section
tween 64C12 and 65E1, on basis of its inclusion in 24F
Df(3L)Vn = Df(3L)64C12~Dl;65D2-El (Mohr, 1938, location: 2-13.
Avhandl. Norske Videnskaps-Akad. Oslo, I. origin: Spontaneous.
Mat.-Natur. Kl. No. 4: 1-7). discoverer: Curry, 39k.
references: 1941, DIS 14: 49.
dvr: divers phenotype: Body small; abdomen narrow and mis-
location: 1-28.1 (located using a W ) . shapen. Body surface dull if not properly dried.
origin: Recovered among progeny of iodine-treated Eyes dull in color and smallish. Wings close
male. textured, small, tend to droop; crossveins close.
discoverer: Sacharov, 1932. Bristles slender. Low viability and fertility. RK3.
MUTATIONS

cytology: Located between 24E2 and 25A2 based on origin: Induced by 2-chlorethyl methanesulfonate
its inclusion in Df(2L)M-zB = Df(2L)24E2- (CB. 1506).
Fl;25Al-2 (Morgan, Bridges, Schultz, and Curry, discoverer: Fahmy, 1956.
1939, Carnegie Inst. Wash. Year Book 38: 273-77). references: 1959, DIS 33: 85.
*dw-b: dwarf-b phenotype: Extremely inviable dwarf; wings
location: 3-12. frequently fail to expand completely. Males fertile
origin: Spontaneous. if they survive to breed. RK3.
discoverer: Bridges, 20b5. dwx: dwarfex
references: Bridges and Morgan, 1923, Carnegie location: 1-33.2.
Inst. Wash. Publ. No. 327: 182, 228, 231 (fig.). discoverer: Bridges, 33c31.
phenotype: Flies about 70 percent as heavy as phenotype: Body small. Wing texture coarse;
wild type. RK3. marginal hairs slightly disarranged. Classifica-
tion sometimes difficult. RK3.
*dw-sc: dwarf with scute *dwxm n: dwarfex-manikin
location: 1-0.7. origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
origin: Spontaneous (arose with sc and separated). phenylalanine (CB. 3026).
discoverer: Bridges, 16a22. discoverer: Fahmy, 1954.
phenotype: Small body. Viability erratic. RK3. synonym: mn.
dwarf unexpanded: see dwu references: 1959, DIS 33: 88.
dwarfex: see dwx phenotype: Fly small with narrow abdomen. Reduc-
dwarfish: see dwh tion in size may be bilaterally asymmetrical and
dwarfoid: see dwf affect abdomen and thorax independently. Male
dwarp: see dwp viability reduced; flies rarely survive more than 48
*dwf: dwarfoid hr. Sterile, probably owing to reduced vigor. RK3.
location: 1-13.3. other information: One X-ray-induced allele.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino- dx: deltex
phenylalanine (CB. 3025). location: 1-17.0.
discoverer: Fahmy, 1955. origin: Spontaneous.
references: 1959, DIS 33: 85. discoverer: Bridges, 22h26.
phenotype: Flies small. Males fertile; viability references: Morgan, Bridges, and Schultz, 1931,
about 50 percent wild type. Homozygous females Carnegie Inst. Wash. Year Book 30: 410.
show extreme expression; fertility and viability phenotype: Veins show thickenings and terminal
low. RK2. deltas, resembling Dl in third chromosome, but
dwg: deformed wings fully viable, fertile, and easily classified. Nearly
location: 1-1.6. suppressed by su(dx), Su(dx), and Su(dx)2. RK2.
origin: Induced by DL-p-NN-di(2-chloroethyl)amino- cytology: Demerec and Sutton show locus to be
phenylalanine (CB. 3007). between 6A3-4 and 6F10-11 (Demerec, Kaufmann,
discoverer: Fahmy, 1954. Fano, Sutton, and Sansome, 1942, Carnegie Inst.
references: 1959, DIS 33: 85. Wash. Year Book 41: 191).
phenotype: Wings broad, round tipped, and have
occasional marginal incisions; sometimes grossly dxst: deltex-sterile
deformed in shape and venation. Extremely fine origin: Spontaneous change of dx to dxst.
bristles. Eyes small and occasionally rough. discoverer: Bridges, 31a3.
Males late in eclosion; viable but sterile. RK2. phenotype: Veins heavy, confluent, and dilated at
dwh: dwarfish junctions; strong deltas at tips. Wings spread
location: 3- (not located), wide; margins and tips snipped and nicked. Ocelli
origin: Spontaneous. sometimes fused, with disturbance of hairs and
discoverer: Bridges, 30dl6. bristles in the region. Acrostichals irregular.
phenotype: Small body. Wings disproportionately Male sterile. Less abnormal phenotype and fertile
broad; eyes irregularly knobby and somewhat dull with Su(dx). RK2.
in color; legs weak and slightly crippled. RK3. dy: dusky
*dwp: dwarp location: 1-36.2 (to the right of m).
location: 1-0. origin: Spontaneous.
origin: Spontaneous, discoverer: Bridges, 1611.
discoverer: Bridges, 23b20. references: Morgan, Bridges, and Sturtevant, 1925,
references: Morgan, Bridges, and Sturtevant, 1925, Bibliog. Genet. 2: 35 (fig.), 224.
Bibliog. Genet. 2: 225. Slatis and Willermet, 1954, Genetics 39: 45-58.
phenotype: Dwarf with warped wings. Body color Dora and Burdick, 1962, Genetics 47: 503-18.
pale. Nearly always lethal, but survivors fertile. phenotype: Wings smaller than normal but of nearly
RK3. wild-type shape, dusky in color. Cell expansion
*dwu; dwarf unexpanded inhibited in prepupal as well as pupal period
location: 1-58.3. (Waddington, 1940, J. Genet. 41: 75-139). RK1.
82 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

cytology: Demerec and Sutton showed that locus phenotype: Similar to other ay's. Fully fertile in
lies just to right of 10E1-2 (Demerec, Kaufmann, both sexes. Shows high complementation with
Fano, Sutton, and Sansome, 1942, Carnegie Inst. m60 and low with dy61a. RK1.
Wash. Year Book 41: 191). dy61a
other information: No reverse mutations among origin: Gamma ray induced.
2 X 10 s progeny of progeny of homozygous females discoverer: Ives, 61a24.
(Pullar). A member of the m-dy pseudoallelic references: 1961, DIS 35: 46.
complex. Dorn and Burdick, 1962, Genetics 47: 503-18.
phenotype: Like dy. Good fertility and fecundity in
both sexes. RK1.
dy62b
origin: Spontaneous,
discoverer: Burdick, 62b.
references: 1963, DIS 37: 47.
phenotype: Similar to other dy's. RK1.
*dy286-9
origin: X ray induced.
discoverer: Demerec, 35b.
phenotype: Lethal in male and cell lethal. RK2.
cytology: Salivary chromosomes normal.
other information: rn + and fw+.

*dyala: dusky-alae
origin: Recovered among progeny of heat-treated
flies.
discoverer: Gottschewski, 34c.
synonym: a/a; ala parvae.
phenotype: Wings about 90 percent as large as dy.
dy: dusky RK1.
Edith M. Wallace, unpublished. *dyb: dusky body
location: 1-44.6.
*dy2 origin: Induced by ethyl methanesulfonate
origin: Spontaneous. (CB. 1528).
discoverer: Bridges, 20a3. discoverer: Fahmy, 1958.
references: Morgan, Bridges, and Sturtevant, 1925, references: 1959, DIS 33: 85.
Bibliog. Genet. 2: 224. phenotype: Dusky body color and browner eyes.
phenotype: Wings not quite so small as dy. RK1. Eye and wing shapes slightly altered. Males
*dy3 viable and fertile; females sterile. RK2.
discoverer: Bridges, 22a2.
references: Morgan, Bridges, and Sturtevant, 1925, e; ebony
Bibliog. Genet. 2: 224. location: 3-70.7.
phenotype: Wings slightly larger than dy. Clean origin: Spontaneous,
separation difficult in females. RK2. discoverer: E. M. Wallace, 12bl5.
*dy31d references: Bridges and Morgan, 1923, Carnegie
origin: X ray induced. Inst. Wash. Publ. No. 327: 50 (fig.).
discoverer: Oliver, 1931. phenotype: Body color shining black. Puparia much
synonym: m-like: miniature-like. lighter than wild type. Classifiable throughout
phenotype: Like dy. RK1. larval period by darkened color of spiracle sheaths
dySSk (Brehme, 1941, Proc. Natl. Acad. Sci. U.S. 27:
origin: Spontaneous. 254—61). Viability lowered to about 80 percent
discoverer: Krawinkel, 58k. wild type. Heterozygote has slightly darker body
references: Burdick, 1961, DIS 35: 45. color than normal. For interaction with other body
Dorn and Burdick, 1962, Genetics 47: 503-18. color mutants, see Waddington, 1941, Proc. Zool.
phenotype: Wings shorter than ay, resembling the Soc. London, Ser. A 111: 173-180. Virtually un-
longer-winged m's. Fertile in both sexes. Shows able to incorporate C 1 4 from labeled beta-alanine
more complementation with tn'» than ay. RK1. into puparium or hardening adult integument,
other information: Recotnbines with all m's except whereas normal flies incorporate heavily; e/+ in-
mD but not with ay. corporates intermediate amounts Qacobs and
dy60k Brubaker, 1963, Science 139: 1282-83; Jacobs,
origin: Spontaneous, 1966, Genetics 53: 777-84). RK1.
discoverer Burdick, 60k. cytology: Placed in salivary chromosome region be-
references: 1961, DCS 35: 45. tween 93B7 and 93F9, on the basis of its inclusion
MUTATIONS 83

in both Dt(3R)el 00.1 72 = Df(3R)93B7-10;93F10- phenotype: Like es when heterozygous with e».
94A1 and Df(3R)el 00.256 = Df(3R)93A5-Bl;93F5-9 Homozygous lethal. RK2A.
(Ward and Alexander, 1957, Genetics 42: 42—54). cytology: Associated with Df(3R)el00.256 -
Df(3R)93A5-Bl;93F5-9.
origin: Spontaneous. *e100.265
discoverer: Sturtevant, 17i27. origin: X ray induced.
references: Bridges and Morgan, 1923, Carnegie discoverer: Alexander.
Inst. Wash. Publ. No. 327: 184. references: Ward and Alexander, 1957, Genetics 42:
phenotype: Body color very dark like e, but 42-54.
abdomen lighter. Most viable and generally best phenotype: Like e. RK1A.
of the dark alleles, such as e and ell. c l a s s i - cytology: Associated with In(3R)e*00.265 =
fiable from hatching throughout larval period by In(3R)93B5-6;95E.
darker spiracle sheaths (Brehme, 1941, Proc. *e100.307
Natl. Acad. Sci. U.S. 27: 254-61). RK1. origin: X ray induced.
*e4.39 discoverer: Alexander.
origin: X ray induced. references: Ward and Alexander, 1957, Genetics 42:
discoverer: Alexander. 42-54.
references: 1960, Genetics 45: 1019-22. phenotype: Like es when heterozygous with e«.
phenotype: Like es when heterozygous with e«. Homozygous lethal. RK2.
Homozygous lethal. RK2A. cytology: Induced simultaneously with
cytology: Associated with Df(3R)e4-39 = In(3L)100.307 = In(3L)62E2-4;64C2-4.
Di(3R)93B;93F. other information: Not shown whether lethality
associated with e or the inversion.
origin: Spontaneous. *e300.96
discoverer: Stern, 25a. origin: X ray induced.
references: 1926, Z. Induktive Abstammungs- discoverer: Alexander.
Vererbungslehre 41: 198-215. references: Ward and Alexander, 1957, Genetics 42:
1934, DIS 1: 35. 42-54.
phenotype: Dark allele similar to e and e4. phenotype: Like e but male sterile. RK2(A).
Tyrosinase formed in adults (Horowitz). Classi- cytology: Induced simultaneously with
fiable throughout larval period by dark spiracle In(3R)300.96 = In(3R)89F2-90Al;99B2-4 but prob-
sheaths (Brehme, 1941, Proc. Natl. Acad. Sci. ably separable from it.
U.S. 27: 254—61). Phenylthiocarbamide inhibits e*: ebony-sooty
development of ell homozygotes more than wild origin: Spontaneous.
type; reverse is true for inhibition by silver discoverer: Sturtevant, 13i20.
chloride; heterozygotes intermediate in both references: Bridges and Morgan, 1923, Carnegie
cases. Mixtures of the two inhibitors affect heter- Inst. Wash. Publ. No. 327: 99 (fig.).
ozygotes to a greater extent, thus exhibiting phenotype: Body color darker than wild type but
"negative heterosis" (Kroman and Parsons, 1960, lighter than e. Viability excellent. Most easily
Nature 186: 411—12), Under ordinary conditions, classified as ea/e4. Classifiable from hatching
viability and fecundity of heterozygote superior to throughout larval period by dark color of spiracle
either homozygote (Moree). RK1.
sheaths, although darkening is less than in e, e4,
eiitti: see tu-bwe
or ell (Brehme, 1941, Proc. Natl. Acad. Sci. U.S.
e60h
27: 254—61). Ability to incorporate beta-alanine-
origin: Spontaneous, derived C intermediate between that of e and +
discoverer: Ives, 60h. (Jacobs and Brubaker, 1963, Science 139: 1282-83;
references: 1965, DIS 40: 35. Jacobs, 1966, Genetics 53: 777-84). RK1.
phenotype: Medium e. RK1. *e**: ebony-striped
* e 700.I72 origin: Spontaneous.
origin: X ray induced. discoverer: Villee, 39k.
discoverer: Alexander. references: 1941, DIS 14: 40.
references: Ward and Alexander, 1957, Genetics 42: 1942, Univ. Calif. (Berkeley) Publ. Zool. 49: 137.
42-54. phenotype: Body color similar to es but with
phenotype: Like e* when heterozygous with e*. definite longitudinal stripes on mesonotum.
a at
Homozygous lethal. RK2A. eat/ea resembles e ; e /e resembles e. Viability
cytology: Associated with Dl(3R)el 00.172 m excellent. RK2.
Df(3R)93B7-10;93F10-94Al. ; ebony-Uganda
*9100.2S6 origin: Spontaneous.
origin: X ray induced. discoverer: Zurcher, 1956.
discoverer Alexander. references: 1958, Arch. Julius Klaus-Stift.
references: Ward and Alexander, 1957, Genetics 42: Vererbungsforsch. Sozialanthropol.
42-54. Rassenhyg. 33: 75.
84 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

1960, DIS 34: 112. e(dpv): enhancer of dumpy-vortex


1963, Genetics 34: 1-33 (fig.). location: 3-40.4.
phenotype: A light ebony allele. Females have origin: Spontaneous.
dark, extremely wide marginal bands on abdominal discoverer: Bridges, 16h7.
tergites; entire abdominal tergite region darkly synonym: vo-3: vortex in chromosome 3.
shaded. Males only slightly darker than wild type. references: 1919, Bridges and Mohr, Genetics 4:
Viability lower than wild type. RK2. 283-306 (fig.).
E(B): Enhancer of Bar 1923, Bridges and Morgan, Carnegie Inst. Wash.
location: 1-57.3. Publ. No. 327: 168.
origin: Spontaneous. 1925, Morgan, Bridges, and Sturtevant, 1925,
discoverer: Bonnier and Nordenskibld. Bibliog. Genet. 2: 41-43 (fig.).
synonym: i; I: Intensifier of Bar; Eb: Exaggeration phenotype: Normal. In combination with dpv
of Bar. produces one or two pairs of pits or volcanolike
references: 1942, DIS 16: 47. protrusions on thorax; hairs and bristles arranged
Bonnier, Nordenskiold, and Bagman, 1943, in whorls. RK3.
Hereditas 29: 113-33 (fig.). *E(f): Enhancer of forked
Rasmuson, 1948, Proc. Intern. Congr. Genet., 8th. location: 2-86.5.
pp. 645—46. origin: X ray induced,
phenotype: E(B) heterozygous with any B allele, discoverer: Belgovsky, 37c4.
including 23+, produces flies similar in phenotype synonym: I-t: Intensifier of forked.
to homozygotes for that allele. B +/+ E(B) eyes references: 1937, DIS 8: 7.
have 80—90 facets but B E(B)/++ eyes have only 1938, Izv. Akad. Nauk SSSR, Ser. Biol. 1017-36.
40. Homozygous lethal. RK2(A). 1940, DIS 13: 52.
cytology: Salivary chromosomes appear normal, but 1944, Zh. Obshch. Biol. 5: 325-56.
there is occasional indication of deficiency for phenotype: Homozygote has short, twisted bristles
faint bands 16A5 and 6. intermediate between f and Bl; postcutellars often
other information: Reduces B-fu crossing over about pale; viability and fertility reduced. Heterozygote
40 percent. is wild type. f/+; E(f)/+ slightly more extreme than
e(bx): enhancer of bifhorax f. f/f; E(f)/+ has an extreme forked phenotype and
location: 1-1.0. hairs are forked, f/f; E(f)/E(f) rarely survives.
origin: Gamma ray induced. RK3.
discoverer: E. B. Lewis, 53b. cytology: Salivary chromosomes normal.
synonym: en-bx. *e(g): enhancer of garnet
references: 1959, DIS 33: 96. location: 1-5.9.
phenotype: Recessive enhancer of most bithorax discoverer: Payne and Denny, 1921.
alleles. By itself, the only abnormal phenotype is synonym: m(g): modifier of garnet.
a slight variegated eye color. Most readily references: 1921, Am. Naturalist 55: 377-81.
classified in combination with bx34e +/+ Ubx. phenotype: Apparently wild type, but in combination
Enhances development of dorsal metanotum with g produces a more orange eye than g alone.
of latter from a few hairs to a broad band of hairy RK3.
tissue somewhat like that seen in homozygous *E(H): Enhancer of Hairless
fox3*8. Enhances bx3 +/+ pbx from wild type to a location: 2-50.5.
slight postbithorax phenotype (slight wing-like discoverer: Nash.
modification of posterior portion of halteres). references: 1963, Proc. Intern. Congr. Genet., 11th.
RK2A. Vol. 1: 174-75.
cytology. Associated with In(l}e(bx) = In(l)3A;4F. Nash, 1965, Genet. Res. 6: 175-89.
Locus within Dp(l;l)w = Dp(l;l)3A;3C, probably phenotype: E(H); H flies nearly devoid of chaetae.
in section 3A. Majority of bristle sites are vestiges. RK2.
e(bx)2 *E(M3g): Enhancer of Minute(3) g
origin: X ray induced, location: 3- (near spindle attachment).
discoverer: E. B. Lewis, 55h, origin: Spontaneous.
synonym: m2~bx. discoverer: Bridges.
references: 1959, DIS 33: 96. phenotype: Specific intensifier of shortness of
phenotype: Similar to e(hx). RK2. bristles of M(3)g. RK3.
cytology: Salivary chromosomes normal. *e(N*): enhancer of Notch-8
E(bx): Enhancer of biihorax location: 3- (not located).
location: 3- (to the left of R). origin: Spontaneous.
origin: X ray induced. discoverer: Mohr, 181.
discoverer. E. B. Lewis. references: 1923, Z. Induktive Abstammungs-
synonym: En-bx. Vererbungslehre 32: 108-232 (fig.).
ph*notyp«: Enhances expression of bx34m, bx3, and phenotype: Produces slight nicking of wings.
Ubx/'*. Lethal homozygous. RK2. Enhances Ns. RK3.
MUTATIONS 85

*e(S): enhancer of Star Variegated position effects do not respond uni-


location: 3- (between 0 and 10; perhaps an allele of formly to E(var)7. RK2(A).
ru or R). cytology: May be small abnormality in 25A (Schultz).
origin: Spontaneous. E(w°): Enhancer of white-apricot
discoverer: Bridges, 16kl8. location: 2- (not located).
synonym: S-i: intensifier of Star. origin: Spontaneous.
references: Bridges and Morgan, 1923, Carnegie discoverer: Scandlyn.
Inst. Wash. Publ. No. 327: 175 (fig.), phenotype: Heterozygote dilutes wB to pale yellow.
phenotype: By itself, homozygous e(S) has normal Homozygote with wa is white. Not tested with
eyes. S/+; e(S)/e(S) has eyes smaller and rougher other white alleles. No effect on eye color in
than S/+, although overlapping somewhat; abdomen presence of w+. Homozygous sterile. RK3.
bulbous, body color darkish. RK3. *efw«); enhancer of white-eosin
E(S): Enhancer of Star location: 1-32.
location: 2-6 [claimed to lie between left break of origin: Spontaneous.
In(2L)Cy and locus of Cy\. discoverer: Green, 55b21.
discoverer: Bridges, 30a27. synonym: en-we.
phenotype: E(S)/+ normal, E(S)/B(S) gives slight references: 1957, DIS 31: 81.
roughening of eye. B(S)/+ strongly reduces size 1959, Heredity 13: 303-15.
and increases roughness of S/+ and S2/+ eyes; phenotype: Enhances intermediate alleles at the
imparts dominance to ast/+, ast%/+, ast3/+, and fourth recombinational site of the w locus, e.g.,
a s ^ / + ( L e w i s , 1945, Genetics 30: 137-66). we, we2, WBE, wh, and wXi6 to produce nearly
S +/+ E(S) occasionally emerges as a late-eclosing white eye color. No enhancement of tested inter-
giant. RK3A. mediate alleles at other w subloci, e.g., wa, w"2)
cytology: Arose in In(2L)Cy = wa3, w*"t, wbi, wch2, wco, wcoi, or wsat> Also
In(2L)22Dl-2;33F5-34Al. suppresses f. e(we) flies occasionally have px-
E(sd): Enhancer of scalloped like venation, or shortened wings, or both. Homo-
location: Autosomal. zygous females sterile. RK2.
origin: Spontaneous. eagle: see eg
discoverer: R. M. Valencia, 1963. eb: ebonized
references: 1965, DIS 40: 37. location: 1-42.0.
phenotype: Almost completely removes wings of origin: Induced by ethyl methanesulfonate
sdBP; not tested with other alleles of sd. No inter- (CB. 1528).
action with Bx or BxT. RK2. discoverer: Fahmy, 1956.
E(spl): Enhancer of split references: 1959, DIS 33: 86.
location: 3- (near ro). phenotype: Heavily pigmented fly, with trident
origin: Spontaneous. pattern and scutellum very dark. Wings slightly
discoverer: Green. shorter; membrane often slightly concave; wing
synonym: En-spl. tips occasionally truncate. Males viable and
references: Welshons, 1956, DIS 30: 157-58. fertile; females sterile. RK2.
Von Halle, 1965, DIS 40: 60. Eb: seeE(B)
phenotype: Both E(spl)/+ and E(spl)/E(spl) cause ebonized: see eb
spl/+ to resemble spl/spl and cause spl/spl and ebony: see e
spt/Y to have an extreme phenotype. ec: echinus
spl/+/Dp(l;2)51b; E(spl)/+ less extreme than location: 1-5.5.
spl/+; E(spl)/+. E(spl) homozygote viable and origin: Spontaneous,
fertile. RK2. discoverer: Bridges, 1516.
e(tu-K): enhancer of tumor K phenotype: Eyes large and bulging. Eye surface
location: 3- (not located). rough; facets large. Wings rather short and broad.
origin: Spontaneous. Body thickset. Tends to remove dorsocentrals
discoverer Bur net and Sang. (posterior more than anterior) and posterior
references: 1964, Genetics 49: 223—35. notopleurals; may also add dorsocentrals anterior
phenotype: Homozygote produces a significant in- to anterior dorsocentrals whether or not posterior
crease in the penetrance of tu-K in both untreated bristles removed (Sturtevant). ec is visible in
flies and those treated in ways known to increase +/ec/ec triploids (Gersh). RK1.
tumor incidence in tu-K. RK3. cytology: Locus placed at 3F1-2 by Demerec and
E(var)7: Enhancer of variegation Sutton (Demerec, Kaufraann, Fano, Sutton, and
location: 2- (not located). Sansome, 1942, Carnegie Inst. Wash. Year Book
origin: X ray induced. 41: 191).
discoverer: Schultz. echinoid: see ed
phenotype: E(ver)7/+ has no phenotype of its own echinus: see ec
but enhances variegation, e.g., w**>* is made much ed: echinoid
lighter and variegation for tmt appears in males. locotion: 2-11.0.
86 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

origin: Spontaneous. el: elbow


discoverer: Bridges, 31al6. location: 2-50.0.
phenotype: Eyes large and rough. Easily classified, origin: Spontaneous.
although not so extreme as ec. RK1. discoverer: E. M. Wallace, 35dl.
cytology: Placed between 24D2 and 24F1 on the phenotype: Wings extended and bent backward (as
basis of its inclusion in Df(2L)M'Zc = Df(2L)24D2- in bt and bat), often warped, shortened; sometimes
5;25A2-3 but not in Df(2L)M-zB = Df(2L)24E2- blistered or nicked. Alula always reduced and
Fl;25Al-2 (Morgan, Bridges, Schultz, and Curry, fused with main wing blade. Venation reduced by
1939, Carnegie Inst. Wash. Year Book 38: 273-77). terminal shortening of L5 and of crossveins.
*e/: elfin Balancers reduced, especially in third segment.
location: 1- (rearrangement). Eye size decreased, varying with other manifesta-
origin: Induced by triethylenemelamine (CB. 1246). tions. RK2.
discoverer: Fahmy, 1952. cytology: Placed in region between 34E5 and 35D1
references: 1959, DIS 33: 86. on the basis of its inclusion in Df(2L)64j =
phenotype: Small fly with slightly excess melaniza- Df(2L)34E5-Fl;35C3-Dl (E. H. Grell).
tion. Wings proportionally smaller, slightly altered elfin: see ef
in shape, and warped. Abdominal tergites often Ellipse: see Elp
broken and abnormally pigmented. Males viable elliptical rough: see e/r
but sterile. RK3A. Elp: Ellipse
cytology: Associated with T(l;2)ef = location: 2-99 (to the left of px).
T(1;2)14C8-D1;2R. origin: Spontaneous.
eg: eagle discoverer: E. H. Grell, 57b.
location: 3-47.3. references: 1960, DIS 34: 50.
discoverer: Morgan, 1930. phenotype: Eyes of heterozygotes rough and more
phenotype: Wings extended. Hairs on thorax some- oval than wild type. Homozygotes have small
what disarranged. Dark pattern on thorax. RK2. eyes with fused facets; resemble homozygous ast.
eg' Homozygotes have low viability. RK1 as hetero-
discoverer: Bridges, 33J16. zygote.
synonym: dh; dihedral. *e/r; elliptical rough
phenotype: Wings extended and uplifted. Females location: 1-25.1.
often sterile. RK2. origin: X ray induced.
eg 57c discoverer: Fahmy, 1956.
origin: Spontaneous. references: 1960, DIS 34: 49.
discoverer: Nicoletti, 57c. phenotype: Eyes slightly elliptical and rough.
references: 1957, DIS 31: 84. Wings slightly broader. Both sexes viable and
phenotype: Wings held out at 45-90° angle from fertile. RK2.
midline. Excellent viability and fertility. RK1. other information: Two other alleles: one induced
by X rays, one by CB. 3025.
en: engrailed
location: 2-62.0.
origin: Spontaneous.
discoverer: Evang, 26k7.
references: Eker, 1929, Hereditas 12: 217—22
(fig.).
Bridges, Skoog, and Li, 1936, Genetics 21: 788—
95.
Brasted, 1941, Genetics 26: 347-73 (fig.).
phenotype: Longitudinal cleft extends from rear
border of scutellum forward; may be reduced to
median nick or posterior flattening of scutellum.
Bristles often javelin- or hooked-like. Wings
larger, broader, and thin textured, with spatulate
end; venation always disturbed (like that of cf),
with gap in L4 and L5 and branching plexus of
extra veins. In males, extra sex comb often
present, smaller than normal and in mirror-image
position on outer side of tarsus. Action of en on
differentiation of secondary sex comb on male
foreleg is autonomous LTokunaga, 1961, Genetics
el: elbow 46: 157—76 (fig.)J. Male genitalia may be malformed
From Bridges and Brehme, 1944, Carnegie inst. Wash. and rotated, resulting in sterility (Curry, 1941,
Pub!. No. 552: 75. DIS 14: 50). eg en/+ + characterized by slight
MUTATIONS 87

degree of L4 interruption and thinning in males at may be segmented and have hairs. Less extreme
low temperature. The triple heterozygote expression produces derangement of central or
eg en/+ +; ci/+ has about 50 percent expression anterior-central facets. Eruption may occur as
of L4 interruption (House, 1961, Genetics 46: encroachment of chitin with bristles and hairs into
871). For other interactions of en with ci alleles anterior edge of eye. RK2.
andfl, see House (1953, Genetics 38: 199-215, other information: Alleles of at least five different
309-27). RK1. strengths present in different wild stocks.
Present in many wild stocks in suppressed
condition.
*Er: Erect
location: 3-50.
origin: Spontaneous.
discoverer: Neel, 41c9.
references: 1942, DIS 16: 50.
phenotype: Posterior scutellars at greater than
normal angle with body; vary from slight effect to
condition in which bristles stand at right angles
to scute Hum. In latter case, bristles usually
appear warped and twisted. Wings incompletely
expanded and crinkled to varying degrees. RK3.
erupt: see er
es: ether sensitive
location: 2- (not located).
origin: Spontaneous.
discoverer: Tinderholt.
references: Kidd, 1963, DIS 37: 49.
phenotype: Hypersensitive to diethyl ether and
chloroform. Homozygotes killed by exposure to
doses of these agents harmless to normal flies.
en; engrailed Sensitivity probably affected by modifiers. A male
From Eker, 1929, Hereditas 12: 217-22. sterility factor seems to be associated but may be
separable. Viability of homozygote about 70 per-
en-; see e( ) cent that of es/SMl and remains low in strains
En-: see E( ) selected for less sensitivity. Not sensitive to
engrailed: see en carbon dioxide. RK3.
enhancer: see e( ) esc; extra sex combs
Enhancer: see E( ) location: 2-54.9 (based on location of escD).
eq: equational producer origin: Spontaneous.
location: 1- (to the right of car — probably hetero- discoverer: Slifer, 40e2.
chromatic). references: 1942, J. Exptl. Zool. 90: 31-40 (fig.).
origin: X ray induced, phenotype: Sex combs may be present on all six legs
discoverer: Schultz, 33a2. of male; at least one extra sex comb present in
references: Morgan, Bridges, and Schultz, 1934, majority of males. Expression affected by culture
Carnegie Inst. Wash. Year Book 33: 280. conditions. When expressivity high, extra trans-
phenotype: Produces 1—2 percent equational non- verse bristle rows appear between sixth and eighth
disjunction of X's in male, producing both X/X longitudinal rows of bristles, mainly on distal por-
and nullo-X, nullo-Y sperm. Original eq male tion of basitarsus and tibia of second and third
when crossed to attached-X female produced legs in both sexes; accompanied by shortening of
89/289 equational exceptional X/X daughters. affected leg segments. Sex comb development
Claimed to generate 66-deficient Y chromosomes. autonomous in mosaics produced by somatic cross-
RK3. ing over [Tokunaga and Stern, 1965, Develop. Biol.
cytology: Both salivary and mitotic chromosomes 11: 50—81 (fig.)]. For interactions with Pc and
appear normal. Sex see Hannah-Alavah, 1958, Genetics 43: 878—
er: erupt 905 (fig.). Males and females sterile. RK2.
location: 3-70.7 (60.7 to 80.7; not an allele of k). cytology: Arose in chromosome carrying In(2L)t »
origin: Spontaneous. In(2L)22D3-El;34A8'9, but mutant and inversion
discoverer: Glass, 1941. separable (Tokunaga).
references: 1943, DXS 17: 50. esc 2 : see escD
1944, Genetics 29: 436-46. *esc°: extra sex combs-Dominant
1957, Science 126: 683-89 (fig.), origin: Spontaneous,
phenotype: Exhibits eruption of underlying hypo- discoverer: Stromnaes, 53f.
dermis in center of one or both eye*. Eruption synonym: ©sc-2; Esc2.
88 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: Hannah and Stromnaes, 1955, DIS 29: system. Est-CF/Est-CF produces a rapidly
121-23. migrating esterase C and Est-CF/Est-Cs produces
phenotype: esc&/+ similar to esc/esc. Homozygous two bands on starch gel; no evidence for hybrid
lethal. RK2. enzyme in heterozygotes. Est-Cs Est-6S/Est-CF
Est-6F: Esterase 6-Fast Est-6F homogenates produce four electrophoretic
location: 3-36.8 (10.3 units to the right of h and 6.4 bands in place of the two found in homozygotes.
units to the left of th). RK3.
origin: Naturally occurring allele. Est-Cs: Esterase C-S/ow
discoverer: T. Wright, 61h. origin: Naturally occurring allele.
references: 1963, DIS 37: 53. discoverer: Beckman and Johnson.
1963, Genetics 48: 787-801 (fig.). references: 1964, Hereditas 51: 212-20 (fig.).
phenotype: Esterase 6 is one of ten positively phenotype: Produces esterase C with slow electro-
migrating esterases demonstrable histochemically phoretic mobility on starch gel. RK3.
with (X-naphthyl acetate and Fast Blue BB after ether sensitive: see es
starch gel electrophoresis of imaginal homoge-
nates. Readily identified in zymograms as the
most heavily staining esterase. Migrates about
5 cm when exposed to a voltage drop of 2.0—2.5 v
per cm for 17 hr at 4°C in a starch gel prepared
with 0.05 M tris buffer, pH 8.7. Flies homozygous
for Est-6F have a single esterase 6 band that
migrates faster than a similar, single esterase 6
band found in flies homozygous for Est-6S. The
enzyme specified by Est-6F is completely inac-
tivated by exposure to 60°C for 10 min (Wright,
1964, DIS 39: 60). Heterozygotes, Est-6F/Est-6S,
exhibit both the fast and slow esterase 6 bands and
do not contain a hybrid esterase 6 with an interme-
diate electrophoretic mobility. No morphological
difference between Est-6F and Eat-6S homozygotes
is apparent. Both have been found together in wild
populations and in numerous wild type and mutant
stocks. Esterase 6 of Wright corresponds to es-
terase D of Beckman and Johnson. RK3.
ex; expanded
discoverer: Maclntyre, 63d. From Stern and Bridges, 1926, Genetics 11: 503-30.
references: Wright, 1964, DIS 39: 60.
phenotype: Produces an esterase 6 with the same ex; expanded
electrophoretic mobility as that produced by location: 2-0.1.
Est-6F but that is stable to treatment with 60°C origin: Spontaneous.
for 10 min. RK3. discoverer: Bridges, 17k21.
Est-6S: Esterase 6-Slow references: Stem and Bridges, 1926, Genetics 11:
origin: Naturally occurring allele. 514 (fig.).
discoverer: T. Wright. phenotype: Wings extremely wide and large, some-
references: 1963, DIS 37: 53. times curved and divergent. Effect produced in
1963, Genetics 48: 787-801 (fig.). prepupal wing, probably by influence on cell
1964, DIS 39: 60. division (Waddington, 1940, J. Genet. 41: 75-139).
phenotype: Produces slowly migrating esterase 6 Eyes slightly reduced in size and roughish. Body
that is stable to treatment with 60°C for 10 min. large. RK.2.
RK3. cytology: Salivary chromosome location in or near
Est-C?: Esterase C-Fast 21C3 (Lewis, 1945, Genetics 30: 137-66).
location: 3-49 (based on 2/68 crossovers with Aph
and 9/43 with Est-6). origin: Recovered among progeny of flies treated
origin: Naturally occurring allele. as larvae with natural gas.
discoverer: Beckman and Johnson. discoverer. Mickey, 48k.
reference*: 1964, Hereditas 51: 212-20 (fig.). references: 1950, DIS 24: 60.
phenotype: Esterase C is one of six postively phenotype: Slightly more extreme than ex. RK2.
migrating esterases that can be demonstrated with Exaggeration of Bat: see E(B)
a-aapiithyl acetate and Fast Blue RR after starch exi: exiguous
gel electrophoresis of single fly homogenates for location: 1-51.5,
3 hr at room temperature at a voltage gradient of origin: Induced by 2-choroethyl methaneaulfonate
6—8 v per cm, using Poulik's discontinuous buffer (CB. 1506).
MUTATIONS 89

discoverer: Fahmy, 1956. 1946, DIS 20: 88) and its presence on Df(4)ll =
references: 1958, DIS 32: 70. Df(4)102E2-10;102F2-10 (Hochman).
phenotype: Small fly with rather dusky body color.
ey-
Not easily classified. Viability and fertility good origin: Spontaneous.
in male, slightly reduced in female. RK3. discoverer: Nonidez, 1919.
expanded: see ex references: Bridges, 1935, Biol. Zh. (Moscow) 4:
*exr: extra venation 401-20 (fig.).
location: 1- (associated with In(l)exr). phenotype: Eye reduced to one-half to one-fourth
origin: Induced by triethylenemelamine (CB. 1246). wild-type area, with relatively little variability.
discoverer: Fahmy, 1952. Cephalic complex smaller than wild type at 25 hr
references: 1958, DIS 32: 70. after hatching from egg at 27°C; subsequent
phenotype: Eyes slightly rough and smaller than growth rate same as wild type [Medvedev, 1935,
normal. Wings have irregularly distributed extra Z. Induktive Abstammungs- Vererbungslehre 70:
vein tissue. Males viable and fertile; females 55-72 (fig.); 1935, Tr. Inst. Genet. Akad. Nauk
viable but sterile. RK3A. SSSR 10: 119-51; Steinberg, 1944, Proc. Natl.
cytology: Associated with In(l)exr = Acad. Sci. U.S. 30: 5-13]. RK1.
In(l)l 2E8-10; 15D1 -3. other information: Most often used ey allele.
*ext: extended
location: 2- (not located),
origin: Spontaneous,
discoverer: Stroher, 1958.
references: Mainx, 1958, DIS 32: 82.
phenotype: Wings held out at about a 75° angle from
body axis, are wavy and gradually curve down-
ward. Distal parts of wings often crumpled and
folded. Halteres normal. Function of wings ey4; eyeless-4
reduced. Viability and fertility good. RK3. Edith M. Wallace, unpublished.
Ext: Extras
location: 1-15.2.
discoverer Schultz, 3318. origin: Spontaneous.
phenotype: Heterozygous female has thickened, discoverer: Li, 25fl0.
branched, and extra veins. Overlaps wild type. references: Bridges, 1935, Biol. Zh. (Moscow)
Lethal in male. RK3. 4: 401-20.
ext-b: see bat phenotype: Eye size and variability intermediate
Ext-sct-3: see Su(sc) between ey and ey2. Optic disk of mature larvae
extended: see exf about 20 percent size of wild-type disk (Chen,
extended-b: see bat 1929, J. Morphol. 47: 135—99). Expression and
Extra sex comb: see Sex fertility of both males and females highly suscep-
extra sex combs: see esc tible to modification of genetic background
extra venation: see exr (Spofford, 1956, Genetics 41: 938-59). RK2.
Extras: see Exf *ev34g
ey; eyeless origin: Occurred among progeny of cold-treated fly.
location: 4-2.0 (located in diplo-4 triploids by discoverer: Gottchewski, 34g26.
Sturtevant, 1951, Proc. Natl. Acad. Sci. U.S. 37: references: 1935, DIS 4: 15.
405-7). phenotype: Strong allele of ey, like ey2. RK1.
origin: Spontaneous.
discoverer: Hoge, 14e. origin: Spontaneous.
references: 1915, Am. Naturalist 49: 47—49. discoverer: Spencer, 36e25.
Bridges, 1935, Biol. 2h. (Moscow) 4: 401-20 (fig.), references: 1937, DIS 7: 8.
phenotype: Eye reduced to three-fourths to one-half phenotype: Like ey, expression variable; viability
normal area, but varies from no eyes to extensive poor. RK3.
overlapping of wild type. Less extreme at low *eY46l
temperatures. Optic ganglia reduced (Richards origin: X ray induced.
and Furrow, 1922, Proc. Oklahoma Acad. Sci. 2: discoverer: King and Poulson, 461.
41—45). Variability in size of eye affected by references: 1948, DIS 22: 54.
environmental and genetic conditions (Morgan, phenotype: Eye smaller than ey2, conical in pro-
1929, Carnegie fast. Wash. Publ. No. 399: file, may be notched or partly covered with
139-68; Baron, 1935, J. Exptl. Zool. 70: cuticle. Sometimes bristle-covered palps protrude
461-90). RK2. from border of eye. Palps often found on bucca.
cytology: Placed between 1O2C15 and 102E10 on Often dark smudges on gena. Extra vibrissae and
basis of the absence of ey + from the 2LD4^P ele- buccal bristles. Anterior verticals usually
ment of T(2;4)h~ T(2;4)25E;102C15'Dl (Morgan, doubled; posterior verticals thickened, shortened,
90 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

and bent or split at tips. ey46i/ey2 intermediate eyK; eyeless from Oregon-K
between the two homozygotes. Viability fair. origin: Spontaneous.
RK2. discoverer: Sang and McDonald.
*eyAD: eyeless of A. Das references: 1954, J. Genet. 52: 392-412 (fig.).
origin: Spontaneous. Sang and Burnet, 1963, Genetics 48: 1683—1700.
discoverer: Das, 63a7. phenotype: Eyes reduced in varying degrees from
references: Sarkar, 1963, DIS 38: 28. eyelessness to wild type. Supernumerary antennae
phenotype: Bye varies from absence to normal. occur in a small proportion of flies, especially in
RK2. flies with no eyes. Eye size depends on diet.
RK2.
eyopt: eyeless-ophthalmoptera
origin: Spontaneous derivative of ey2. On chromo-
some 4; possibly separable from ey2.
discoverer: E. Goldschmidt.
references: Goldschmidt and Lederman-Klein, 1958,
J. Heredity 49: 262-66 (fig.).
phenotype: Eyes small with variable outgrowths.
RK2.
ey R : eyeless-Russian
origin: Spontaneous.
discoverer: Sacharov, 23hl3.
references: Serebrovsky and Sacharov, 1925, Zh.
eyD: eyeless-Dominant Eksperim. Biol. 1: 75—91.
Left: head. Right: first pair of legs. Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20.
From Patterson and Muller, 1930, Genetics 15: 495- phenotype: Eye reduced to one-half or one-fourth
577. normal area, reduction occurring chiefly in
anterior part. Similar to ey2. RK1.
eyD; eyeless-Dominanf *eyfu; eyeless-fumorous
origin: X ray induced. origin: Spontaneous,
discoverer: Muller, 27k. discoverer. Datta, 63c26.
references: Patterson and Muller, 1930, Genetics synonym: tu-h^^: tumor head-63.
IS: 495-577 (fig.). references: Sarkar, 1963, DIS 38: 28.
Bridges, 1935, Biol. Zh. (Moscow) 4: 401-20. phenotype: Abnormal growths in various regions of
1935, Tr. Dinam. Razvit 10: 463-73. the head; protuberances on or near eyes; small
phenotype: Eyes small, outline irregular, displaced eyes; aggregations of hairs in antennal region.
toward top and rear. Head large, often with dupli- RK2.
cated antennae or ocelli. Sex combs always very other information: Allelism shown by Fahmy.
large. Second joint of tarsi shortened to give *eyW59: eyeless of Wfc/fe
lumps, sometimes very conspicuous. Fully origin: Spontaneous.
dominant in triplo-4 flies (Sturtevant, 1936, discoverer: White, 59f.
Genetics 21: 448). Eye size of B; ey-*V+ males references: Meyer, 1959, DIS 33: 97.
larger than of B alone. Produces extreme pheno- phenotype: Eyes small or absent. More extreme at
type in combination with D, JD/+; eyD/+ almost higher temperatures. Good viability and fertility.
completely lethal (Sobels, Kruijt, and Spronk, RK1.
1951, DIS 25: 128). Homozygous lethal. Homo- *Eye: Eyeless dominant in chromosome 2
zygotes die during larval period and many eyD/+ location: 2-62.7.
flies die as pupae (Hochman). RK2. origin: Probably ultraviolet induced.
cytology: Salivary chromosomes show duplication discoverer: Edmonds on, 51g.
of about a dozen bands inserted into middle of synonym: ey-HD.
fourth chromosome as a reversed repeat. Source references: 1952, DIS 26: 60.
of duplication unknown (Bridges, 1935). phenotype: Eyes may be greatly reduced in size,
other information: May not be an allele of ey. with frequent doubling of antennae. Overlaps wild
*6yD39k type, especially in old vials. Recessive in
origin: X ray induced. triploids. Eye/+'f &yD/+ has smaller eyes than
discoverer: Suttoo, 39k. either alone. Homozygous lethal. RK3.
references: Hinton, 1940, DIS 13: 49. eye gone: see eyg
phenotype: Eyes appear as a knot or two separated eyeless: see ey
knots. Homozygote overlap* wild type in 75 per- Eyeless dominant in chromosome 2: see Eye
cent of flies. Heterozygote overlaps wild type in Eyeluf. see Eyl
only 50 percent of flies (T. Hinton, 1942, Am. eyes reduced', see eyr
Naturalist 76: 219-22). RK3. eyg: eye gone
cytology: Salivary chromosomes normal. location: 3-35.5.
MUTATIONS 91

origin: Spontaneous. affected, but this is visible only at high magnifi-


discoverer: Ives, 40g20. cations. Treatment with methylurea causes normal
references: 1942, DIS 16: 48. bristle formation (De Marinis). Developmental
phenotype: Eyes and head much smaller than studies [Lees and Waddington, 1942, Proc. Roy.
normal. Considerable pupal mortality, probably Soc. (London) Ser. B 131: 87-110 (fig.); Lees and
from inability to push open pupa cases. Adults Picken, 1945, Proc. Roy. Soc. (London), Ser. B
normal in viability and productivity. Character 132: 396—423 (fig.)] show nature of pupal bristle
subject to genetic modifiers and possibly environ- secretion is affected. Suppressed by su(f). RK1.
mental influences. Expression varies from com- cytology: According to Demerec and Sutton, bands
plete absence of facets to formation of about 100 15F1-5 are involved (Demerec, Kaufmann, Fano,
facets. RK2. Sutton, and Sansome, 1942, Carnegie Inst. Wash.
*Eyl: Eyeluf Year Book 41: 191). This can be narrowed to
location: 1-18. 15F1-3 on the basis of Df(l)f2S7-S -
origin: Spontaneous. Dt(l)l 5E7-F1.-15F2-4.
discoverer: Marzluf. other information: Green (1955, Proc. Natl. Acad.
phenotype: One or both eyes reduced in size. Sci. U.S. 41: 375-79; 1956, Proc. Natl. Acad.
Expression varies from slight reduction to Sci. U.S. 42: 73—77) showed the forked mutants
absence of eye. Sometimes extraneous materials can be assigned to either of two pseudoallelic
protrude through eye; frequently one or more series, f is a member of the right series. Back
duplicated antennae present. Penetrance incom- mutations to f+ occur spontaneously and their
plete; viability good. In aged and crowded cul- incidence is not increased by X rays (Green,
tures, both penetrance and expressivity increased. 1959, Proc. Natl. Acad. Sci. U.S. 45: 16-18;
Third chromosome carries important modifiers Lefevre and Green, 1959, Genetics 44: 769—76).
affecting penetrance, and different wild-type and
mutant stocks carry different modifiers. Pene-
trance lower at 18°C than at 25°C. RK3.
eyr: eyes reduced
location: 3-103.
origin: Found among flies grown on food containing
copper sulfate.
discoverer: Edwards and Gardner, 1962.
references: 1963, DIS 37: 47.
1966, Genetics 53: 785-98.
phenotype: Eyes vary from normal to absence of
ommatidia. Shows some degree of dominance;
many heterozygotes have some eye abnormality,
usually a nick in anterior region of one or both
eyes; an abnormal growth of wing tissue may be
associated with the nick, eyr; ey4 flies have very
small heads, usually without ommatidia. Viability
greatly reduced. RK2.

X4 XI
X3 h h forked
X2 36 a Edith M. Wallace, unpublished.
3N 5
3 I
l I
origin: Spontaneous,
discoverer: Bridges, 19kl4.
Map of the / locus references: 1938, DIS 9: 46.
Drawn from Green, 1956, Proc. Natl. Acad. Sci. U.S. Green, 1955, Proc. Natl. Acad. Sci. U.S. 41:
42: 73-77. 375-79.
phenotype: Slight allele of /. Ends of bristles
f: forked twisted or split. Not suppressed by su(f). RK2.
location: 1-56.7. other information: A member of the left pseudo-
origin: Spontaneous. allelic series.
discoverer: Bridges, 12kl9. f3N
references: Morgan and Bridges, 1916, Carnegie origin: Spontaneous derivative of /.
Inst. Wash, Publ. No. 237: 58 (fig.), discoverer: Green.
phenotype: Bristles shortened, gnarled, and bent, references: 1955, Proc. Natl. Acad. Sci. U.S. 41:
with ends split or sharply bent. Hairs similarly 375-79.
92 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

1959, Proc. Natl. Acad. Sci. U.S. 45: 16-18. cytology: Associated with In(l)i257"4 =
Lefevre and Green, 1959, Genetics 44: 769—76. In(l)15F2-16Al;16D2-El.
phenotype: Expression similar to /. Unlike f, does f2S7-5
not respond to su(f). RK1. origin: X ray induced.
other information: At the left f sublocus (Green). discoverer: Demerec, 33k.
Reverts spontaneously, and X rays delivered to phenotype: Lethal in male and cell lethal. RK2A.
oogonia and oocytes cause high incidence of cytology: Associated with Dt(l)f257-5 -
reversion. Irradiation of postmeiotic stages in the Df(l)15E7-Fl;15F2-4 (Sutton).
male does not cause back mutation (Green 1959; *{2S7-6
Lefevre and Green 1959). orjgin: Spontaneous; recovered originally as a Bar
{5 reversion.
origin: Spontaneous. discoverer: Bridges, 14i25.
discoverer: Bridges, 21b. references: 1917, Genetics 2: 445—65.
references: 1938, DIS 9: 47. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
phenotype: Bristles and hairs much curled and Genet. 2: 164, 226.
twisted. A rather extreme allele of f. Suppressed Sutton, 1943, Genetics 28: 99.
by su(f). RK1. phenotype: Lethal and cell lethal. RK2A.
other information: A member of the right f pseudo- cytology: Associated with Dt(l)P57-6 =Df(l)15E4-
allelic series (Green, 1955). F1;16A7-1.
*f34b other information: Identity of Dt(l)f2*7-6 w i t h the
origin: X ray induced. Bridges f B deficiency claimed in the original
discoverer: Stone, 34b. edition, but this does not seem well established.
references: 1935, DIS 4: 63. Bridges's case was the first recorded deficiency;
phenotype: Subliminal allele of f. Wild type in in 1925 it was claimed to be lost. Di(l)f257-6
males and homozygous females, but gives a weak obtained by Demerec from Pasadena in 1933 as a
forked phenotype when heterozygous with / or f5. t B deficiency.
RK3. *f2S7-9
*(34» origin: X ray induced; recovered with simultaneous
origin: X ray induced. reversion of BiB*.
discoverer Oliver, 34e4. discoverer: Demerec, 34c.
references: 1939, DIS 12: 48. phenotype: Lethal in male and cell lethal. RK2A.
phenotype: Like f. RK1. cytology: Associated with Df(l)f2S7-9 =
{36a Df(l)l 5E7-Fl;16D2-4.
discoverer: Ives, 36a27. *f2S7-J5
phenotype: The most extreme f allele. Hairs and origin: X ray induced.
bristles extremely crooked. RK1. discoverer: Demerec, 35a.
other Information: A member of the right / pseudo- phenotype: Lethal. RK2A.
allelic series (Green). cytology: Associated with T(l;2)f257mls =
*{42 T(l;2)13E9-10;15E2-3;24F (Sutton).
origin: Spontaneous,
discoverer: Anderson, 42c30. /257-I9
references: Oliver, 1942, DIS 16: 53. origin: X ray induced.
phenotype: An extreme allele like is. RK1. discoverer: Hoover, 35h.
*fSla phenotype: Lethal, extreme forked. RK2.
origin: X ray induced. cytology: No major band missing (Kaufmann).
discoverer: Green, 51a. *f2S7-22
references: Lefevre and Green, 1959, Genetics 44: origin: X ray induced in BiB* X chromosome.
769-76. discoverer: Demerec, 36c.
phenotype: An extreme f like f36a. Not suppressed phenotype: Lethal. RK2A.
by au{f). RK1. cytology: Associated with T(l;2)t2^7~22 -
other information: Not observed to revert sponta- T(1;2)4D2-3;8F;15E4-F1;39E;41F-42A (Kaufmann).
neously; reversions not induced by X rays. *f2S7-24
/56« origin: X ray induced.
origin: Spontaneous. discoverer: Demerec, 36e.
discoverer; Williams, 56e. phenotype: An extreme /. Lethal. RK2.
references: 1956, DIS 30: 79. cytology: No major band missing (Kaufmann).
phenotype: Like i. RK1. *{257-27
origin: X ray induced.
origin: X ray induced. discoverer: Demerec, 381.
discoverer: Demerec, 33j. phenotype: Lethal- RK2A.
phwitotyp*: Bristles and hairs strongly forked. cytology: Associated with Df(l)l357-27 - Df(l)14F6-
Viability and fertility good. RKIA. 15Al;l5FS-6 (Sutton).
MUTATIONS 93

*f257-28 fB27
origin: X ray induced. origin: X ray induced in BM2 male.
discoverer: Sutton, 40h. discoverer: Belgovsky, 361.
phenotype: Lethal. RK2A. references: 1940, DIS 13: 47.
cytology: Associated with Df(l)f257-28 = Df(l)15E7- phenotype: Males have mostly normal bristles; a
F1;16E5-F1 (Sutton). few reduced like a Minute; rarely forked. fB27/f
*/257-29 are mosaic for forked. fB27/fB27 females rarely
origin: X ray induced. survive, those that do sometimes have reduced
discoverer: Bishop, 401. bristles or notched wings or both, and are sterile.
phenotype: Very slight f effect. Male sterile. RK2A. More extreme in combination with E(f). RK3A.
cytology: Associated with T(l;3)f257-2 9 -
T(1:3)15F5-16A1;64 (Sutton). * / « : forked of Hexter
*{257-3 0 origin: Spontaneous.
origin: X ray induced. discoverer: Hexter.
discoverer: Bishop, 41a. synonym: f*3.
phenotype: Forked bristles; viability and fertility references: Green, 1956, Proc. Natl. Acad. Sci.
good. RK1. U.S. 42: 73-77.
cytology: Salivary chromosomes apparently normal phenotype: Like f but not suppressed by su(f). RK1.
(Sutton). other informotion: A member of the right f sublocus.
*f2S7-31 /*"; see f+ih
origin: X ray induced. fa*: see fx
discoverer: Bishop, 41a.
phenotype: Lethal. RK2A. fx: forked from X irradiation
cytology: Associated with Df(l)f257-31 =Df(l)15E7- origin: X ray induced, simultaneously with f+il*.
F1.-J5F5-6 (Sutton). synonym: f"; forked-mottled = fx / + ^ .
discoverer: Muller.
f+//»; forked-wild type in heterochromatin references: 1946, DIS 20: 88-89.
origin: X ray induced simultaneously with fx. 1947, DIS 21: 71.
synonym: P": forked-mottled = fx f+if>. Muller and Oster, 1957, DIS 31: 141-44.
discoverer: Muller. Oster, Ehrlich, and Muller, 1958, DIS 32: 144-45.
references: 1946, DIS 20: 88-89. phenotype: A medium /• Suppressed by su(f). RK1.
1947, DIS 21: 71. cytology: Salivary chromosomes appear normal
Muller and Oster, 1957, DIS 31: 141-44. (J. I. Valencia),
Oster, Ehrlich, and Muller, 1958, DIS 32: 144-45. other information: Located to right of f3-. Does not
phenotype: f +ih with any f allele has normalizing cross over with /. Is not induced to revert by X
effect. Patches of bristles and occasionally whole rays. See last two references for relation between
fly is wild type. An extra Y chromosome enhances fx and
the normalizing effect. RK2A.
cytology: Salivary chromosomes appear normal *fX7
Q. I. Valencia). origin: X ray induced.
other information: Apparently, £**** is all or part of discoverer: Green.
the normal allele of f transposed to the proximal references: 1956, Proc. Natl. Acad. Sci. U.S. 42:
heterochromatin of the X chromosome, where it 73-77.
variegates. phenotype: Like f but not suppressed by su(f). RK1.
*fB; forked of Belgorsky other information: In right f sublocus.
origin: X ray induced in In(l)scs or Jn(l)BM2. *fX2
discoverer: Belgovsky, 1936-1937. origin: X ray induced.
references: 1937, DIS 8: 7. discoverer: Green.
1938, Izv. Akad. Nauk SSSR, Ser. Biol., 1017-36. references: 1956, Proc. Natl. Acad. Sci. U.S. 42:
1940, DIS 13: 47-48. 73-77.
other information: A series of several dozen / phenotype: Like f but not suppressed by su(f). RK1.
alleles of different strengths. Analyzed genet- other information: In left f sublocus.
ically but not cytologically. +fX3
fBIS origin: X ray induced.
origin: X ray induced in B ^ 2 male, discoverer: Green.
discoverer Belgovsky, 361. references: 1956, Proc. Natl. Acad. Sci. U.S. 42:
references: 1940, DES 13: 47. 73-77.
phenotype: Shows variegated expression of f. More phenotype: Like /but not suppressed by su(f). RK1.
extreme in combination with E(f). RK2A. other information: In left sublocus of forked.
cytology: G#netic data indicate that the mutation *fX4
is associated with a reinversion of the BM3 origin: X ray induced.
inversion. BMS phenotype reverted. discoverer Green.
94 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: 1956, Proc. Natl. Acad. Sci. U.S. 42: references: Muller and Altenburg, 1921, Anat.
73-77. Record 20: 213.
phenotype: Like f but not suppressed by su(f). RK1. Muller, 1935, DIS 3: 30.
other information: In left sublocus of forked. phenotype: fa^/fa resembles fa/fa; not notched.
fa: facet Homozygous lethal. RK2.
location: 1-3.0.
origin: Spontaneous.
discoverer: Bridges, 14b.
references: Morgan and Bridges, 1916, Carnegie
Inst. Wash. Publ. No. 237: 76.
phenotype: Eyes of all males moderately rough
owing to irregularity in size, shape, and arrange-
ment of facets. Eyes of females less rough, with
about 10 percent overlap of wild type. Eye
abnormality caused by overgrowth of secondary
pigment cells, which compresses cones and
causes overlying corneal facet to bulge
(Waddington and Pilkington, 1942, DIS 16: 70).
Wings have apical nicks in 0.25 percent of males
and 0—5 percent of females. N/fa has rough eyes
of fa as well as a Notch phenotype; that is, fa has
a pseudodominant effect with Notch. RK2.
cytology: Salivary chromosome locus placed at 3C7
by Slizynska (1938, Genetics 23: 291-99).
Salivary chromosomes normal (Welshons). fa": facet-notched
other information: A member of the pseudoallelic From Glass, 1933, J. Genet. 27: 233-41.
series at the Notch locus (Welshons and
Von Halle, 1962, Genetics 47: 743-59; Welshons, fa": facet-notched
Von Halle, and Scandlyn, 1963, Proc. Intern. origin: X ray induced in In(l)dl-49.
Congr. Genet., 11th, Vol. 1: 1-2), located discoverer: Glass, 1929.
between JV 55eJjr and fano. references: 1933, J. Genet., 27: 233—41 (fig.).
phenotype: Wings have apical nicks or notches in
fa3: see spl 90—100 percent of males, but only about 8 percent
*fado-vg: facet-dominigene for vestigial of homozygous females. Eyes not rough. fa/fan is
origin: Spontaneous. wild type. Viability and fertility excellent. RK2
discoverer: Goldschmidt. in male.
references: 1935, Z. Induktive Abstammungs- cytology: fa" is on an In(l)dl-49 chromosome but
Vererbungslehre 69: 38-131 (fig.)- should be separable from the inversion.
1935, Biol. Zentr. 55: 535-54 (fig.). fano; facet-notchoid
Gardner, 1942, Univ. Calif. (Berkeley) Publ. location: About 0.05 unit to right of fa.
Zool. 49: 85-102. origin: X ray induced.
phenotype: By itself it is wild type. fad°-v6/fa discoverer: Bauer, 1943.
shows rough-eye character of fa. fado~v6/fado"vi', references: 1943, Z. Induktive Abstammungs-
v&/+ produces some wing notching. RK3. Vererbungslehre 81: 374-90 (fig.).
other information: Presumed by Goldschmidt to Welshons, 1958, Proc. Natl. Acad. Sci. U.S. 44:
enhance dominance of vg and thus termed a 254-58.
' * d ominigene''. 1958, Cold Spring Harbor Symp. Quant. Biol. 23:
fa9: facet-glossy 171-76.
origin: Spontaneous. Welshons and Von Halle, 1962, Genetics 47:
discoverer: Pratt, 1962. 743-59.
phenotype: Eyes have facets more irregular than fa, phenotype: Wings of both sexes notched at ends of
but surface is smoothed, giving a glossy effect. L3 and LA veins; other longitudinal veins
Pigment distribution may be uneven, contributing enlarged and show deltas. Somewhat less extreme
to an impression of altered eye color. No wing ef- at elevated temperatures. Up to 5 percent of fano
fect. Eyes of fa*//a intermediate between the two males from aged cultures show hyper- and hypo-
. Complementary with apt, fano, nd, developed external genitalia (Kroeger, 1960,
(Vtm Halle and Welshons). RK.1. J. Morphol. 107: 227—32). Heterozygote shows
cytology: Salivary dtarotBOSoroe* normal (Welshons). extremely weak dominance. fetno/N almost com-
other infaraMrttoa: Loc@t©d to the left of fe**0; not pletely lethal. Rare survivors have exaggerated
y«t ••pttnted from fa (Wmlshoas,). Notch phenotype. taao/fa closely resembles wild
*hh fwet-htlml type; nicks in wings appear infrequently. RK1.
dJ«c«v«r*r: Matter, 19b. cytology: Salivary chromosomes normal (Welshoos).
MUTATIONS 95

other information: A member of pseudoallelic series normal eggs also laid. Hatchability reduced and
at the Notch locus, located between fa and variable. RK3.
N264-40, fg: see spdl&
facetious: see rgp *fi: frail
/as/: see rgp location: 1-53.
fat: see ft origin: Recovered among progeny of flies treated
faulty chaetae: see fc with Janus green.
*fb: fine bristle discoverer: Muller, 28e20.
location: 1-1.0. references: 1935, DIS 3: 30.
origin: Induced by D-p-NN-di-(2-chloroethyl)amino- phenotype: Wings nearly as small as tn, thin and
phenylalanine (CB. 3026). frail. Bristles fine. Fly weak. Viability 10—30
discoverer: Fahmy, 1954. percent wild type. RK3.
references: 1958, DIS 32: 70. *fil: fine lash
phenotype: Thin, slightly shortened bristles. location: 1-56.8.
Occasional scalloping of wing margins. Delayed origin: Induced by L-p-NN-(2-chloroethyl)amino-
emergence. Good viability and fertility both phenylalanine (CB. 3025).
sexes. RK3. discoverer: Fahmy, 1953.
*fc: faulty chaetae references: 1959, DIS 33: 86.
location: 1-0.9. phenotype: Thin, slightly shorter bristles. Eyes
origin: Induced by DLT3-NN-di-(2-chloroethyl)amino- reduced in size; posterior border very close to
phenylalanine (CB. 3007). orbital bristles. Both sexes viable and fertile.
discoverer: Fahmy, 1954. RK3.
references: 1958, DIS 32: 70. other information: Two alleles induced by CB. 1528.
phenotype: Short, thin bristles. About one-third of fin: finer
flies show either absence or duplication of one location: 1-29.6.
scute liar bristle. Viability and fertility good both origin: Induced by D-p-NN-di-(2-chloroethyl)amino-
sexes. RK2- phenylalanine (CB. 3026).
*fd: furled discoverer: Fahmy, 1954.
location: 1- (rearrangement). references: 1959, DIS 33: 86.
origin: Induced by P 3 2 . phenotype: Fly slightly smaller than normal, with
discoverer Bateman, 1949. shorter, thinner bristles. Delayed eclosion. Males
references: 1950, DIS 24: 54. viable but sterile. RK3.
1951, DIS 25: 77. fine bristle: see fb
phenotype: Like vestigial, but with immovable fine chaetae: see fnc
mouth parts and fully extended proboscis. Dies fine lash: see fil
early, perhaps owing to failure to ingest. Viability fine macros: see fm
at eclosion good. RK3A. finer: see fin
cytology: Associated with T(l;3)fd= T(1;3)7A;86E +
In(3R)89C;96A (Darby).
Female lethal: see Fl
female sterile: see fs( )
Female sterile(2) Dominant: see Fs(2)D

fern: see is(2)B


fes(2)K: see is(2)K
ff: fluff
location: 1-57.7.
origin: Induced by 2-chloroethyl rnethanesulfonate
(CB. 1506).
discoverer: Fahmy, 1955.
references: 1959, DIS 33: 86.
phenotype: Extremely fine short bristles. Wings fj: four jointed
slightly rounded at tips. Males and females viable Second and third legs.
and fertile; eclosion delayed. RK3. Edith M. Wallace, unpublished.
other information: One allele induced by CB. 1414.
*fft: fused filament //; four jointed
location: Not located, location: 2-81.
origin: Spontaneous. origin: Spontaneous.
discoverer: Robertson and Reeve. discoverer: Schultz, 31dl.
references: 1954, DIS 28: 78. phenotype: Tarsi four instead of five jointed. Legs
phenotype: Chorionic filaments of eggs laid by fit short and stocky. Enhanced by s s a and ss*B
female usually fused into a single structure. A few (Villee, 1945, Genetics 30: 26-27). Development
96 GENETIC VARIATIONS OF DROSOPHILA MELANOGAS

similar to that of dachs LWaddington, 1943, J. phenotype: Fertility of Fls/+ females may be
Genet. 45: 29-43 (fig.)]- Wings shorter, broader, reduced. Fls/Fl* females sterile. Fl/Fls fema
with crossveins conspicuously closer together; lethal. RK3.
veins diverge at greater angle. Effect visible in fla: flat eye
prepupal wing (Waddington, 1940, J. Genet. 41: location: 1-2.4.
75—139). Eyes smaller, ellipsoid, coarse origin: Induced by L-p-NN-di-(2-chloroethyl)amin
textured; head foreshortened. RK2. phenylalanine (CB. 3025).
discoverer: Fahmy, 1953.
origin: Spontaneous. references: 1958, DIS 32: 70.
discoverer: Ives, 40e. phenotype: Smaller fly, with smaller and less-
references: 1941, DIS 14: 39. curved eyes. Wings extremely variable, from
phenotype: Venation irregularities more extreme normal, through incised margins, to crumpled
than those described for fj, but variable and may vestigial stumps. Not easily classified. Viabil
overlap wild type. Production of fj*Oe flies low and fertility good in males but reduced in femal
in competition with wild-type flies in crowded RK3.
cultures, but normal in a homozygous stock. flap wing: see flw
Developmental period about 20 percent longer flateye: see fla
than wild type. RK2. flipper: see flp
*///: ftyless
location: 3- (not located).
origin: Spontaneous,
discoverer: Cercos, 41gl5.
references: Andres, 1943, DIS 17: 48.
phenotype: Wings apparently normal, but fly cann
keep them spread and cannot fly more than a fex
inches. RK3.
*flp: flipper
//: fluted location: 2-30.
Edith M. Wallace, unpublished. origin: Spontaneous.
discoverer: Mohr, 18h5.
fl: fluted references: Bridges and Mohr, 1919, Genetics 4:
location: 3-59.9. 304.
origin: Spontaneous, phenotype: Wings fail to expand; remain compact,
discoverer: Redfield, 211. very dark, extended, and curved slightly down-
phenotype: Wings creased lengthwise and dark. ward. Fly a wizened dwarf. Body surface dull
Overlaps wild type slightly at 25° but not at and dark. Both sexes sterile, RK3.
19°C. RK3. flp: see flw
*f!2 fluff: s e e ff
origin: Spontaneous. fluted: s e e fl
discoverer: Spencer, 36dl5. *flw: flap wing
phenotype: Like tl, RK3. location: 1-31.
Fl: Female lethal discoverer: Waletzky, 1937.
location: 1-19.1 (based on crosses with F/ s ). synonym: flp, a preoccupied symbol.
origin: Spontaneous. phenotype: Wings held out; consistently concave
discoverer. Muller and Zimmering, 1960. upward; slightly pointed. Darkened longitudinal
references: 1960, Genetics 45: 1001—2. stripe along thorax, underneath chitin; occasion,
phenotype: Viability of F1/+ females varies from bubbling, with maximum pigmentation in anterior
zero to normal, depending on maternal genotype. part of stripe. Anterior scutellars sometimes
Where viability is low, surviving females often missing or doubled. Eyes bulging; slightly
show diverse morphological abnormalities. Fl/Fl roughened. Head compressed anteroposteriorly.
females lethal; die as early embryos (Oster). Third antennal joint shortened. RK2.
tnftrm has no effect on survival of Fl/Fl or F1/+
(Zimmering and Muller, 1961, DIS 35: 103-4). Ft/Y flyless: see HI
males have normal viability and phenotype. RK2A. *fm: fine macros
cytology: Present in X containing In(l)dt-49 = location: 1-66.1.
ln(l)4D7-El;llF2-4. origin: Induced by 2-chloroethyl methansesulfonate
(CB. 1506).
Fl*: Female lethal-sterile discoverer: Fshmy, 1956.
origin: Spontaneous. references: 1959, DIS 33: 86.
discovmmr, Zimmering and Muller, 1961. phenotype: Small fly with narrow abdomen and
synonym; F!%. extremely short, thin bristles. Males fertile;
references: 1961, DIS 35: 103-4. viability about 50 percent wild type. RK3.
MUTATIONS 97

*fnc: fine chaetae margin irregularity. Viability variable, from 16—90


location: 1-34.9. percent wild type. Females rarely fertile. Char-
origin: Induced by S-2-chloroethylcysteine acter less extreme at low temperature. RK3.
(CB. 1592). *fr°: fringed-zero
discoverer: Fahmy, 1957. origin: Spontaneous.
references: 1959, DIS 33: 86. discoverer: Bridges, 15a20.
phenotype: Extremely fine short bristles. Body references: Bridges and Morgan, 1919, Carnegie
parts disproportionately reduced; reduction least Inst. Wash. Publ. No. 278: 257 (fig.).
marked on head and most marked on abdomen. phenotype: Wing margins have bare spots;
Wings broad and slightly rounded at tips, remaining hairs frayed and irregular. Wings
occasionally with incisions of margin. Eyes smaller, discolored, somewhat divergent. Like fr
slightly brighter red than normal. Males viable but but less extreme. RK3.
sterile. RK3. other information: Lost before fr, fr2, and frdi were
fo: folded found.
location: 1-63. fr2
discoverer: Grossman, 1932. origin: Spontaneous.
references: 1934, DIS 1: 30. discoverer: Novitski, 37a22.
phenotype: Wings remain unexpanded in a varying synonym: tan: trimmed.
percentage of flies. Balancers shriveled and post- references: 1937, DIS 8: 10, 13.
scutellars bent forward. Overlaps wild type. RK3. Lewis, 1938, DIS 10: 55-56.
Fo; Forkoid phenotype: Wings scalloped and fringed on all
location: 2-107 (between or and sp). margins, slim and coarse textured, often divergent
origin: X ray induced. and uplifted. Eyes rough. Some extra bristles
discoverer. Mohler, 5 8 c l 8 . present. Abdomen slightly offset, as in fr.
references: 1960, DIS 34: 52. Female sterile. Classification at 25°C excellent;
phenotype: Heterozygote shows reduction in size of characters less extreme at 19°C. Viability
bristles and weak forking of head and posterior generally good, but erratic. RK2.
thoracic bristles. Using Dp(2;3)P, it may be shown *frdl; fringed-dibro
that the expression of +/+/Fo <+/Fo <+/Fo/Fo; origin: Spontaneous.
+/F0/F0 shows extreme forking of all bristles and discoverer: Bridges, 17kl9.
is sterile. Homozygous lethal. Fo interacts with f synonym: dihro.
alleles to produce extreme f bristles. RK1. references: Lynch, 1920, Genetics 4: 527—28.
cytology: Located between 58E3 and 60B10, on phenotype: Spread wings with scalloped margins.
basis of its inclusion in Dp(2;3)P = Dp(2;3)58E3- Eyes rough. Very inviable; both sexes sterile.
F2;60D14-E2;96B5-Cl but not in Df(2R)Px = RK3.
Df(2R)60B8-10;60Dl-2 (Mohler) or in the deficiency frail: s e e fi
for the tip of 2R derived from T(l;2)Bld =
T(1;2)1C3-4;6OB12-13 (Armentrout). Frd: Freckled
location: 2-103.1 (Nicoletti).
focal melanosfs: see me origin: X ray induced.
*fol: folded wings discoverer: M. G. Davis, 1961.
location: 2-39. references: Erlich, 1963, DIS 37: 47.
origin: Spontaneous. Barigozzi, 1963, Proc. Intern. Congr. Genet., 11th.
discoverer: Goldschmidt, 1937. Vol. 1: 207.
phenotype: Expanded wing folded. Overlaps wild 1965, DIS 40: 64.
type. RK3. phenotype: Pupa and young fly characterized by ac-
folded: see fo cumulation of dark pigment; in older fly, pigment
folded wings: see /b/ becomes concentrated in black specks scattered
forked: see f throughout body, head, and legs. Homozygous
Forkoid: see Fo lethal. RK2.
four jointed: see // other information: Barigozzi claims it has an extra-
chromosomal or episomal component in its trans-
fr: fringed mission.
location: 2-80. fringed: see fr
origin: Spontaneous. frizzled: see fz
discoverer: Bridges, 22c30. fs 2.: see fs(2)E
references: 1938, DIS 9: 48. fs(l)N: female sterile (1) of Nasrat
phenotype: Wings often spread; wing margins location: 1-0.0 (closely linked to me).
snipped and bristles irregular and fringelike. origin: Induced by an unspecified chemical
Eyes small and rough. Midline of abdomen at mutagen.
slight angle to ioagitudinal axis of fly. Much discoverer: Nasrat, 1952.
variability in expression; safest criterion is wing synonym: /®f2)" ae .
GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
98

phenotype: Females sterile but no other distin- origin: Ultraviolet induced.


guishing characteristics. Males fertile. Develop- discoverer: Edmondson, 50j.
mental study by Counce and Ede [l957, J. synonym: fs2.1.
Embryol. Exptl. Morphol. 5: 404-21 (fig.)]. Eggs of references: Meyer and Edmondson, 1957, DIS 25:
ta(l)N/i8(l)N females will not support develop- 72.
ment of normal embryos. About half the eggs phenotype: Sterile females do not lay eggs. Gonads
contain little or no yolk; development may or may rudimentary. RK3.
not begin in such eggs but never progresses *fs(2)E2
beyond a highly abnormal cleavage. In eggs location: 2-22.0.
containing more yolk, major effect is on synchrony origin: Ultraviolet induced.
of cleavage and blastoderm mitoses. Twenty per- discoverer: Edmondson, 1951.
cent of these embryos cease development before synonym: fs2.2.
blastoderm formation. The remainder have references: 1952, DIS 26: 61.
abnormal blastoderms and aberrant gastrulation. phenotype: Females produce eggs that appear
Final pattern of damage determined by degree of normal but do not hatch. Fertile in heterozygotes
abnormality of earlier stages, but some embryos with ms(2)E3 (2-28.0). RK3.
show larval differentiation. A few of the least *fs(2)E3
abnormal embryos may emerge but never move location: 2-47.5.
about or feed. Formation of polar granules origin: Ultraviolet induced.
abnormal. RK3. discoverer: Edmondson, 1951.
cytology: No detectable chromosomal rearrange- synonym: fs2.3.
ments (Slizynska). references: 1952, DIS 26: 62.
fs(2)5*6: see is(2)E9 phenotype: Female sterile; narrow curved wings.
ts(2)adp: see adpte No eggs laid. Fertile in heterozygotes with
h(2)B: female sterile (2) of Bridges ms(2)E4 (2-47.9), fs(2)E4 (2-48.5), fs(2)E5 (2-50.4),
location: 2-5. fs(2)E6 (2-54.4), ms(2)E5 (2-54.8), ms(2)E6
origin: Spontaneous. (2-54.8), ms(2)E7 (2-54.8), ts(2)E7 (2-55.2),
discoverer: Bridges, 29c2S. ms(2)E8 (2-55.6), and ms(2)E9 (2-57.0). RK3.
synonym: fes, *fs(2)E4
references: King, Sang, and Leth, 1961, Exptl. Cell. location: 2-48.5.
Res. 23: 108-17 (fig.). origin: Ultraviolet induced.
King, Koch, and Cassens, 1961, Growth 25: 45-65 discoverer: Edmondson, 1951.
(fig.). synonym: is2.4.
Koch and King, 1964, Growth 28: 325-69 (fig.). references: 1952, DIS 26: 62.
phenotype: External morphology normal. Male fer- phenotype: Very few eggs laid; female sterile. Fer-
tile; female sterile. Ovarioles of homozygous fe- tile in heterozygotes with fs(2)E3 (2-47.5), ms(2)E4
male subdivided into a series of sausage-shaped (2-47.9), fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), ms(2)E5
cell aggregates, each surrounded with an ill- (2-54.8), ms(2)E6 (2-54.8), ms(2)E7 (2-54.8),
defined follicular epithelium and filled with hun- is(2)E7 (2-55.2), ms(2)E8 (2-55.6), and ms(2)E9
dreds to thousands of mitotically active oogonia- (2-57.0). RK3.
like cells [King, Burnett, and Staley, 1957, Growth *fs(2)E5
21: 239—61 (fig.)]. These cells occasionally dif- location: 2-50.4.
ferentiate into cells resembling nurse cells, which origin: Ultraviolet induced.
may have polytene chromosomes, and rarely into discoverer: Edmondson, 1951.
oocytes. la(2)B ovaries transplanted into wild-type synonym: fs2.5.
hosts in late larval stages and reciprocal trans- references: 1952, DIS 26: 62.
plant® develop autonomously (Clancy and Beadle, phenotype: Female semisterile. A few larvae hatch
1937, Biol. Bull, 72: 47-56; Bodenstein and King, normally, but most embryos degenerate. Fertile in
1963, DIS 37: 65; 1965, Z. Naturforsch. 20b: 292- heterozygotes with fs(2)E3 (2-47.5), ms(2)E4
97). RK3. (2-47.9), fs(2)E4 (2-48.5), fs(2)E6 (2-54.4), ms(2)E5
Fs(2}D: Female sterile(2) Dominant (2-54.8), ms(2)E6 (2-54.8), ms(2)E7 (2-54.8),
location: 2- (not located). ts(2)E7 (2-55.2), ms(2)E8 (2-55.6), and ms(2)E9
origin: Induced by ethyl methanesulfonate. (2-57.0). RK3.
discoverer: E. H. Grell, 65e. *fs(2)E6
pfieraotype: Heterozygous female entirely sterile, location: 2-54.4.
with underdeveloped ovaries. Heterozygous male origin: Ultraviolet induced.
ba» normal fertility. Bristles short; thorax broad discoverer: Edmondson, 1950.
sad flattened with air bubbles under cuticle. RK3. synonym: f&2.6.
eth*r information: Sterility makes genetic mapping references: 1952, DIS 26: 62.
phenotype: Female produces normal-appearing
*fs(2)Eh female sterile (2) of Edmondson eggs, which do not hatch. Fertile in heterozygotes
location: 2-57.6. with tm(2)E3 (2-47.5), ms(2)E4 (2-47.9), ts(2)E4
MUTATIONS 99

(2-48.5), fs(2)E5 (2-50.4), ms(2)E5 (2-54.8), origin: X ray induced.


ms(2)E6 (2-54:8), ms(2)E7 (2-54.8), fs(2)E7 discoverer: Gill, 59a.
(2-55.2), ms(2)E8 (2-55.6), ms(2)E9 (2-57.0), and synonym: fs(3)2*9*.
is(2)E8 (2-62.6). RK3. references: 1960, Anat. Record 138: 351.
*fs(2)E7 1961, Ph.D. Thesis, Yale Univ.
location: 2-55.2. 1962, DIS 36: 37.
origin: Spontaneous. 1963, J. Exptl. Zool. 152: 251-78 (fig.).
discoverer: Edmondson, 1950.
phenotype: Females almost sterile; produce rare
synonym: fs2.7.
surviving progeny. Oogenesis incomplete; usually
references: 1952, DIS 26: 62.
stops in early phases of vitellogenesis. Most (89
phenotype: Females produce collapsing eggs.
percent) follicles contain 32 cells instead of
Apparently, there is a weakness in the vitelline normal 16 as a result of an extra oogonial divi-
membrane, since these eggs cannot be success- sion. The 32 cells of an incipient cyst enclosed
fully dechorionated; when chorion is removed, egg in two chambers in 6 percent of the cases. Posi-
contents flow out. Fertile in heterozygotes with tion of oocyte in follicle abnormal in 28 percent of
fs(2)E3 (2-47.5), ms(2)E4 (2-47.9), fs(2)E4 (2-48.5), cases. Males partially sterile. Viability low.
fs(2)E5 (2-50.4), fs(2)E6 (2-54.4), ms(2)E5 (2-54.8), RK3.
ms(2)E6 (2-54.8), ms(2)E7 (2-54.8), ms(2)ES
(2-55.6), ms(2)E9 (2-57.0), and fs(2)E8 (2-62.6). fs(3)G3
RK3. location: 3-25.
*fs(2)E8 origin: X ray induced.
location: 2-62.6. discoverer: Gill, 59a.
origin: Ultraviolet induced. synonym: fs(3)3^9a.
discoverer: Edmondson, 49k. references: 1960, Anat. Record 138: 351.
synonym: fs2.8. 1961, Ph.D. Thesis, Yale Univ.
references: 1952, DIS 26: 62. 1962, DIS 36: 37.
phenotype: Females do not lay eggs. Fertile in 1963, J. Exptl. Zool. 152: 251-78 (fig.),
heterozygotes with fs(2)E6 (2-54.4), ms(2)E5 phenotype: Oogenesis incomplete; most follicles
(2-54.8), ms(2)E6 (2-54.8), ms(2)E7 (2-54.8), stop development during yolk deposition (after
fs(2)E7 (2-55.2), ms(2)E8 (2-55.6), ms(2)E9 stage 9). Males fertile. RK3.
(2-57.0), ms(2)E10 (2-66.5), ms(2)Ell (2-68.0), and
ms(2)E12 (2-68.2). RK3. fs(3)G5
*fs(2)E9 location: 3-49.
location: 2-35.6. origin: X ray induced.
origin: Ultraviolet induced. discoverer: Gill, 59a.
discoverer: Edmondson, 1958. synonym: fs(3)5S9a.
synonym: fs(2)ss&. references: 1960, Ar"'. Record 138: 351.
references: 1960, DIS 34: 49. 1961, PhJD. Thesis, Yale Univ.
phenotype: Numerous infertile eggs produced. 1962, DIS 36: 37.
Viability normal. Males fertile. RK3. 1963, J. Exptl. Zool. 152: 251-78.
*fs(2)K: female sterile (2) of Kikkawa phenotype: Oogenesis incomplete; ovarioles
location: 2-100. contain excessive numbers of follicles, which
origin: Spontaneous, usually stop developing at or before stage 9.
discoverer Kikkawa, 1960. Males fertile. RK3.
synonym: ies(2)K. ft: fat
references: 1960, DIS 34: 51. location: 2-12.0.
phenotype: Female sterile. Male fully fertile. RK3. origin: Spontaneous.
fs(3)Gh female sterile (3) of Gill discoverer: Mohr, 2Obi5.
location: 3-47. references: 1923, Studia Mendeliana (Brunae)
origin: X ray induced. pp. 266-87.
discoverer: Gill, 59a. 1929, Z. Induktive Abstaimmngs- Vererbungslehre
synonym: fs(3)l*9». 50: 113-200 (fig.).
references: 1960, Anat. Record 138: 351. phenotype: Abdomen short and fat. Thorax broad.
1961, Ph.D. Thesis, Yale Univ. Wings short and broad, with crossveins much closer
1962, DIS 36: 37. together than normal. Scutellutn shortened;
1963, J. Exptl. Zool. 152: 251-78 (fig.). scutellar bristles far apart. Viability good. Second-
phenotype: Eggs die in early cleavage stage. and third-instar larvae, particularly when there is
Meiosis precocious in 2 percent of oocytes; first little yeast in the food, show vacuoles in cyto-
meiotic spindle parallel to egg surface. Males plasm of salivary gland cells. Two waves of
fertile. RK3. vacuole formation. Cells with vacuoles exhibit a
slight puff in 24D-E, which is negatively corre-
fs(3)G2
lated with puffing in 21 and 22. Tip of X disfigured.
location: 3-11.
TOO GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

possibly as a result of several small puffs inter- In(l)ClLy4R = Jn(l)4A5-Bl;17A6-Bl^lA8-


mingled with hard nonpuffed bands. In about 1 per- B1;18A3-4R] are more extreme than those in fu
cent of larvae, salivary glands distally expanded homozygote (King, 1959, DIS 33: 142-43). fu/fu
and crooked [Slizynski, 1964, Cytologia (Tokyo), ovaries transplanted into fu + hosts develop auton-
29: 330-36 (fig.)]. RK1. omously in regard to fertility (Clancy and Beadle,
cytology: Placed between 24D2 and 24F1 on the 1937, Biol. Bull. 72: 47-56; Sobels, 1950, Ex-
basis of its inclusion in Di(2L)M-zc = Df(2L)24D2- perientia 6: 139—40) and tumor formation (Boden-
5;25A2-3 but not in Df(2L)M-zB = Df(2L)24E2- stein and King, 1963, DIS 37: 65). The few normal-
Fl;25Al-2 (Morgan, Bridges, Schultz, and Curry, appearing eggs that are laid by fu/fu females
1939, Carnegie Inst. Wash. Year Book 38: 273-77). produce adults only if they have been fertilized by
ftj+-bearing sperm (Lynch, 1919, Genetics 4: 501 —
33). Eggs fertilized by fu- or F-bearing sperm de-
velop into embryos that become abnormal 5-5V2 hr
after fertilization. A general asymmetry in germ
layers is responsible for many ensuing abnormali-
ties. Such embryos never hatch but survive long
after normal embryos have become larvae LCounce,
1956, Z. Induktive Abstammungs- Vererbungslehre
87: 462-81 (fig.)], fu eggs from fu/+ mothers de-
velop normally. Heterozygous daughters from homo-
zygous mother have high incidence of abnormal ab-
dominal segmentation and, as embryos, have
abnormal musculature. This is a maternal effect
not found in the reciprocal cross, and it is temper-
ature sensitive (Armstrong and Sobels). RK1.
cytology: Salivary chromosome location of fu
appears to be in 17D or E (Green, 1953, Genetics
38: 91—105; 1953, Z. Induktive Abstammungs-
Vererbungslehre 85: 435—49).

origin: Ultraviolet induced.


discoverer: Edmonds on, 51e.
fu: fused references: Meyer and Edmondson, 1951, DIS 25: 72.
Edith M. Wallace, unpublished. phenotype: Like fu, but possibly slightly more
extreme. RK1.
fu; fused fvS7a
location: 1-59.5. origin: Induced by azo-mustard.
origin: Spontaneous. discoverer: Purdom, 57a.
discoverer: Bridges, 12k4. references: King, Burnett, and Staley, 1957, Growth
references: Morgan and Bridges, 1916, Carnegie 21: 239-61.
Inst. Wash. Publ. No. 237: 55-58 (fig.). phenotype: Veins L3 end L4 fused from origin to
Lynch, 1919, Genetics 4: 501-33. beyond anterior crossvein. Ocelli and ocellar and
phenotype: Veins L3 and L4 fused from base to anterior scutellar bristles sparse or absent. Length
beyond anterior crossvein, with elimination of of scute Hum generally reduced. Female sterile as
anterior crossvein and first basal cell; L3 and L4 with hi. Ovaries tumorous but to a lesser extent
fused at tip; this fusion may reach back to basal than with fu. Wings not outstretched but held in
cell. Wings usually extended. Ocelli reduced or normal position. RK1.
absent; bristles of ocellar region small or absent.
Eyes small and slightly rough. Anterior scute liar *fu57f
bristle® reduced in number, and scute Hum short- origin: Induced by azo-mustard.
ened. Female late to eclose and has decreased discoverer: Purdom, 57f.
longevity. Ovaries histologically normal at eclo- references: King, Burnett, and Staley, 1957, Growth
sioo but with half the normal number of ovarioles 21: 239-61.
(BMtty, 1949, Proc. Roy. 8oc. Edinburgh, B 63: phenotype: Wing veins L3 and L4 often completely
249—70); fecundity 7 percent normal. Dewioping fused. Wings outstretched. Ovaries tumorous.
egg chambers may fuse or become timorous with RK1.
at® [King, Boraett, and Staley, 1957, Growth 21:
239—61 (fig.)]. Proportion of tumorou* egg chambers origin: Spontaneous.
increases by 6 percent pan day. Female raised at discoverer: R. F. Grell, 1959.
1S°C stews only 10 percent th« tumor development
references: King and Smith, 1963, DIS 37: 49.
of that rmlaed at 25°. Ovarian effects i& female
phenotype: Wings like fu. Rate at which ovarian
cairying fa and • deficiency for §u [i.e.,
tumors develop is the lowest of all alleles tested;
MUTATIONS 101

number of tumorous egg chambers increases by 1


percent per day. RK1. origin: X ray induced.
*fu6lfl discoverer: MuHer, 31a.
origin: Gamma ray induced. phenotype: Extreme fw. Female sterile. RK2A.
discoverer: Fahmy, 62f. cytology: On a chromosome containing In(l)dl-49.
references: Smith and King, 1963, DIS 38: 39. fw34e
phenotype: Like fuS7a. RK1. discoverer: Duncan, 34e20.
* fa 62 i2 phenotype: Originally showed eye surface medium
origin: Gamma ray induced. folded; bristles much gnarled. Schultz and Curry
discoverer: Fahmy, 62f. report that stock in 1940 showed gnarled bristles
references: Smith and King, 1963, DIS 38: 39. and eye small but no vertical fold. RK2.
phenotype: Wings and ocelli show typical abnormali- fw49c
ties. Ovaries develop tumors. RK1. origin: Induced by P^2.
fu62f3 discoverer: R. C. King, 49c28.
origin: Gamma ray induced. references: 1949, DIS 23: 62.
discoverer: Fahmy, 62f. phenotype: Eyes furrowed; distal portions of aristal
references: Smith and King, 1963, DIS 38: 39. branches hooked; wings divergent and often
phenotype: Like fa. RK1. stringy; scutellar groove reduced. Bristles split,
*fuff: fused-formalin food bent, and often erect; acrostichal hair pattern dis-
origin: Induced by formaldehyde. tributed with whorls and naked areas. Late
discoverer: Auerbach, 1951. hatching, poorly viable, and mostly sterile.
references: Counce, 1956, Z. Induktive f w 49c/f w phenotypically intermediate but more like
Abstammungs- Vererbungslehre 87: 462—81. fw/fw than fw49c/fw49c^ R K 3
King, Burnett, and Staley, 1957, Growth 21:
239-61.
phenotype: Like fu but wings not outstretched. origin: X ray induced.
Ovaries contain tumors. Development of lethal discoverer: Garcia-Be Hid o, 59i21.
embryos roughly similar to that of fu (Counce, references: 1963, Genet. Iberica 15: 1—102.
1956). RK1. phenotype: Eyes rough and creased; facets irregular,
*fu9: fused-glider 15 percent fewer than normal. Eyes browner than
origin: Found among heat-treated flies, normal; pterine concentration reduced in the eyes
discoverer Grossman, 1932. and, except for isoxanthopterine, increased in
references: 1934, DIS 1: 30. testis sheath. Riboflavin accumulates in Mal-
phenotype: Like fu except wings more divergent and pighian tubules. Large bristles of head and thorax
females more fertile, fu^/fu females 40 percent short, thick, angled, and blunt; occasionally re-
fertile. fu&/fu6 females 20 percent fertile. In duced to stumps. Arista thick with contorted and
general, pattern of development of lethal embryos supernumerary branches. Scute Hum small with
is like that of fu (Counce, 1956, Z. Induktive groove between it and thorax reduced. Hatchability
Abstammungs- Vererbungslehre 87: 462—81). RK1. and larval development normal; larval anal plates
furled: see fd swollen and surrounded by melanotic halo. Mela-
furrowed: see fw notic anal region persists in pupa; pupa also has
fused: see fu melanotic spots elsewhere that may result in non-
fused filament: s e e fft pigmented areas on the imaginal integument. Ex-
fuzzy: see fy trusion of anterior and posterior spiracles in pre-
fw: furrowed pupa incomplete. Many fn>5 9 flies die either after
location: 1-38.3. 24-30 hr of pupal development or at the time of
origin: Spontaneous, eclosion. Fecundity of female reduced owing to
discoverer: Duncan, 14k. reduced number of ovarioles. RK2.
references: 1915, Am. Naturalist 49: 575—82.
Morgan and Bridges, 1916, Carnegie Inst. Wash. origin: X ray induced.
Publ. No. 237: 80. discoverer: Garcia-Bellido, 60k8.
Nachtsheim, Z. Induktive Abstammungs- references'. 1963, Genet. Iberica 15: 1-102.
Vererbungslehre 20: 118-56. phenotype: Like fw59 but with lower penetrance and
phenotype: Eyes with vertical fold and furrows. expressivity. RK2.
Head and scute Hum shortened. Bristles gnarled
and shortened, especially the postscutellars. Best *fww; furrowed-weak
classification character is short, blunt noto- origin: Spontaneous.
pleurals. RK2. discoverer Ives, 43b24.
cytology: Placed between 10E2 and 11A7 on the references: 1946, DIS 19: 46.
basis of its being to the right of dy, which is to phenotype: Affects only bristles, particularly the
the right of 10E2 and to the left of the X break- scutellars and postaiars. Eyes normal. Normal
point of T(l;4)A8 » T(l;4)llA6-7. fertility and viability. RK2.
102 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

fwwr: {urrowed-wrinkled discoverer: Bridges, 15bl9.


origin: X ray induced in In(l)scslLscSR+dl.49. synonym: salmon.
discoverer R. M. Valencia, 1959. references: 1916, Genetics 1: 151.
synonym: wr, Chovnick, 1958, Proc. Natl. Acad. Sci. U.S. 44:
references: 1959, DIS 33: 100. 333-37.
1965, DIS 40: 36. 1961, Genetics 46: 493-507.
phenotype: Eye surface in folds. Some bristles phenotype: Eye color deep purplish ruby, like pr;
shortened, thickened, or curved; many doubled and pinkish in young and brownish in old flies. Eyes
may be fused. Viability low. RK2. have 38 percent normal red and 56 percent normal
*fy: fuzzy brown pigment (Nolte, 1959, Heredity 13: 233—41).
location: 2-33. RK1.
origin: Spontaneous. cytology: Placed in salivary chromosome region
discoverer: Ives, 39a. 12B9-12C7 by deficiency analysis (J. I. Valencia).
references: 1940, DIS 13: 49. other information: A pseudoallelic locus composed
phenotype: Hairs on abdomen and thorax irregular of four demonstrated sites; g2 occupies the left-
and directed toward midline. Hairs on wing margins most, gSOe £he second, g and g3 the third, and gsi
erect. Resembles fz. Fertility and viability below the right-most site (Chovnick, 1961).
normal. RK2.

50e sf
_J I

Map of the g locus


From Chovnick, 1961, Genetics 46: 493-507.

origin: Spontaneous.
discoverer: Bridges, 18c28.
references: Morgan, Bridges, and Sturtevant, 1925,
Bibliog. Genet. 2: 226 (fig.).
Sturtevant and Beadle, 1939, An Introduction to
Genetics, Saunders, p. 64 (fig.).
phenotype: Eye color translucent yellowish ruby,
somewhat lighter than g. Eyes contain 16 percent
normal red pigment and 32 percent normal brown
pigment (Nolte, 1959, Heredity 13: 233-41). Mal-
pighian tubes very pale yellow at base (Beadle,
fz: frizzled 1937, Genetics 22: 587-611). Eye color auton-
From Bridges and Brehme, 1944, Carnegie Inst. Wash. omous in transplant into wild-type host (Beadle
Pobl. No. 552: 85. and Ephrussi, 1936, Genetics 21: 230). With st,
gives dark yellow to orange color; with bw, gives
fz: frizzled reddish yellow to brownish rose (Mainx, 1938,
location: 3-41.7. Z. Induktive Abstammungs- Vererbungslehre 75:
origin: Spontaneous. 256—76). Dominant in females homozygous for
discoverer: Bridges, 38bl8. car, according to Schultz; i.e., pigmentation of
phenotype: Hairs on thorax directed irregularly g2 car/g2 car — g2 car/+ car < car/car. RK1.
toward midline. Thoracic bristles also in turned other information: Occupies left-most pseudoallelic
and often wavy. Postverticals may turn outward. site of the g locus (Chovnick, 1961, Genetics 46:
Hairs on wing edge and feet nearly erect. Wing 493-507).
may be reduced. Sex combs may be irregular. Eyes 1
always rough. Resembles in. RK2. origin: Spontaneous.
*fz46f discoverer. Bridges, 22d22.
origin: Spontaneous. references: Morgan, Bridges, and Sturtevant, 1925,
discoverer: Ives, 46120. Bibliog. Genet. 2: 226.
references: 1946, DIS 20: 65. phenotype: Like g, but gives lighter orange in
phenotype: Like fit. RK2. combination with v. Has 21 percent normal red
pigment and 47 percent normal brown pigment
9; qornmt (Nolte, 1959, Heredity 13: 233-41). Malpighian
location: 1-44.4. tubes extremely pale yellow (Brehme and Demerge,
origin: Spontaneous. 1942, Growth 6: 351-56). RK1.
MUTATIONS 103

other information: Occupies third pseudoallelic site g53d

from left of g locus (Chovnick, 1961, Genetics 46: origin: Spontaneous.


493-507). discoverer: Hexter, 53d.
94 references: 1958, Proc. Natl. Acad. Sci. U.S. 44:
origin: X ray induced in In(l)dl-49. 768-71.
discoverer: Glass, 1929. 1956, DIS 30: 72.
references: 1934, DIS 2: 7. 1963, Proc. Natl. Acad. Sci. U.S. 50: 372-79.
phenotype: Eye color, like g2, has 22 percent phenotype: Eye color orange, like wB. RK1.
normal red pigment and 23 percent normal brown other information: £53d, when heterozygous with g,
pigment (Nolte, 1959, Heredity 13: 233-41). g2, g3, or g4, yields wild-type progeny, some
Malpighian tubes extremely pale yellow (Brehme associated with crossing over in a manner con-
and Demerec, 1942, Growth 6: 351-56). RK1. sistent with ordering, and some not associated
cytology: Probably inseparable from In(l)dl-49. with recombination 6f outside markers. A double
mutant is not produced by the event that yields
g17Ba6 wild types.
origin: X ray induced in In(l)scslL'sc8ridl-49. *aSSk
9
discoverer: Muller, Valencia, and Valencia, 1946-53. origin: Spontaneous.
references: Valencia, 1966, DIS 41: 58. discoverer: Williams, 55k.
phenotype: Male viable but sterile. RK1A. references: Muller, 1956, DIS 30: 80.
cytology: Associated with In(l)gx 7Bfl<* = In(l)12B14- phenotype: Eyes translucent yellowish ruby. RK1.
15;19F (J. I. Valencia). *a271-2
*g26-10 origin: X ray induced,
origin: X ray induced. discoverer: Demerec, 1933.
discoverer: Sobels, 57j. phenotype: Male lethal, cell lethal. RK2.
references: 1958, DIS 32: 84. other information: ty but not s, wy, or pi also
phenotype: Eye color darker than g2. RK1. affected.
g26-75 *a271-6
origin: Induced by mustard gas. origin: X ray induced.
discoverer: Sobels, 57j. discoverer: Demerec, 34a.
references: 1958, DIS 32: 84. phenotype: Male lethal and cell lethal. RK2.
phenotype: Eye color like g2. RK1. other information: ty but not s or wy also affected.
*g26-41 *g27J-9
origin: Induced by mustard gas. origin: X ray induced,
discoverer: Sobels, 57j. discoverer. Hoover, 35h.
references: 1958, DIS 32: 84. phenotype: Male lethal. RK2.
phenotype: Weak garnet allele. Eye color strikingly other information: ty+.
different from wild type in newly emerged flies. *g271-T0
RK2. origin: X ray induced.
*g28-10 discoverer: Hoover, 35h.
origin: Induced by mustard gas. phenotype: Lethal. RK2.
discoverer Sobels and Jansen, 1957. other information: ty+, s + .
references: Sobels, 1958, DIS 32: 84. g(+): s e e g™
phenotype: Eye color like g2 but darkens more with
age. RK1. origin: Appeared among progeny of cold-treated
g28-40 flies.
origin: Induced by mustard gas. discoverer Gottschewski, 34gl7.
discoverer Sobels, 1957. references: 1935, DIS 4: 8, 15.
references: 1958, DIS 32: 84. phenotype: Eye color of males like g2, of females
phenotype: Eye color darker than g2. RK1. probably lighter than g*. RK1.
g49h 9**: garnet of Schalet
origin: Induced by P 3 2 . origin: Spontaneous,
discoverer: R. C. King, 49h. discoverer: Schalet.
references: 1950, DIS 24: 58. references: Chovnick, 1961, Genetics 46: 493—507.
phenotype: Eye color light purple. Viability of male other information: Occupies right-most pseudoallelic
normal, female 60 percent normal. RK2. site of the g locus.
gSOm gw: garnet-wild
references: Chovnick, 1958, Proc. Natl. Acad. Sci. origin: X ray induced,
U.S. 44: 333-37. discoverer: Muller.
1961, Genetics 46: 493-507. synonym: |f( + ).
phenotype: Eye color orange. RK1. references: 1946, DIS 20: 67.
other information: Occupies second pseudoalleiic Chovnick, 1958, DK 32: 88.
site from left of g locus. 1961, Genetics 46: 493-507.
GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER
104

phenotype: Homozygote and hemizygote indistin- phenotype: Does not show G phenotype. Allelic to
guishable from wild type; however, gw/g2 (and ft, but does not exaggerate ft. Lethal in combina-
probably other g alleles) is brownish. RK2A. tion with G. RK2.
cytology: Associated with In(l)gw (breakpoints un- g, Inh: see g*
known), which lies between dy and f. gap: see gp
garnet: see g
gx: garnet from X irradiation *Gd: Gulloid
origin: X ray induced. location: 3-78.
discoverer: Muller. origin: Spontaneous in Dp(2;3)P.
synonym: g, Inh. discoverer: Bridges, 22g26.
references: 1946, SIS 20: 67. phenotype: Gd/+ wings shorter, blunter, slightly
phenotype: Like g2. Male fertile; homozygous fe- more spread, and have crossveins closer together
male has low fertility. RK2A. than wild type. Homozygous lethal. RK3A.
cytology: Associated with /n(i;g* = In(l)12;19-20. cytology: Inseparable from Dp(2;3)P =
Dp(2;3)58E3-F2;60D14-E2; 96 B5- Cl.
*ge: genitalless
location: 1-0.1.
origin: Induced by methyl methanesulfonate
(CB. 1540).
discoverer: Fahmy, 1955.
references: 1958, DIS 32: 70.
phenotype: External male genitalia absent or
grossly deformed. Bristles fine; wings often small
and deformed. Tergites abnormal; abdomen
frequently contains melanotic tumors. Males viable
but sterile. RK3.
gespleten: see gs
*99: goggle
location: 1-23.1 (no crossovers with oc among 4300
flies).
G: Gull origin: Spontaneous.
From Mohr, 1929, Z. Induktive Abstammungs- discoverer: Nichols-Skoog, 34el4.
Vererbimgslehre 50: 113-200. phenotype: Eyes protruding and bulging; placed far
back on a narrow head. Facets very large in rough
G: Gull areas. Wings smaller, with fringed marginal hairs;
location: 2-12.0. dusky; pebbly appearance caused by large cells.
origin: Spontaneous. Bristles coarse and irregular; hairs sparse and
discoverer: Mohr, 19k23. irregular, especially on abdomen. Body small in
references: 1923, Studia Mendeliana (Brunae), late counts. Viability 20 percent wild type.
pp. 266-87 (fig.). Females usually sterile; males usually fertile.
1927, Proc. Intern. Congr. Genet., 5th. Vol. 2: RK3.
1136. gg2
1929, Z. Induktive Abstammungs- Vererbungslehre discoverer: Waletsky, 371.
501 113-200 (fig.). phenotype: Like gg, but many bristles on posterior
pti*nofyp«: Wings Urge, held out from sides at 45— lateral margin of head missing; others on head and
90°«gl«!, cttnred downward, and somewhat pointed. thorax reduced or missing. Females sterile; at
Vein LI thickened; crosaveins closer together, 19 C, ovaries small and contain no oocytes; at
s©«teti»es broken. Thoracic and vertical bristles 25°C ovaries contain a few eggs, but none are laid
tfoplicated in majority of flies. G/ft has exagger- (Beatty, 1949, Proc. Roy. Soc. Edinburgh, B 63:
ated It pfamotypt. Partially inhibited by d«/+ and 249-70). RK3.
ja»ei» inhibited by dc/rfc. Homozygous lethal. RK2. 99 3
cy*@J®fy: Placed between 24D2 und 24F1 on the origin: Spontaneous.
b*fti* of its inclasioo in Dt(2L,yt~zc = Df(2L)24D2- discoverer R. F. Grell, S3d.
&2SA2-3 bat »ot fa Df2L>f-x« *D§(2L)24E2~ references: 1953, DIS 27: 59.
Pli25Al'2 {**!»&„ Bridges, Scfaults, and Curry, phenotype: Similar to gg, but both sexes fertile.
1939, Otraegic last. Wash. Y®ar Book 38: 273-77). Viability low. RK2.
®f§*«? tafor»#tie«: Causes local shortening of map by
99*
abet* 1.1 units. I* a d^fk-iesicy for or an allel* of
origin: Spontaneous.
it.
discoverer: Mohler, 54J28.
references: 1956, DIS 30: 78.
!te » of 0.
phenotype: Similar to gg3. Varies from eyelessness
in first flies to emerge to nearly normal in aging
MUTATIONS 105

cultures. Always s e p a r a b l e from wild type on phenotype: Like gl3, perhaps with eye color more
basis of p e b b l y a p p e a r a n c e of wings. RK2. normal. RK1.
giant: see gt
Giant: see Gt location: 3-64.
giantoid: see gtd origin: Spontaneous.
gl: glass discoverer: Oliver, 41el.
location: 3-63.1. synonym: gl-l: glass-like.
origin: Spontaneous. references: 1942, DIS 16: 53.
discoverer: Muller, 18b. phenotype: Eye texture smooth to rough. Eye
references: Bridges and Morgan, 1923, Carnegie smaller than wild type. Eye color orange; lighter
Inst. Wash. Publ. No. 327: 188 (fig.). around rim of eye. Viability and fertility good.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. RK2.
Genet. 2: 214 (fig.), 226. *glS1k
phenotype: Eyes reduced to one-half normal area; origin: Spontaneous.
diamond shaped with glassy texture from fused discoverer Edmondson, 51k.
facets and irregular surface. Eye color blotchy, references: 1952, DIS 26: 60.
ranging from scarlet to orange in males and orange phenotype: Like gl. RK2.
to apricot in females; colorless rim and eroded *glS4g
patches, especially in female. Malpighian tubes of origin: Spontaneous.
larva paler than wild type (Brehme). RK2. discoverer: Hexter, 54g.
cytology: Located in region between 90C2 and 91A3 references: 1956, DIS 30: 72.
on basis of its inclusion in Df(3R)P14 = phenotype: More extreme than gl. Facet area less
Df(3R)90C2-Dl;91A2-3 (E. B. Lewis). than one-half of normal surface; very irregular.
gl2 Eye color white with some orange specks in both
origin: Spontaneous. sexes. Ocelli colorless. Malpighian tubes normal.
discoverer: R. L. King, 1927. RK1.
phenotype: Slightly less extreme than gl. Eyes gi62d
reduced to two-thirds normal size; ovoid; glassier origin: Spontaneous.
and smoother than gl. Eye color blotchy scarlet in discoverer: Tano, 62d.
both sexes, with rim and eroded patches of color- references: Burdick, 1963, DIS 37: 47.
less material. Ocelli colorless and papillose. phenotype: Less extreme than gl2. Viability normal.
Larval Malphigian tubes normal yellow (Brehme). RK1.
Cephalic complex smaller than wild type at 36 hr *gl63a14
after hatching (27°C); growth rate subsequently origin: Spontaneous.
normal [Medvedev, 1935, Z. Induktive discoverer: Ashburner and Hudson, 63al4.
Abstammungs- Vererbungslehre 70: 55—72 (fig.)J references: 1966, DIS 41: 60.
1935, Tr. Inst. Genet. Akad. Nauk SSSR 10: 119- phenotype: Similar to gl, but eye color darker and
5l]. RK2. pigmented area larger. Male eyes darker than fe-
9I3 male. Malpighian tubules wild type, gl63*14/gl
origin: Spontaneous. has more eye pigment than either homozygote. Via-
discoverer: Stern. bility and fertility good. RK1.
synonym: rh: rauhig. gl63d
references: Csik, 1929, Biol. Zentr. 49: 419—21. origin: Gamma ray induced.
phenotype: Eye small and elliptical, with surface discoverer: Ives, 63d29.
less rough than gl or gl2. Eye color homogeneous references: 1965, DIS 40: 35.
scarlet in both sexes. Ocelli colorless. Viability phenotype: Eyes small, nearly colorless, with a
and fertility good. Malpighian tubes of larvae wild typically glassy surface. Viability of homozygote
type (Brehme). RK1. good except in competition with other genotypes.
*gt4 RK2A.
origin: Spontaneous. cytology: Associated with T(2;3)gl63d; breakpoints
discoverer: Villee, 40d. unknown.
references: 1941, DIS 14: 40. *qJ63f6
1942, Univ. Calif. (Berkeley) Publ. Zool. 49: 137. origin: Spontaneous.
phenotype: More extreme than gl. Eyes reduced to discoverer: Ashburner and Hudson, 63f6.
less than one-half normal area; ovoid; narrower, references: 1966, DIS 41: 60.
glassier, and smoother than gl. Eye color white to phenotype: Like gl63t**4 and may be identical. RK1.
apricot in females, pink in males. Ocelli colorless. Gl: Glued
Viability about 65 percent wild type. RK2. location: 3-41.4 [0.9 unit from Ly (Mossige, 1935,
*gl40h DIS 4: 59; 1938, Hereditas 24: 110-16)1.
origin: Spontaneous. origin: Recovered among progeny of heat-treated
discoverer: Ives, 4Qh. flies,
references: 1941, DIS 14: 39. discoverer: Ives, 31 f5.
106 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: 1934, DIS 1: 34. references: Eyes rough but of normal size; facets
1934, DIS 2: 35. and hairs irregular. RK2.
Plough and Ives, 1935, Genetics 20: 42—69 (fig.) Go; Gold tip
phenotype: Eyes rough, smaller, and oblong; facets location: 2-64.3 (57.5 to 71.1; between en and eg).
rounded; surface smooth and shiny like gl. origin: Spontaneous.
Bristles generally shortened slightly and straighter discoverer: Sturtevant, 1948.
than normal. Viability and fertility good. references: 1948, DIS 22: 55.
Homozygous lethal. RK1. phenotype: Tips of many bristles and hairs pale and
cytology: Salivary chromosomes normal (Bridges in curved. Bristles often short (tips broken off ?).
Morgan, Bridges, and Schultz, 1937, Carnegie Inst. Wild-type bristles sometimes have pale tips, thus
Wash. Year Book 36: 301). interfering with positive classification. Lethal
gl-1: see gl*i« when homozygous. Expression best at low temper-
atures. RK2.
Gla: Glazed goggle: see gg
location: 2- (rearrangement). Gold tip: see Go
references: Morgan, Bridges, and Schultz, 1936, gouty legs: see gy
Carnegie Inst. Wash. Year Book 35: 293.
phenotype: Eye reduced to one-fourth normal area
and narrowed to a point ventrally. Eye color gen-
erally diluted but with some black patches. Omma-
tidia coalesce into gleaming, smooth sheet. Mal-
pighian tubes of larva somewhat lighter than wild
type; difficult to classify (Brehme and Demerec,
1942, Growth 6: 351—56). Homozygous lethal.
RK2A.
cytology: Associated with In(2LR)Gla =
In(2LR)27D;51E, superimposed on In(2L)Cy =
ln(2L)22Dl-2;33F5-34Al or In(2L)t = gp- gap
In(2L)22D3-El;34A8-9. From Bridges and Morgan, 1919, Carnegie Inst. Wash.
glass: see gl Publ. No. 278: 209.
glass-like: see gl41 e
Glazed: see Gla gp; gap
gleam: see gm location: 2-74.
*gli: glide origin: Spontaneous.
location: 1-38.0. discoverer: Bridges, 12alO.
origin: Induced by DL~p-NN-di-(2-chlorethyl)amino- references: Bridges and Morgan, 1919, Carnegie
phenylalanine (CB. 3007). Inst. Wash. Publ. No. 278: 208 (fig.).
discoverer: Fahmy, 1954. Morgan, Bridges, and Sturtevant, 1925, Bibliog.
references: 1958, DIS 32: 70. Genet. 2: 212 (fig.), 226.
phenotype: Wings held horizontally at right angles phenotype: Vein L4 weak or has section missing
to body. Pigmentation of tergites frequently inter- beyond posterior crossvein. Overlaps wild type
rupted along mid-dorsal line; tergites occasionally when homozygous; semidominant as heterozygote.
show a nick in the posterior border. Males sterile; RK3.
viability about 70 percent wild type. RK2. *gr: gracile
glisten: see gn location: 1-36.4.
glossy: see Iz origin: Induced by L-p-NN-di-2-chloroethyl)amino-
Glued: see Gl phenylalanine (CB. 3025).
gly: see Iz6 discoverer: Fahmy, 1953.
gm: gleam references: 1959, DIS 33: 86.
location: 3- (not located), phenotype: Small fly with narrow abdomen. Wings
origin: Spontaneous, frequently held atypically, either upward or down-
discoverer Bridges, 27cl. ward. Very inviable, many dying less than 24 hr
phenotype: Eyes small and rough; irregular hairs after eclosion; males sterile. RK3.
and facets cause glints. Body small. Viability *gre: green body color
about 10 percent wild type but variable. RK3. location: 1- (not located).
cytology: Associated with In(3L)P, according to origin: Spontaneous.
Bridges (Morgan, Bridges, and Schultz, 1937, discoverer: Bridges, 13e.
Carnegie Inst. Wash. Year Book 36: 301). references: Morgan and Bridges, 1916, Carnegie
*gm: glisten Inst. Wash. Publ. No. 237: 73.
Socotlon: 3-67.3. phenotype: Body color tinged greenish black, with
origin: Gamsaa my induced. marked trident pattern. Overlaps wild type. RK3.
r. WcUbrun, 61i6.
other Information: Possibly an allele of ptg.
MUTATIONS 107

gro: groucho gland chromosomes of double thickness in some


location: 3-90 (no recombinants with Pr obtained). cells (Bridges, 1935, J. Heredity 26: 60-64). RK3.
origin: Spontaneous. other information: Used by Bridges (1935) in the
discoverer: E. H. Grell, 64k. construction of salivary chromosome maps.
phenotype: Clumps of extra bristles above each eye *Gt-2: Giant in chromosome 2
give impression of bushy eyebrows; also extra location: 2- (not located),
bristles on humerus. Top of head tends to be mal- origin: Spontaneous,
formed; ocelli often enlarged and run together. In discoverer: Bridges, 14128.
selected stocks, penetrance approaches 100 per- phenotype: Heterozygote normal but, in presence of
cent but is low in unselected stocks. Probably no homozygous gt-3, gives giant male-sterile flies.
gross chromosomal rearrangement since recombina- Homozygous lethal. RK3.
tion is normal in vicinity of mutant. RK2. *gt-3: giant in chromosome 3
location: 3-64.
grooved: see gv origin: Spontaneous.
groove/ess: see gvl discoverer: Bridges, 14i28.
groucho: see gro references: Bridges and Morgan, 1923, Carnegie
Inst. Wash. Publ. No. 327: 120 (fig.).
gs: gespleten phenotype: Body size much larger than normal. Late
location: 3-35.1. hatching. Entirely sterile in male. Giant character
origin: Spontaneous. produced only in flies homozygous for gt-3 and
discoverer: Smelink-den-Hollander, 561. heterozygous for Gt-2. RK3.
references: 1957, DIS 31: 85. gt-4
phenotype: Thorax cleft by medial groove. Eyes location: 2-24.0.
small; sometimes missing. Hair pattern on thorax origin: Spontaneous,
disturbed. Viability and fertility excellent. RK1. discoverer Bridges, 30bl4.
other information: Probably an allele of gv (3-36.2). phenotype: Giant flies hatch very late. Viability
variable but around 15 percent wild type. RK3.
*gtd: giantoid
location: 1-0.5.
origin: Spontaneous.
discoverer: Bridges, 21cl2.
references: Bridges and Gabritchevsky, 1928, Z.
Induktive Abstammungs- Vererbungslehre 46: 232
(fig.).
phenotype: Body size larger, especially head. Late
hatching. Viability erratic, about 50 percent wild
type. Separation difficult in females, easier in
males. RK3.
Gull: see G
Gulloid: see Gd
*gv: grooved
gt: gianf location: 3-36.2.
Left: wild-type female. Right: giant female. origin: Spontaneous,
From Bridges and Gabritschevsky, 1928, Z. Induktive discoverer Ives, 43128.
Abstammungs- Vererbungslehre 46: 231—47.
references: 1946, DIS 20: 65.
phenotype: A longitudinal medial groove in thorax;
in extreme individuals, thorax nearly cleft. Eyes
gt: giant
reduced. Irregular and often extra alar bristles.
location: 1-0.9.
Viability good. RK1.
origin: Spontaneous.
other information: Probably allelic to &*.
discoverer: Gabritschevsky, 25i2.
references: Bridges and Gahritschevsky, 1928, Z.
Induktive Abstammungs- Vererbungslehre 49: gvl: groove/ess
location: 4-0.2 fin diplo-4 triploids (Sturtevant,
231-47 (fig.).
1951, Proc. Natl. Acad. Sci. U.S. 37: 405-7)].
Gabritschevsky and Bridges, 1928, Z. Induktive
origin: Spontaneous.
Abstammungs- Vererbungslehre 49: 248—84.
discoverer: Bridges, 33elO.
phenotype: Larval development 4 days longer than
references: 1935, Biol. Zh. (Moscow) 4: 401-20.
normal, resulting in giant larvae, pupae, and
phenotype: Sharp transverse groove between scu-
imagos. Adult weight 1.7 times normal. But not all
tellum and thorax is nearly eliminated; no overlap
genetically giant flies show the character; the rest
of wild type. Black scars appear on scutellar
have normal size. Distribution sharply bimodal.
groove at si«4e&, in pleural region, and behind
Percentage giant greatest in well-fed cultures;
stemopletirais. Viable and fertile. RK1.
also raised by modifying action of bh11. Salivary
108 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

gy: gouty legs (Stern, 1944, DIS 18: 56). Shown byboth Sturtevant
location: 4- (not located). and Rasmussen to recombine with and He to the
origin: Spontaneous. right of h2.
discoverer: Muller. h2
references: 1965, DIS 40: 36. origin: Spontaneous.
phenotype: Legs shortened and thickened, espe- discoverer: Bridges, 28d23.
cially the metatarsi of the hind legs, which are phenotype: Extra hairs on wings, but fewer than in
often swollen. Usually classifiable; viability and h. Extra hairs not present on scutellum, pleurae,
fertility good. gy/eyD is gy. RK2. or top of head. Bristles cylindrical, with javelin
heads. Some abnormal abdomen effect. RK2.
other information: Shown by both Sturtevant and
Rasmussen to recombine with and lie to the left of
h.
*h!00.12
origin: X ray induced.
discoverer: Alexander.
references: Ward and Alexander, 1957, Genetics 42:
42-54.
phenotype: Homozygous viable. RK2A.
cytology: Associated with In(3L)h100-12 =
In(3L)61A2'3;66D.
*/,700.239
origin: X ray induced.
discoverer: Alexander.
references: Ward and Alexander, 1957, Genetics 42:
42-54.
phenotype: Homozygous semilethal. RK2A.
cytology: Associated with In(3L)h100'239 =
In(3L)66Dll-12;80C.
*hl00.271
h: hairy origin: X ray induced.
From Bridges and Morgan, 1923, Carnegie Inst. Wash. discoverer: Alexander,
PubJ. No. 327: 202. references: Ward and Alexander, 1957, Genetics 42:
42-54.
h: hairy phenotype: Homozygous lethal. RK2A.
location: 3-26.5. cytology: Associated with T(2;3)ht00.271 =
origin: Spontaneous. T(2;3)41;66D14'E1.
discoverer. Mohr, 18111. */,7OO.39O
references: 1922, Z. Induktive Abstammungs- origin: X ray induced.
Vererbungslehre 28: 17. discoverer: Alexander.
Bridges and Morgan, 1923, Carnegie Inst. Wash. references: Ward and Alexander, 1957, Genetics 42:
Publ. No. 327: 202 (fig.). 42-54.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Homozygous lethal. RK2A.
(tenet. 2: 214 (fig.), 227, cytology: Associated with Df(3L)h100'390 =
Neel, 1941, Genetics 26: 52-68 (fig.), Di(3L)66D2'5;66D14-El.
pfoenotype: Extra hairs cm scutellum, along veins, ft*; hairy-subliminal
on pleorae, and on top of head. Additional hair- origin: X ray induced.
forming cells present in 19-hr pupa (Lees and discoverer: Green.
Waddiagtoa, 1942, DIS 16: 70). Interacts synergis- references: 1960, Proc. Natl. Acad. ScL U.S. 46:
tically with ppd and Hw or Hw variegateds [e.g., 524-28.
JTnf/)*e*]; In{l)sc7 suppresses h (Steinberg, 1942, phenotype: Homozygote nearly lethal but has no h
DIS 16: 68; Neel, 1941). RK1. phenotype. Heterozygote with h and h2 also wild
ey*of*§y: Pla-eed is aalivsry chromosome region be- type. he/+ has extra hairs on wings, head,
tween 66D2 and 66E1, on basis of its inclusion in pleurae, halteres, and occasionally on scutellum if
DK3LftteO.3f0 mDI(3L)6$D2-5;66DI4'E1 (Ward also heterozygous for certain X-chromosome inver-
•ad AUxmndm, 1957, Genetics 42: 42-54). sions that variegate for Hw, including In(l)scs,
e*h«r in§ormotion: As with c/ + , expression of h + may In(l)acS*, and ln(l)y3P. Presence of y+Y also in-
b® altered i© tht direction of ft by rearrangements duces extra hairs, RK3.
with bmstkm in the vicinity of the h locus [Dubinin H: Hairless
@«j Sldorov, 1934, Biol. Zh. (Moscow) 3: 307-31 ]. location: 3-69.5.
Unlike tike ci case, however, rearranged ft chrotno* origin: Spontaneous.
do act sfeow evidence of altered gene action discoverer: Bridges, 16c4.
MUTATIONS 109

references: Bridges and Morgan, 1923, Carnegie references: Plunkett, 1926, J. Exptl. Zool. 46:
Inst. Wash. Publ. No. 327: 161 (fig.). 181-244.
Morgan, Bridges, and Sturtevant, 1925, Bibliog. phenotype: Bristle effect more extreme than in H
Genet. 2: 170 (fig.), 227. and more easily separated from wild type. Venation
Nash, 1965, Genet. Res. 6: 175-89. effect slighter than in H. Interactions with mutants
phenotype: Bristles, especially postverticals and at the N locus similar to those of H (Von Halle).
abdominals, missing. Bristle sockets present at Homozygous lethal. H2/H lethal. RK1.
some sites, not at others. Veins L4 and L5 do not H3
reach wing margin; occasionally true of L2 also. discoverer: Sturtevant.
Eyes larger than wild type; body color somewhat phenotype: Like H. RK1.
paler. Lees and Waddington [l942, Proc. Roy. Soc. 4
(London), Ser. B. 131: 87-110 (fig.)J show that origin: Spontaneous.
trichogen cell forms a socket instead of a bristle at discoverer: Bridges, 30b20.
some sites. Interactions with other mutants studied phenotype: Like H. RK1.
by House (1953, Genetics 38: 199-215, 309-27;
1959, Genetics 44: 516; 1955, Anat. Record 122: origin: Gamma ray induced.
471; 1959, Anat. Record 134: 581-82). H sup- discoverer: Ives.
presses wing notching of /V, fa, fano, and nd; en- references: 1959, DIS 33: 95.
hances Ax, and enhances eye effect of spl; H re- phenotype: Extreme bristle effects; anterior lateral
moves more bristles in combination with spl acrostichals removed. L5 incomplete distally.
(House, Von Halle). Shows some superadditive in- RK2A.
teraction with en, ci, ciw, and ci® relative to de- cytology: Shown genetically to be associated with
gree of L4 interruption. L2 interruption augmented
in combinations with ve and ri; L3 interruption aug- other information: Allelism to H inferred from
mented in combinations with ve and tt. Triploid, phenotype.
H/+/+, intermediate between wild type and H/+. Hairless of Dobzhansky
H/H/+ most extreme type, with bristles absent origin: X ray induced,
from head, thorax, and abdomen LGowen, 1933, Am. discoverer: Dobzhansky, 1930.
Naturalist 67: 178—80 (fig.)J. Homozygous lethal. phenotype: Slight allele of H with no shortening of
RK1. L4 or L5. Bristle effect slighter, particularly on
cytology: Salivary chromosomes normal. abdomen. Homozygous lethal. RK2.

*HP2: Hairless from P32


origin: Found among descendants of male fed P 3 2 .
discoverer: Bateman, 1949.
references: 1950, DIS 24: 55.
phenotype: Bristle effect like H, but venation quite
distinct. Veins not interrupted, but knotted. Homo-
zygous lethal. RK1.
*ha: hair bristles
location: 1-22.7.
origin: Induced by L-p-NN-di-(2-chloroethyl)amino-
phenylalanine (CB. 3025).
discoverer. Fahmy, 1954.
references: 1958, DIS 32: 70.
phenotype: Small fly with extremely fine, short
bristles. Males viable and fertile. Females less
viable and highly infertile. RK3.
Hairless: see H
hairy: see h
Hairy wing: see Ww
Haltere mimic: see Hm
hdp: holdup
location: 1-59.5.
origin: Induced by DL-p-NN-di-{2-chloroethyl)amino-
H: Hairless phenylalanine (CB. 3007).
From Bridges and Morgan, 1923, Carnegie Snst. Wash. discoverer: Fahmy, 1954.
Publ. No. 327: 161. references: 1958, DIS 32: 70.
phenotype: Wings held up to various degrees. May
overlap wild type. Viability and fertility good in
origin: Spontaneous. both sexes. RK2.
discoverer: Stwrtevant. other information; One X-ray-induced allele.
no GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

*/i</prwg; heldup-reduced wings roughened. Wings usually divergent and may be


origin: Induced by 2-chloroethyl methanesulfonate smaller. Body sometimes small and chunky. Less
(CB. 1506). extreme expression at 19°C, especially the wing
discoverer: Fahmy, 1956. character, but classification reliable. RK2.
synonym: rwg. cytology: Salivary chromosome locus between 37B2
references: 1958, DIS 32: 74. and 40B2 (Schultz and Curry).
phenotype: Wings short; upheld or outspread to
various degrees. Small regions of deranged wing
hairs, giving shaded streaks parallel to longitu-
dinal veins. Male infertile; viability about 70 per-
cent normal. RK2.
heavy vein: see hv
holdout', see ho
heldup: see hdp
Henna: see Hn
Hermaphrodite: see traD
*hi: high
location: 2- (not located).
origin: Found in Florida natural population.
discoverer: Ives, 1943.
references: 1943, Genetics 28: 77.
1950, Evolution 4: 236-52.
phenotype: Male homozygous for hi produces sperm
containing 10 times normal frequency of mutations.
Heterozygous hi/+ causes a mutation rate 2—7
times normal. Ratio of sex-linked lethal to visible
mutations about 8 to 1. Inversions associated with
about 5 percent of mutations. RK3.
cytology: Salivary chromosomes normal.
other information: Homozygous hi constructed by hk: hook
crossing two balanced lethal stocks, 11 M/CyX From Mohr, 1927, Hereditas 9: 169-79.
12 hi/Cy. Since these stocks have developed a hk2
common lethal, it is now difficult to obtain hi origin: Spontaneous.
homozygotes. discoverer: Bridges, 33a31.
*Hi: Hirsute phenotype: Bristles cylindrical, javelinlike, or bent,
location: 3- (rearrangement). but effect less extreme than hk. RK2.
origin: X ray induced. Hm: Haltere mimic
discoverer: Bishop, 1939. location: 2- or 3- (rearrangement).
phenotype: All bristles except postscutellars and origin: X ray induced.
postdorsocentrals multiplied, especially on head discoverer Slatis, 49b5.
and anterior thorax. Eyes smaller and facets phenotype: Wing resembles a large haltere, but stalk
irregular. Homozygous lethal. RK2A. is more winglike. In combination with vg, wing is
cytology: Associated with In(3LR)Hi - further reduced and haltere almost completely
In(3LR)71A;91F. eliminated. Homozygous lethal. RK3A.
Hia: Hiatus cytology: Appears to be associated with complex
location: 2- (not located), translocation, T(2;3)Hm.
origin: Spontaneous. tin31: s e e UnT<*
discoverer: Bridges, 2%12. Hn; Henna
phenotype: Terminal interruption of L2. More location: 3-23.0.
obvious in heterozygous male than in heterozygous origin: X ray induced.
female. Homozygous viable. RK3. discoverer: Van Atta, 30k.
high: see hi references: 1932, Am. Naturalist 66: 93—95.
Hirsute: see Hi 1932, Genetics 17: 637-59.
hk: hook phenotype: Eye color homogeneous dark, dull brown.
location: 2-53.9. Classification difficult; best in aged flies. Homo-
origin: Spontaneous. zygous lethal. Heterozygotes viable and fertile.
discoverer; Mohr, 24a4. RK2A.
references: 1927, Hereditas 9: 169-79 (fig.)- cytology: Placed in 66A-B, on basis of its associa-
pltttnotyp*: Bristles nearly ail hooked at tip or tion with Df(3L)Hn = Df(3L)66A;66B (Lewis, 1956,
blunted; some bent at right angles. Scute liars and DIS 30: 130). Hn was induced together with, but is
verticals especially affected. Acrostichal hairs separable from T(2;3)Hn = T(2;3)53E-
fewer mad outer rows separated. Eyes slightly 54A;77A;94F;96A.
MUTATIONS in
*HnS3k references: Novitski and Rifenburgh, 1938, Proc.
origin: Ultraviolet induced. Indiana Acad. Sci. 47: 256-60.
discoverer: Meyer and Verderosa, 53k. phenotype: Wings extended at right angles to body.
references: Meyer, 1954, DIS 28: 76. RK1.
phenotype: Heterozygote has brick-red eye color, cytology: Located in or near 22E (Lewis, 1945,
but classification difficult. Homozygote viable; Genetics 30: 137-66).
eye color dark brown like sepia. RK1 as recessive.
Hnr: Henna-recessive origin: X ray induced.
origin: Spontaneous. discoverer: E. B. Lewis, 1940.
discoverer. Bridges, 33c20. synonym: In-ho.
references: Mohr, 1937, DIS 8: 12. references: 1945, Genetics 30: 137-66.
phenotype: Eye color dull, dark brown, like cl. Best phenotype: Wings reduced to tiny stubs. Eyes
separability in aged flies. Hnr/Hn more extreme reduced, with anterior indentation. Male lacks
than Hnr/Hnr. Eye color autonomous in transplant genitalia and anal apparatus; female fertile.
into wild-type larval host (Beadle and Ephrussi, ho^O/ho resembles ho/ho. RK2A.
1936, Genetics 21: 230). Larval Malpighian tubes cytology: Associated with In(2L)ho'*0 =
bright yellow as in wild type (Beadle, 1937, In(2L)21D4-El;22E2-3.
Genetics 22: 587—611). Eyes have 58 percent hook: see hk
normal red pigment and 120 percent normal brown Hooked veins: see Hv
pigment (Nolte, 1954, J. Genet. 52: 127-39). RK2. hp: humped
location: 3- (rearrangement).
discoverer: Nordenskiold, 39b9. origin: Spontaneous.
synonym: bu; brunette; hn^. discoverer: Bridges, 31a22.
references: 1937, DIS 7: 18. phenotype: Thorax shortened and strongly humped,
phenotype: Eye color darker than HnT in young flies. with thoracoscutellar groove almost absent. Eyes
RK2. sharply reduced; may be absent at 19°C. Bristles
Hnr3 Minutelike and occasionally missing. Viability 10
origin: Spontaneous. percent wild type. RK3A.
discoverer: Weinstein, 1927. cytology: Associated with In(3R)hp.
synonym: sed: sepiaoid. *hpa: hyperantenna
phenotype: Eye color dull chocolate. Classification location: 1-50.1.
easier than for Hnr. 79 percent normal red pigment, origin: Induced by DL-p-NN-di-(2-chloroethyl)amino-
100 percent normal brown pigment (Nolte, 1955, J. phenylalanine (CB. 3007).
Genet. 53: 1—10). Eye color autonomous in trans- discoverer: Fahmy, 1954.
plants of optic disk into wild-type hosts (Beadle references: 1959, DIS 33: 86.
and Ephrussi, 1936, Genetics 21: 230). Larval phenotype: Antennae enlarged or have duplicated
Malpighian tubes bright yellow as in wild type parts, sometimes an extra antennal base near the
(Beadle, 1937, Genetics 22: 587-611). RK2. eye. Grossly deformed head and eyes. Wings have
other information: Allelism by Lewis (1956, DIS 30: rounded tips and incised inner margins. An occa-
130). sional bristle absent or shorter. Phenotype vari-
*Hn'53/ able and minimal expression slightly altered eye
origin: Ultraviolet induced. shape and blunt wing tips. Males viable and
discoverer: Meyer, 53j. infertile; females sterile. RK3.
references: 1954, DIS 28: 76. Ht: see traD
phenotype: Similar to se. RK1. Hu: Humeral
*Hnrh Henna-recessive of Ives location: 3-51 (48-54).
origin: Spontaneous. origin: X ray induced.
discoverer: Ives, 45J17. discoverer: Ruch, 1931.
references: 1946, DIS 20: 65. phenotype: Extra bristles on humeral patches of
phenotype: Eye color brownish at hatching, darkens Hu/+. Humeral bristles more numerous in homozy-
to black. Resembles se. Viability good. RK1. gote, with a streak of hairs below humerus toward
*HnrN: Henna-recessive from Nebraska base of first leg. Hu/Hu has viability 90 percent
origin: Spontaneous. wild type. RK2A as heterozygote.
discoverer: Williamson, 53j. cytology: Associated with In(3R)Hu = In(3R)84B2-
references: 1955, DIS 29: 75. 3;84F2-3;86B4-C1.
phenotype: Indistinguishable from se. Larval Mal- humped: see hp
pighian tubules somewhat darker yellow than wild humpy: see hy
type. RK1. hv: heavy vein
ho: heldout location: 2-104.0.
location: 2-4.0. discoverer: Curry, 36115.
origin: Spontaneous. phenotype: Veins thick and knotty, especially at
discoverer Novitski, 35g. ends of crossveins; posterior crossvein oblique
112 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

and may show break in middle; extra cross veins Hw*


sometimes present. Wings broad, thick, dark, origin: Spontaneous derivative
warped, divergent, and droopy. Eyes small and discoverer: Nichols-Skoog, 35a9.
bulging. Posterior scutellars blunt, short, and phenotype: Females homozygous for Hw2 show only
crossed. Overlaps wild type at 25°C but useful at occasional extra hairs along wings. Overlaps wild
19°C. RK2. type. RK3A.
*Hv: Hooked veins cytology: Salivary chromosome analysis by Schultz
location: 1-66. (Morgan, Schultz, and Curry, 1941, Carnegie Inst.
discoverer: Tanaka, 35a4. Wash. Year Book 40: 284) shows small inversion
references: 1935, DIS. 4: 16. of the region from 1A3 through 1B1 of the first
1936, D1S 5: 8. 1B1-2 doublet of Dp(l;l)Hw; i.e., associated with
1937, DIS 8: 11. In(l)Hw2 =In(l)lA2-3;lBl-2.
phenotype: Heterozygous female shows small Hw49c
branches from posterior crossvein and L5. Eyes origin: Induced by P32 simultaneously with sc45<=.
small and rough. Homozygous female lethal. discoverer: R. C. King, 49c21.
RK3A. references: Poulson and King, 1949, DIS 23: 62-63.
cytology: Associated with In(l)Hv. phenotype: More extreme than Hw. Homozygous
female has doubling and tripling of many bristles;
extra wing veins; gap in posterior crossvein; extra
hairs on vein L2 and in wing cells. Heterozygous
female has normal bristles but extra hairs on L2
and L3 and in wing cells; often an extra free vein
from posterior crossvein. Hw4^c male much like
homozygous female, but bristle duplication less
extreme. Male and heterozygous female fertile;
homozygous female sterile. RK1.
Hx: Hexaptera
location: 2- (not located).
origin: Spontaneous.
discoverer: Herskowitz, 47j.
references: 1949, Genetics 34: 10-25 (fig.).
phenotype: Expression same in Hx/-f and Hx/Hx;
varies from absence of a detectable difference from
normal, through various intermediate types, to
presence of large appendage on prothorax. Entire
abnormal structure may remain beneath exoskeleton.
Appendage varies from small amorphous mass to
highly differentiated wing. May also produce
haltere- and leglike appendages. Penetrance same
Hw: Hairy wing in homozygote and heterozygote; enhanced by
Edith M. Wallace, unpublished. crowding and by high temperature (20°C, male 1.5
Hw: Hairy wing and female 3.3 percent; 25°, male 6.5 and female
location: 1-0.0. 24.2 percent); affected by genotype, e.g., sup-
origin: Spontaneous in stock containing y and never pressed by In(2L+2R)Cy and by ln(2LR)bwvl.
separated. RK3.
discover**; Bridges, 23cl2. hy; humpy
reference*: N«el, 1941, Genetics 26: 52-68 (fig.). location: 2-93.3.
pHenotyp*: M®ie has extra bristles along wing vein, origin: Spontaneous.
©a btftd (especially occipitals), and on thorax. discoverer: Bridges, 18j22.
Ala©, extra hairs on wing veins, back of head, and references: 1937, Cytologia (Tokyo), Fuji! Jub. Vol.
SMKopletvae. Homoxygous female sterile, expres- 2: 745-55.
sion men mximmm than male, and has 40—80 per- phenotype: Thorax strongly ridged, with commas
cent wiid-typ® viability. Heterozygous female has anteriorly and two pairs of vortices. Wings
good viability. Phenotype similar to male. Classi- obliquely truncated to one-half normal length. An
fiable an stagte do«# in triploids (Schultz, 1934, irregular contraction of larval muscles at time of
DB 1: 55). Interacts synergistically with ft and pupation (Waddington, 1941, Proc. Zool. Soc.
Pf4 (M«®!( 1941). Suppressed by su(Hw) and London Ser. A 111: 181-88). Viability low and
mu(Hw)2. REtA as heterozygous female and as erratic. Both sexes highly infertile. RK2.
cytology: Placed in region 57 on basis of its being
cyteleoy: Salivary chromosome analysis by Denserec to the right o(In(2R)NS =In(2R)52A2-Bl;56F9-13
cad Hoover shows repeat for doublet 1B1-2, i.e., and to the left of Df(2R)M'l = Di(2R)57Fll-
Dp(lilti»l\9*9, Gteetics 24: 68). 58A1;58F8-59A1 (Bridges, 1937).
MUTATIONS 113

apex. Eyes smaller and sometimes slightly rough.


Bristles thin. Males small, late eclosing; viability
reduced. Female sterile. RK3.
in: inturned
location: 3-47 (left of centromere).
origin: Spontaneous.
discoverer: Bridges, 26k20.
phenotype: Hairs and bristles on thorax directed
irregularly toward midline. Marginal hairs of wing
stand out from wing margin; wings slightly spread
and tend to be long and narrow. RK1.
cytology: Tentatively placed in salivary region
77B-C (Hannah and*Arajarvi).
//?-/: see E(f)
In-ho: see ho40
*inb: incised balloon
location: 2-55.
hy: humpy origin: Spontaneous.
Edith M. Wallace, unpublished. discoverer: Neel, 41d9.
references: 1942, DIS 16: 50.
hyperantenna: see hpa phenotype: Wings held at 45° angle to body. Wing
margins incised, varying from slight nicks to
i: see E(B) extreme reduction to small fluid-filled sacs. RK2.
I: see E(B) cytology: Salivary chromosomes normal.
Ic D: see bwVD *lnd: Indented
*if: inflated location: 2-63.
location: 1-55. origin: Spontaneous.
origin: Spontaneous. discoverer Cole, 40e.
discoverer: Weinstein, 1916. references: Whittinghill and Parker, 1945, Genetics
references: 1918, Genetics 3: 157 (fig.). 30: 27-28.
phenotype: Wings inflated with lymph and smaller Whittinghill, 1947, DIS 21: 72.
than normal; venation defective. Wings later dry phenotype: Eye usually kidney shaped with indenta-
and blistered. Viability and fertility poor. RK3. tion anteriorly; shape sometimes normal, but facets
if* irregular. Often indented posteriorly as well as
origin: Spontaneous. anteriorly, sometimes dividing eye into two spots,
discoverer: Curry, 38b. or with only upper lobe persisting. Rarely eyeless.
references: 1939, DIS 12: 45. More extreme at 28° than at 25°C. RK2.
phenotype: Resembles if. Longitudinal veins inflated: see if
thickened, especially at wing base. Anterior infra bar: see B'
cross vein thickened. In most cases, wings inflated infrabar Bat: see BBi
or blistered. RK2. intensifies: see e( )
If: Irregular facets Intensifiet: see E( )
location: 2-107.6 (0.6 unit to the right of sp, ac- interrupted margin: see im
cording to Ives). Interruptus: see ciw
origin: Spontaneous. inter sex: see ix
discoverer: Casey, 65116. intersex on chromosome 3: see dsx^oi
phenotype: In heterozygote, eye area about one-half intersex-62c: see dsx
of normal; narrow and pointed ventrally; facets ir- inturned: see in
regular and often missing across middle of eyes, Irregular facets: see If
sometimes fused or absent in ventral portion. In It: see ciw
homozygote, eyes are narrow slits, with smooth ix: intersex
glossy surface. Viability and fertility good. RK1. location: 2-60.5.
*im: interrupted margin origin: Spontaneous.
location: 1-3.1. discoverer: L. V. Morgan, 1943.
origin: Induced by 2-chloroethyl methanesulfonate references: Morgan, Redfield, and Morgan, 1943,
(CB. 1506). Carnegie Insst. Wash. Year Book 42: 171-74.
discoverer: Fahmy, 1956. Kroeger, 1959, Arch. Entwicklungsmech. Organ.
references: 1959, DIS 33: 86-87. 151: 301-22 (fig.).
phenotype: Wing margin nicked to various degrees, phenotype: Females changed into sterile inters exes
with costal vein frequently interrupted. Extra wing with a set of reduced male and a set of irregular
venation often present, and occasionally anasto- female external genitalia. Gonads also mixed.
moses, giving a plexus, particularly at the wing They have no sex combs; pigmentation of abdomen
114 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

intermediate between male and female. A large references: 1927, Biol. Zentr. 47: 361-69.
mass of chitinized tissue protrudes from vaginal 1934, DIS 1: 35.
opening. Males not affected. RK2. phenotype: Wings upturned but less extreme than ;.
iV Curl of wings dependent on hatching and unfolding
origin: Ultraviolet induced. at 25°C or higher. RK3.
discoverer: Meyer, 50k. *j49j
synonym: torn: tomboy. origin: Spontaneous.
references: Meyer and Edmonds on, 1951, DIS 25: 73. discoverer: Mossige, 49J18.
Meyer, 1958, DIS 32: 83. references: 1947, DIS 25: 69.
phenotype: Females homozygous for ix2 have male- phenotype: Less extreme than ;. Some overlapping
like pigmentation of posterior tergites, rudimentary at 21°, none at 30°C. Viability and fertility good.
ovaries, and are sterile. Expression extreme and RK2.
viability reduced at 27°C; at 17°C, expression iSO*
less extreme but viability greater. Homozygous origin: Spontaneous.
males appear normal but have nonmotile sperm. discoverer: Mossige, 5Oe5.
RK2. references: 1951, DIS 25: 69.
other information: The possibility that the male phenotype: Like j49i. RK2.
sterility is at another locus has not been excluded. *j58i
ix62c: s e e fax origin: Spontaneous.
ix-3: see discoverer: Andrew, 58i.
references: 1959, DIS 33: 82.
phenotype: Expression variable, although pene-
trance complete at temperatures above 25°C. In
most-extreme cases, wings bend sharply upward in
region of anterior crossvein. A small dark blot
occurs near L3 vein at level of the deflection.
Anterior crossvein partly or wholly absent in some
cases. RK2.

J: Jammed
location: 2-41.0.
origin: Spontaneous.
discoverer: Bridges, 23d3.
phenotype: Wings often compressed into narrow
strips; sometimes filled with fluid. Alula larger
and square tipped, with clumped bristles and bare
regions. Alula modification is characteristic least
likely to overlap wild type. Completely overlaps
wild type at 19°, almost never at 28° or 30°C. Not
lethal when homozygous; viability, as in heterozy-
j: jaunty gote, about 70 percent wild type. Classifiable in
From Bridges and Morgan, 1919, Carnegie Inst. Wash. single dose in triploids (Schultz, 1934, DIS 1: 55).
Publ. No. 278: 148. RK1 at 28°-30°C; RK2 at 25°C.
cytology: Salivary chromosomes apparently normal.
/: (Bridges in Morgan, Bridges, and Schultz, 1937,
location* 2-48.7. Carnegie Inst. Wash. Year Book 36: 301).
origin: Spontaneous, J34*
A'scovemr. Bridges, 11111. origin: Spontaneous,
rtfwrmcos: Bridges mnd Morgan, 1919, Carnegie discoverer*. Duncan, 34e3.
fast. Wash. Pttbl. No. 278: 160 (fig.). phenotype: Resembles / but produces more vigorous
C!a»se<ra, 1924, J. Exptl. Zool. 38: 423-36. homozygous stock. RK1 at 28°C.
Stem, 1927, Biol. Zeatr. 47: 361-69.
phmmw«- Distal half of wing upturned. Curling is
s t r « f if wteg unfolds at 25-30°C, but weak or
wild type if wing unfolds below 25QC.

Placed to region between 34E5 and 35DI


un th* basis of its iaclaaioo in D§(2L)$4j «
Df'2Lt34E5-Fl,35C3-m (E. H. Gmll).

Jag; Jagged
<Ii£c#Y#f#r Stern. 25431.
Edith M. Wallace, unpublished.
MUTATIONS 115

Jag: Jagged origin: Spontaneous.


location: 2-54.9 (0.1 unit from Bl). discoverer: Bridges, 12f26.
discoverer: L. V. Morgan, 34b20. references: Bridges and Morgan, 1923, Carnegie
phenotype: Jag/+ has end of wing cut off; better in Inst. Wash. Publ. No. 327: 72 (fig.).
early counts and above 25°C. Jag/Jag has reduced Morgan, Bridges, and Sturtevant, 1925, Bibliog.
and roughened eyes and extremely jagged wings. Genet. 2: 214 (fig.), 227.
RK2 as heterozygote; RK3 as homozygote. phenotype: Eye size reduced by indentation of front
Jammed: see J margin. Tuft of vibrissae and hairs below eye.
jaunty: see / Variable; overlaps wild type. RK3.
jaunty x: see jyx *fc2
javelin: see jv origin: From progeny of heat-treated flies.
javelinlike: see jvl discoverer: Goldschmidt, 1927.
*je: jelly references: Gottschewski and Ma, 1937, Z.
location: 3-46. Induktive Abstammungs- Vererbungs lehre 73:
origin: Spontaneous; arose simultaneously with mu 584_97.
(3-50). phenotype: Eye reduced in size. Penetrance 50—80
discoverer: Mohr, 37121. percent. Expression variable. RK3.
references: Mossige, 1939, DIS 12: 47.
phenotype: Dark pinkish eye color. RK1. origin: Spontaneous.
jv: javelin discoverer: Gottschewski, 1937.
location: 3-19.2 (0.9 unit to left of dv). references: Gottschewski and Ma, 1937, Z.
discoverer: Mohr, 31j29. Induktive Abstammungs- Vererbungs lehre 73:
references: 1937, DIS 8: 12. 584-97.
Mohr and Mossige, 1943, Skrifter Norske phenotype: A weak allele; penetrance 10—20 per-
Videnskaps-Akad. Oslo, I: Mat.-Naturv. KL, No. cent. RK3.
7. 51 pp. (fig.). *kD: kidney-Dominant
phenotype: All bristles and hairs cylindrical instead origin: Spontaneous in chromosome containing k.
of tapered, with small enlargement before tip. discoverer: Puro, 60cll.
RK2. references: 1964, DIS 39: 65.
cytology: Placed between 64C12 and 65E1 on the phenotype: Eyes of heterozygote reduced at anterior
basis of its inclusion in Df(3L)Vn = edges. Expression variable; in extreme cases, eye
D[(3L)64C12-D1;65D2-E1. size about one-third normal. kD/k more extreme.
jvl: javelinlike Eyes of homozygote reduced about as much as
location: 3-56.7. kP/k, but occasionally one or both eyes missing;
origin: Spontaneous. antennae usually slightly deformed with thickened
discoverer: Ives, 4012. aristae. RK1.
references: 1942, DIS 16: 48. K-pn: Killer of prune
phenotype: Resembles ;v, bristles sometimes more location: 3-102.9 (0.2 to right of bv; recalculated
crooked. Viability and productivity somewhat from Sturtevant).
lower than normal. RK2. origin: Spontaneous.
*jyx: jaunty x discoverer: Sturtevant, 54a.
location: 1-24. references: 1955, DIS 29: 75.
origin: Spontaneous, 1956, Genetics 41: 118—23.
discoverer: Bridges, 14112. phenotype: No phenotypic effects, either when
phenotype: Wings curved up at tips. Viability about homozygous or when heterozygous, except that all
60 percent wild type. RK3. pn; K-pn flies die at end of second larval instar.
Kills all seven alleles of pn that have been tested.
K-pn eye disks transplanted to pn hosts develop
autonomously, as do the reciprocal transplants
(Grell, 1958, DIS 32: 123-24). RK3.
kar: karmoisin
location: 3-51.7.
origin: Spontaneous.
discoverer: Pariser.
references: Gottschewski, 1935, DIS 4: 15.
phenotype: Eye color like st but less bright. Ocelli
ic: kidney white. Eyes contain 29 percent wild-type brown
From Bridges and Morgan, 1923, Carnegie Inst. Wash. pigment (Nolte, 1954, J. Genet. 52: 111-26).
Publ. No. 327: 77. Larval Malpighian tubes considerably lighter than
wild type but difficult to classify in living larvae
k: kidney (Brehme and Demerec, 1942, Growth 6: 351-56).
location: 3-64. RK1.
116 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

cytology: Placed in region 87D-F, on basis of its in- Viability somewhat reduced but fertility near
clusion inDf(3R)ry = Df(3R)87D-E;87E-F. normal. RK1 as heterozygote.
kar2 kidney: see k
origin: Spontaneous. kidney eye: see ke
discoverer: Bridges, 38blO. Killer of prune: see K-pn
phenotype: Like kar except that larval Malpighian Kinked: see Ki
tubes are bright yellow, similar to wild type kinked femur: see kf
(Brehtne and Demerec, 1942, Growth 6: 351—56),
RK1. *kk: kinky
kar31: karmotsin-3 lethal location: 1-42.
origin: X ray induced. origin: Spontaneous.
discoverer: Schalet. discoverer: Philip.
references: 1964, DIS 39: 64. references: 1937, DIS 8: 10.
phenotype: Heterozygote of kar3I/kar resembles kar. phenotype: Bristles slightly bent or forked. RK3.
Homozygote lethal. RK2. other information: May be an allele of fw.
other information: kar-ry crossing over normal.
*ke: kidney eye KL: Male fertility complex in the long arm of the
location: 1-28.6. Y chromosome
origin: Induced by 2-chloroethyl methanesulfonate The male fertility complex of the long arm of the Y
(CB. 1506). chromosome, originally called KI by Stern (1929,
discoverer: Fahmy, 1956. Z. Induktive Abstammungs- Vererbungslehre 51:
references: 1959, DIS 33: 87. 253—353) and subsequently called KL by Brosseau
phenotype: Eyes small and extremely rough; anterior (1960, Genetics 45: 257—74), is subject to muta-
border indented, giving a kidney shape. Wings tions or deficiencies leading to male sterility,
small, abnormal, outspread, or upheld. Veins which are symbolized ms(Y)L. By complementation
thick, and often interrupted or fail to reach wing analysis, Brosseau divided KL into five different
margin, which is usually incised. Deformed complementing units designated kl-1 through kl-5.
antennae. Bristles straggly; occasionally one is By studying recombinants between the X and the
missing. Flies short lived; 50 percent die less y, i.e., detachments of attached X's, he ordered
than 24 hr after eclosion. Sterile, probably be- the complementation groups with respect to the
cause they are too weak to mate. RK3. centromere; he assigned the symbol kl-1 to the
*kf: kinked femur proximalmost and kl-5 to the distalmost.
location: 1-20.2. kn: knot
origin: Induced by DL-p-NN-di-(2-chloroethyl)amino- location: 2-72.3.
phenylalanine (CB. 3007). discoverer: Nichols-Skoog, 31hl.
discoverer: Fahmy, 1954. phenotype: Veins L3 and L4 shifted closer together
references: 1959, DIS 33: 87. in region of anterior crossvein, which is either
phenotype: Small fly with slightly dark, dull red eye extremely thick or eliminated by regional fusion of
color. Wings seldom fully expanded: when they do L3 and L4. Frequently extra crossvein between L3
expand, they are opaque and abnormal in shape. and L4 near end of wing. Wing narrowed. Head
Femurs kinked. Flies seem unable to move narrowed and flattened, so that long axis of eye is
normally and die on the food soon after eclosion. at oblique angle. May overlap wild type at high
RK3. temperatures and in late counts. Best at 19°C.
*Kg: Kugel RK2.
location: 3-48,2. *kno: knobbyhead
origin: Spontaneous. location: 1-63.9.
discoverer: B©nx, 1953. origin: Induced by triethylenemelamine (CB. 1246).
references: 1956, Rev. Suisse Zool. 63: 208-16. discoverer. Fahmy, 1951.
phenotype: Larva, pupa, and adult shorter and references: 1958, DIS 32: 70.
thicker than normal. Most striking in pup®. Homo- phenotype: Abnormal head; one or both eyes irreg-
xygote more estrero« than heterozygote. Homozy- ularly shaped, often drastically reduced in size.
gote viability 68 percent of wild type and fertility Occipital region frequently has hairy tufts, often
somewhat reduced. RK2. carried on protuberances. Males highly infertile;
Ki: Kicked viability about 10 percent wild type. RK2.
location: 3-47.6 (to the left ©f p). other information: One altele induced by CB. 2511.
origin: Spontaneous. knot: see Jen
discoverer: R. F. Grell, 571. Kr: KrUppel
references: 1958, DIS 32: 80. location: 2-108 (published value of 113 must be in-
pJwftotyfHK All bristles and hairs of beterozygote correct because the chromosome is only 108 units
shortened and twisted. Resembles am. Viability long)'
aKd fertility excellent; classification easy. Hooso- origin: Spontaneous,
has more-extretne bristle and limit effects. discoverer: Graber.
MUTATIONS
117
references: Gloor, 1950, Arch. Julius Klaus-Stift. phenotype: Eyes of L2/+ as small as or smaller than
Vererbungsforsch. Sozialanthropol. Rassenhyg. L/L. L2 homozygotes have tiny eyes and are
25: 38-44 (fig.). poorly viable or completely lethal, depending on
1954, Arch. Julius Klaus-Stift. Vererbungsforsch. background. Best used as heterozygote. Eyes
Sozialanthropol. Rassenhyg. 29: 277—87. further reduced in combination with M(3)w,
phenotype: Kr/+ adult sometimes has thoracic mal- M(3)h33i, and M(2)l2 (Dunn and Coyne, 1935, Biol.
formation; a leg or wing may be absent. Pene- Zentr. 55: 385—89). Classifiable in single dose in
trance low. Kr/Kr lethal before hatching. Primary triploids (Schultz, 1934, DIS 1: 55). Reduced
body segments of embryo abnormal, particularly number of cells enter into formation of eye disks
median segments. Ventral chain of ganglia discon- (Steinberg, 1944, Proc. Natl. Acad. Sci. U.S. 30:
nected. Tracheal system defective. Malpighian 5—13). RK1 as heterozygote.
tubules missing. Salivary glands normal. RK2.
KS: Male fertility complex in the short arm of the
Y chromosome
The male fertility complex of the short arm of the Y
chromosome, originally called K2 by Stern (1929,
Z. Induktive Abstammungs- Vererbungslehre 51:
253—353) and subsequently called KS by Brosseau
(1960, Genetics 45: 257—74), is subject to muta-
tions and deficiencies leading to male sterility,
which are symbolized ms(Y)S. By complementation
analysis, Brosseau divided KS into two comple-
menting units, ks-1 and ks-2. He believes that the
most probable order of factors on Ys from the tip L2: Lobe-2
toward the centromere is ks-2, ks-1, bb. Edith M. Wallace, unpublished.
Kugel: see Kg
kz: kurz
location: 1-0.9 (to the right of pn). origin: Spontaneous.
origin: Spontaneous. discoverer: Bridges, 24dlO.
discoverer: Stern, 26a23. references: Morgan, Bridges, and Sturtevant, 1925,
references: 1930, Z. Induktive Abstammungs- Bibliog. Genet. 2: 230.
Vererbungslehre 53: 279-86. phenotype: Eye reduction intermediate between L
1934, DIS 1: 35. and L2, but variability high. RK2.
phenotype: Bristles shorter and finer, like a slight
Minute. Postcutellars often absent. Hatches some-
what late. Viability fair; both sexes fertile. RK2. origin: Spontaneous.
cytology: Salivary chromosome location in region discoverer: Sturtevant, 23f.
2E1 through 2F6 (Demerec, Kaufmann, Fano, synonym: Lc.
Sutton, and Sansome, 1942, Carnegie Inst. Wash. phenotype: Heterozygote has fairly constant reduc-
Year Book 41: 191). tion in eye size, but not so great as to exclude its
use in combination with most eye colors. Expres-
L: Lobe sion more extreme at 25° than at 19°C. Hoaoxygot©
location: 2-72.0. has smaller eyes but lowered viability. Six® of
origin: Spontaneous. eyes of L4/+ reduced in combination with M(3)w,
discoverer: Bridges, 18i24. M(3)h33i, and M(2)!2 (Dunn and Coyne, 1935, Biol.
references: Morgan, Bridges, and Sturtevant, 1925, Zentr. 55: 385—89). Development similar to L3
Bibliog. Genet. 2: 230 (fig.). (Steinberg, 1944, Proc. Natl. Acad. Sci. U.S. 30:
phenotype: Heterozygous L eyes slightly smaller, 5—13). Reduced size of cephalic complex detect-
with nick in anterior edge, and lower half of eye able in 24-hr larva, but subsequent growth rate
reduced more than upper; overlaps wild type. similar to wild type (Medvedev, 1935, Z. Induktive
Homozygous L, eyes much smaller and less vari- Abstammungs- Vererbungslehre 70: 55—72, Tr.
able. Size of L/+ eyes reduced in combination with Inst. Genet. Akad. Nauk. SSSR 10: 119-51). RK2
M(3)w, M(3)h33), and M(2)l2 (Dunn and Coyne, as heterozygote.
1935, Biol. Zentr. 55: 385-89). Best used as a 5
recessive. RK2. origin: Spontaneous,
L.2 discoverer: Mohr, 3!k26.
origin: Spontaneous. references: Dunn, 1935, DIS 4: 14.
discoverer: Mohr, 20b2. pKenotyp©: Heterozygote ha* small nick in eyes,
references: 1924, Z. Induktive Abstammungs- overlap* wild type. Hotnoajygote has small ef®s
Vererbungslehre 32: 216. and tendency to antenna reduplication. May h«
Morgan, Bridges, and Sturtevant, 1925, Bibliog. used as a recessive but not as a dominant. Mor#
Genet. 2: 230. extreme than Lr but less so than L*. Development
118 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

similar to that of L? (Steinberg, 1944, Proc. Natl. phenotype: Eye reduction strong, with little varia-
Acad. Sci. U.S. 30: 5-13). RK3. tion in heterozygote. Homozygote more extreme;
L34 viability and fertility high. RK2.
origin: Spontaneous. cytology: Salivary chromosomes normal.
discoverer: Glass, 1934. Lr: Lobe-recessive
references: 1939, DIS 12: 47. origin: Spontaneous.
phenotype: Between L and L4. Dominance varies in discoverer: L. V. Morgan, 29h23.
different crosses. RK2. phenotype: Homozygote has small kidney-shaped
L52c eyes. Overlaps wild type at 19°; generally good at
origin: Spontaneous. 25°C. Heterozygote rarely shows seam or nick.
discoverer: Nakayama, 52c. RK2 as homozygote.
references: 1953, DIS 27: 59. Lrt>: Lobe-rough
phenotype: Like L. RK2. origin: Ultraviolet induced.
discoverer: Edmondson, 49k.
LB: Lobe of Becker references: Meyer, Edmondson, Byers, and Erickson,
origin: Spontaneous in In('2L)Cy +In(2R)Cy. 1950, DIS 24: 60.
discoverer: Becker. phenotype: Eye reduction similar to L4, but eye
references: 1957, Z. Induktive Abstammungs- surface slightly rough.
Vererbungslehre 88: 333—73 (fig.). Homozygote has extreme reduction of eyes; few or
phenotype: Not separable from Cy; homozygote no facets. L4/LTO has similar reduction in size of
therefore cannot be tested. Lower half of eyes eyes. Viability and fertility excellent. RK1 as
reduced or absent in heterozygote; more extreme at homozygote; RK2 as heterozygote.
25° than at 18°C. Sectors of ommatidia replaced by Lsi: Lobe-sinuate
chitin and bristles. Lower half of eyes apparently origin: Spontaneous.
produced from fewer than the normal 9 or 10 pre- discoverer: Morgan, 1932.
sumptive ommatidia-producing cells. Lower half of phenotype: Eyes of heterozygote flat, smooth, nearly
head also reduced at 25° but not 18°C. Tempera- full size, with sinuate margin; overlaps wild type
ture-sensitive period for ommatidia formation first only slightly. Eyes of homozygote smaller, with
and second ins tars; third instar as well for head flat or concave contour, smooth surface, and
reduction. RK2A. sinuate lower margin. RK3.
other information: Allelism to L inferred from pheno-
type and linkage to Cy alone. /( ): lethalf )
Lc: see L4 General term used to describe recessive mutations
*Ld: Lobe-duplicating that lead to death of most or all homozygous car-
origin: Spontaneous. riers. The symbol / is followed parenthetically by
discoverer: Kodani. the chromosome and then by the designation of the
references: Zimm, 1951, J. Exptl. Zool. 116: 289— particular mutant. Unfortunately, it is not practi-
319 (fig.). cable, except in special cases, to test allelism of
phenotype: Partially dominant. Characterized by sex-linked lethals, and it has not been common to
incomplete penetrance and variable expression: retain and test allelism of autosomal lethals. Con-
reduced single or bipartite eyes, kidney-shaped sequently, little information on allelism of lethals
eyes with bristles, large eyes with palps, or dupli- with similar genetic location is included.
cated antennae. Modifiers present on chromosomes 1-mah see l(l)m
I and 3. Penetrance affected by temperature during *l(l)h lethcl(l) J
development. RK3. location: 1-1.1.
origin: Spontaneous.
*Ld*: Lobe-diminished discoverer: Rawls, 12b.
origin: Spontaneous. references: 1913, Biol. Bull. 24: 115-24.
discoverer: Kadel and Jenkins, 55g. Morgan and Bridges, 1916, Carnegie Inst. Wash.
synonym: dq. Publ. No. 237: 31.
references: Kadel, 1956, DIS 30: 73-74. other Information: First recessive lethal found in
1957, DIS 31: 83. D. melanogaster.
phenotype: LdQ/+ normal. Eyes of Ld<3/Ldti irregu- KD3C3
larly deformed; occasionally divided into two or location: 1-1.6 (between w and rst).
root* lobes. Expression variable; some overlap of origin: Synthetic.
wild type, especially in old cultures. Aristae discoverer: Lefevre and Wilkins.
reduced and deformed. RK2. references: 1964, Genetics 50: 264.
phenotype: Male lethal. l(l)3C3/w is normal. RK2.
LK: Lobe of Krivsh&nko cytology: Associated with the deficiency for band
origin: Spontaneous. 3-C3 obtained as a single recornbinant carrying the
discover or: Krivshenko, 1957. left ©nd of T(l;4)w**J « T(t;4)3C2-3;2O;lQ2C and
references: 1958, DB 32: 81. the right end of InfDnt3 * In(l)3C3-4;20B.
MUTATIONS 119

cytology: Associated with T(l;3)l-184 -


location: 1- (0.4 to the left of y). T(1;3)18A;81.
origin: Spontaneous. *I(J)272-J3
discoverer Bridges, 14d9. location: 1- (rearrangement).
references: 1916, Genetics 1: 149. origin: X ray induced.
1(1)7: see discoverer: Demerec, 1940.
references: Sutton, 1943, Genetics 28: 210—217.
location: 1-0. phenotype: Lethal. 1(1)272-13/'sc is scute. RK2A.
origin: Spontaneous in dor1. cytology: Associated with In(l)l-272-13 =
discoverer: Bridges, 1928. In(l)lA6-Bl;llA7-8;llF2-12Al;18A4-Bl.
phenotype: Probably a specific modifier of
1(1)7'e dor1 dies earlier than dor1. RK3. origin: Spontaneous.
cytology: Included in 2RDXP element of T(l;2)Bld discoverer: Demerec, 1936, 1937.
T(1;2)1C3-4;6OB12-13 whereas dor1 is not. references: Slizynski, 1938, Genetics 23: 283—90.
cytology: Salivary analysis (Slizynski, 1938) shows
location: 1-21.3 (19.0 to 23.6). some normal, some deficient, and some inverted for
discoverer: Sobels. single-lettered subdivisions of the X chromosome.
references: Gloor, 1962, Rev. Suisse Zool. 69: other information: A series of 13 independently
409-63 (fig.). occurring and genetically located lethals.
phenotype: Larvae lethal in third instar; survive up *l(l)294-
to 10 days. Testes and lymph glands degenerate. origin: X ray induced.
Imaginal disks develop normally after transplanta- discoverer: Demerec, 36d.
tion. Protein metabolism disturbed; free amino references: Slizynski, 1938, Genetics 23: 283—90.
acids and peptides abnormally high. RK2. cytology: Three normal, one deficient for 10B, and
one deficient for IOC.
l(l)48j: see l(l)mys other information: A series of five independently
*I(J)52 induced and genetically located lethals.
location: 1- (to the right of B). *l(l)296-
discoverer: Sobels. origin: Spontaneous.
references: Gloor, 1962, Rev. Suisse Zool. 69: discoverer: Demerec, 1936.
409-63 (fig.). references: Slizynski, 1938, Genetics 23: 283-90.
phenotype: Larvae die in second instar. Growth cytology: Salivary chromosomes normal.
retarded. Histology of nervous system, testes, and other information: A series of six independently
imaginal disks abnormal. Number of nuclei in occurring and genetically located lethals.
salivary glands increased. Amino acids and pep- *l(l)302-
tides increased. Transplanted testes and imaginal origin: Neutron induced,
disks autonomously lethal. RK2. discoverer: Demerec, 36k.
references: Slizynski, 1938, Genetics 23: 283-9Q.
location: 1- (claimed to be about 0.6 unit to the left cytology: Three normal; one deficient for 3F.
of y, making it the leftmost known locus on the other information: A series of four independently
first chromosome). induced and genetically located lethals.
discoverer: Burdick, 55a. */(7)304-
references: 1956, DIS 30: 69. origin: X ray induced.
1957, DIS 31: 86. discoverer: Demerec, 37d.
phenotype: Presumed to be a lethal. Heterozygote references: Slizynski, 1938, Genetics 23: 283-90.
claimed to have viability about 1.5 times normal. cytology: Three normal; one deficient for 4C.
Not allelic to 1(1 )J1. RK2. other information: A series of four independently
other information: Map location of this mutation induced and genetically located lethals.
must be near y; however, the published data (1957, l(l)bt: see brl-
DIS 31: 86) permit the following alternative to a KDC
location to the left of y: l(l)55a is not completely location: 1-6 (between ec and bi).
lethal, and the 1.29 percent scored as crossovers origin: Spontaneous in sc t2 v si B chromosome.
to the left of y are actually surviving noncross- discoverer: Muller, 20j.
overs. references: 1928, Genetics 13: 279-357.
1(1)76: see dor13 phenotype: Dies as late embryo or, more commonly,
as first-instar larva (Brehme, 1937, Am. Naturalist
location: 1- (rearrangement), 71: 567). RK2A.
origin: X ray induced. cytology: Associated with the left breakpoint of
discoverer: Lindsley, Edington, and Von Halle, In(l)Cl «= In(l)4A5-Bl;17A6-Bl.
references: 1960, Genetics 45: 1649—70. *I(1)DM: lethol(l) of D. Mor/wofcj
phenotype: Almost completely lethal. The few sur- origin: X ray induced.
vivors have dark rough eyes. RK2A. discoverer: Moriwaki, 1932.
120 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: 1934, Japan J. Zool. 5: 585-602. I(1)EN7


1940, DIS 13: 50. location: 1- (rearrangement).
other information: Five independently induced and origin: X ray induced.
roughly located lethals. discoverer: Novitski.
references: 1963, DIS 37: 52.
I(1)ENT: lethal(l) from Eugene nonautonomous phenotype: Usually dies as early pupa; survives as
location: 1-46. patches of hemizygous tissue in gynandromorph.
origin: X ray induced. Fat bodies beaded instead of ribbonlike in third
discoverer: Novitski. larval instar. More free glutamine but less free
references: 1963, DIS 37: 51-52. tyrosine than normal. RK2A.
phenotype: Dies between first instar and prepupa; other information: Crossing over in X greatly re-
survives as patches of hemizygous tissue in gynan- duced.
dromorph. Salivary glands and gastric ceca small, I(1)EN8
and fat bodies usually absent in third-ins tar larva; location: 1- (close to the left of cv).
excess of free alanine or closely related sub- origin: X ray induced.
stance, and less free tyrosine than normal. RK2. discoverer: Novitski.
I(1)EN2 references: 1963, DIS 37: 52.
location: 1-0.3. phenotype: Usually dies in second larval instar; 25
origin: X ray induced. percent survive to third instar and a few to pupa-
discoverer Novitski. tion. Survives as patches of hemizygous tissue in
references: 1963, DIS 37: 52. gynandromorph. Fat bodies, Malpighian tubes, and
phenotype: Dies at end of third instar or in early salivary glands reduced. Rare adult survivors have
pupa; survives as patches of hemizygous tissue in soft exoskeleton and die within a few days. More
gynandromorph. More free glutamine but less free free glutamine but less free tyrosine than normal.
glutamic and as part ic acids than normal. RK2. RK2.

location: 1- (near car). I(1)EN9


origin: X ray induced. location: 1-10.
discoverer: Novitski. origin: X ray induced.
references: 1963, DIS 37: 52. discoverer: Novitski.
phenotype: Dies shortly after pupation; survives as references: 1963, DIS 37: 52.
patches of hemizygous tissue in gynandromorph. phenotype: Usually dies as third-instar larva; sur-
Larva usually has red-black pigmented areas in or vives as patches of hemizygous tissue in gynan-
on the cuticle. More free glutamine than normal. dromorph. Larva becomes transparent. Salivary
RK2. glands, Malpighian tubes, and fat bodies much re-
I(1)EH4 duced. Unknown fluorescent substance accumu-
location: 1-52. lates in larval cuticle. More free glutamine but
origin: X ray induced. less free tyrosine than normal. RK2(A).
discoverer Novitski. other information: Crossing over suppressed at tip
references: 1963, DIS 37: 52. of-X.
phenotype: Roughly 40 percent eclose but die imme- !(1)ENJO
diately; survives as patches of hemizygous tissue location: 1-59.
in gynandromorph. No morphological abnormalities origin: X ray induced.
observed in larva or pupa. More free glutamine discoverer: Novitski.
than normal. RK3. references: 1963, DIS 37: 52.
I(1)EN5 phenotype: About half die as pupae and half as few-
location: 1-47. day-old adults. Survives as patches of hemizygous
origin: X ray induced. tissue in gynandromorph. More free glutamine than
discoverer: Novitski. normal; free tyrosine nearly absent; low in free
references: 1963, DIS 37: 52. proline. RK3.
phenotype: Usually dies before third instar; sur- KDENJOa
vive® as patches of hemizygous tissue in gynan- location: 1-50.
droaaorpfa. More free glutamine than normal. RK2. origin: X ray induced.
1(1)EN6 discoverer: Novitski.
location: 1- 63 [between cmr and au(f)]. references: 1963, DIS 37: 52.
origin: X ray induced. phenotype: Dies as pupa. Third-instar larva shows
discoverer. Novitski. large excess of free glycine. RK2.
r«f©rtmc«s: 1963, DIS 37: 52. KDEHll
pbenotyp*: Dies at any stage; survives as patches location: 1-43.
of bemizygous tissue in gynandromorph. Larval fat origin: X ray induced.
bodies and Malpighian tubes reduced. More free discoverer: Novitski.
glutamine than normal. RK2. references: 1963, DIS 37: 52.
MUTATIONS 121

phenotype: Dies from second instar through pupa. movements but is unable to break through vitelline
Survives as patches of hemizygous tissue in membrane. Muscular activity persists several
gynandromorph. Melanotic spots on some larvae hours, but hatching does not occur and cell degen-
and inside pupae. Culture with dying larvae has eration begins at about 25 hr. Differentiation
distinct urinous odor. More free phenylalanine and abnormal in several ways: pharyngeal apparatus
less free tyrosine than normal. RK2. reduced and distorted; brain forms irregular mass;
I(1)ENU constriction forms behind head; segmentation dis-
location: 1-3. torted; and body wall usually incomplete dorsally.
origin: X ray induced. RK2.
discoverer: Novitski. *I(1)GSB: lethal(l) of Gershenson, Shapiro, and
references: 1963, DIS 37: 52. Borissenko
phenotype: Usually dies in third larval instar or origin: X ray induced in In(l)scs.
pupa. Rare survivors reach eclosion. Survives as discoverer: Gershenson, Shapiro, and Borissenko,
patches of hemizygous tissue in gynandromorph. 1931.
Flies that reach eclosion have soft exoskeleton references: Gershenson, 1934, DIS 1: 54.
with little pigmentation; appear almost translucent. other information: A series of 51 independently
Low in free tyrosine. RK2. induced and genetically located mutants.
KDEN13 *l(l)l: lethal(i) of Ives
location: 1-13.4. origin: Recovered from heat-treated lines.
origin: X ray induced. discoverer: Ives.
discoverer: Novitski. references: Plough and Ives, 1934, DIS 1: 32.
references: 1963, DIS 37: 52. 1935, Genetics 20: 42-69.
phenotype: Some survivors. No gross larval or pupal other information: A series of 13 independently
abnormalities. Low in free tyrosine. RK3. occurring mutants.
KVENU i(l)Jh lethal(i) of Jacobs-Duller
location: 1- (rearrangement). location: 1-0.0 (to the left of y).
origin: X ray induced. origin: X ray induced simultaneously with scJ*.
discoverer: Novitski. discoverer: Jacobs-Muller.
references: 1963, DIS 37: 52-53. references: Muller, 1932, Proc. Intern. Congr.
phenotype: Dies mostly in second, but also in third- Genet., 6th. Vol. 1: 225.
instar larva; survives as patches of hemizygous Muller, 1935, Genetica 17: 237-52.
tissue in gynandromorph. Accumulates propanol- phenotype: Lethal. Not cell lethal (Ephrussi, 1934,
ammonia-insoluble fluorescent substance in larval Proc. Natl. Acad. Sci. U.S. 20: 420-22). One
cuticle. Less free tyrosine and proline than recorded surviving male had rough eyes and was
normal. RK2A. sterile. RK2A.
other information: Crossing over in X greatly re- cytology: Probably in 1A6. Associated with
duced. In(l)scJ1 =In(l)lA4-5;lB4-5 (Muller, Prokofyeva,
KDEN15 and Raffel, 1935, Nature 135: 253-55).
location: 1- (near car). I(1)J12S9
origin: X ray induced. origin: X ray induced in y-bearing chromosome.
discoverer Novitski. discoverer Lindsley, Edington, and Von Halle,
references: 1963, DIS 37: 53. references: Frye, 1959, Genetics 44: 511.
phenotype: Dies as third-instar larva or early pupa; *l(l)jl: lethal(l) /aw/ess
survives as patches of hemizygous tissue in location: 1-14.
gynandromorph. Less free tyrosine and proline origin: Ultraviolet induced,
than normal. RK2. discoverer: McQuate, 1951.
t(l)ENU references: Oster, 1952, Heredity 6: 403-7.
location: 1-24. phenotype: Dies during first larval instar. Mouth
origin: X ray induced. parts poorly formed and sometimes absent. RK2.
discoverer Novitski. cytology: Salivary chromosomes normal (Valencia
references: 1963, DIS 37: 53. and McQuate, 1951, Genetics 36: 580).
phenotype: Dies between first-instar larva and pupa. *I(1)K: lethalO) of King
Less free tyrosine and proline than normal. RK2. origin: Recovered among progeny of males fed P32.
*!(l){fn: lethal(l) formalin food discoverer: R. C. King, 1948 and 1949.
location: 1- (not located). references: 1950, DIS 24: 58.
origin: Induced by formaldehyde. other informotion: Four independently induced and
discoverer: Auerbach. genetically located lethals.
synonyms: Ltlll. *IO)LB: lethal(l) of Luers and Belitz
references: Ede, 1956, Arch. Entwicklungsmech. discoverer: Luers and Belitz, 1951-1956.
Organ. 148: 416-36 (fig.). references: Belitz, 1954, Z. Induktive Abstarnrnungs-
phenotype: Develop® to late embryonic stage, at 22 Vererbungslehre 86: 173-84.
hr (normal hatching time) shows vigorous muscular 1956, DIS 30: 104.
122 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

other information: A series of nearly 500 mutations phenotype: 20-hr embryos (25°C) show mid-dorsal
recovered from untreated males or from males herniation of brain or mid-gut, or both, abnormal
treated with Miracil [l-(2-diethylaminoethylamino)- somatic, visceral, and pharyngeal muscles, and
4-methylthioxanthineJ, Mirasan, triethylenemela- incomplete morphogenesis of yolk-filled mid-gut.
mine, aminopterin, hesperidine, or 2,5-bisethylene- Development of embryo normal up to 13 hr.
imine-l,4-benzoquinone. All lethals were located Between 13 and 14.5 hr, first muscular contrac-
genetically. tions occur, while basement membrane is incom-
*l(l)m: lethal(l) malignant plete. This results in dorsal rupture of hypoderm,
location: 1- (not located). retraction of myogenic elements of somatic and
origin: Induced by mustard gas. pharyngeal muscles into spheroidal masses. Con-
synonym: l-m&l. tinuation of myogenesis produces spheroidal
references: El Shatoury, 1955, Arch. Entwicklungs- muscles with a cortex of disoriented fibrillae sur-
mech. Organ. 147: 496-522 (fig.). rounded by a medulla of nucleated sarcoplasm.
El Shatoury and Waddington, 1957, J. Embryol. RK2.
Exptl. Morphol. 5: 143-52 (fig.). cytology: Salivary chromosomes normal.
phenotype: Cells originating from lymph glands in
late third instar first spread to, and cause, *l(l)nd: lethal(l) no differentiation
destruction of imaginal buds and later may move location: 1- (not located).
along ventral nerve cord to attack posterior fat origin: Induced by mustard gas.
bodies and testes. The tumor cells eventually references: El Shatoury, 1955, Arch. Entwicklungs-
become melanotic after destruction of various mech. Organ. 147: 523-38 (fig.).
healthy tissues. Death occurs in late larval or phenotype: Some or all imaginal buds fail to differ-
early pupal stages. Claimed to be the only true entiate during larval third instar, apparently as a
malignancy in Drosophila melanogaster, RK2. result of abnormal proliferation of imaginal disk
mesoderm. Death in pupal or prepupal stage. RK2.
*I(1)MA: lethal(l) of Mailer and Altenburg
origin: Spontaneous. *l(l)ne: lethal(l) nonevaginated
discoverer: Muller and Altenburg. location: 1-0.1.
references: 1919, Proc. Soc. Exptl. Biol. Med. 17: origin: Induced by urethane.
10-14. discoverer: Vogt, 1949.
other information: A series of about 50 mutants of references: 1951, DIS 25: 76.
which only a few were located. Florschutz-de Waard and Faber, 1952, DIS 26: 99.
1(1 )ml: lethal(l) melanoma!ike Faber, Sobels, Florschiitz-de Waard, and
location: 1-10. Oppenoorth, 1954, Z. Induktive Abstammungs-
origin: Ultraviolet induced. Vererbungslehre 86: 293—321 (fig.).
discoverer: McQuate, 1951. phenotype: Lacks imaginal thoracic hypoderm.
references: Oster, 1952, Heredity 6: 403-7. Cephalic complex and thoracic imaginal disks fail
Oster and Sobels, 1956, Am. Naturalist 90: 55-60. to evaginate. The unaffected abdominal hypoderm
phenotype: Larvae die in third instar. At death, develops but ends anteriorly in a free edge that
they have internal melanotic masses (usually one folds back on itself and forms a darkly pigmented
or two, sometimes as many as ten). RK2. ring around the pupa. Genital disk capable of
cytology: Salivary chromosomes normal (Valencia normal evagination but vasa deferentia do not
and McQuate, 1951, Genetics 36: 580). connect to testes, which do not spiralize. Death
*l0)mt: lethal(l) midget occurs 3—5H days after prepuparium formation.
location: 1-2.5. Pupae darker than normal, with sticky, irregular
origin: Ultraviolet induced. surface and distinctly meandering tracheal trunks.
discoverer: McQuate, 1951. RK2.
references: Oster, 1952, Heredity 6: 403-7. *l(l)nib; lethal(l) no imaginal buds
phenotype: Dies as undersized third instar larva. location: 1- (not located).
RK2. references: El Shatoury and Waddington, 1957, J.
cytology. Salivary chromosomes normal (Valencia Embryol. Exptl. Morphol. 5: 143—52 (fig.).
and McQuate, 1951, Genetics 36: 580). phenotype: Dies in third larval instar. Imaginal
l(l)mys: lethal(l) myospheroid buds small or absent. Excessive proliferation of
location: 1-21.7. stomach epithelium leads to occlusion of gut. Pro-
origin: Induced by P 3 2 . liferations degenerate into melanotic masses.
discoverer: Poulson, 48j. RK2.
synonyms: 1(1)48). 1(1 )Q: lethal(l) Quinacrine mustard induced
references: Rizki, 1956, J. Exptl. Zool. 131: origin: Induced by 2-methoxy-6-[3-(ethyl-2-chloro-
203-22 (fig.). ethyl)aminopropylaminojacridine (ICR 100).
Wright, 1958,, Proc. Intern. Congr. Genet., 10th. discoverer: Carlson.
Vol. 2: 323. references: Carlson, Sederoff, and Cogan, 1967,
1960, J. Exptl. Zool. 143: 77-99 (fig.). Genetics 55: 295-313.
MUTATIONS 123

other information: A series of 64 independently Number Location


induced and genetically located lethals. Their * 67 20.3
numbers and locations are tabulated below. * 68 40.4
Number Location * 69 54.1
* 1 13.1 * 70 51.9
* 2 40.9 * 71b 54.4
* 3 26.5 * 72 10.6
* 4 52.0 * 73 53.9
* 5 0.0 * 74 18.2
* 6 30.2 75 29.3
* 7 28.8 * 76 49.1
* 8 65.5 * 77 0.0
* 9 1.4 * 78 0.0
* 10 28.6 * 79 48.0
* 11 16.0 * 80 54.2
* 12 42.0 * 81 6.8
* 13 49.7 * 82 18.7
* 14 64.2 * 83 12.1
* 15 19.6 * 85 0.0
* 16 52.6 86 56.7
* 17 23.6 87 0.0
* 18 56.7 * 89 51.5
* 19 62.5 *201 13.4
* 20 0.0 202 40.1
21 20.5 203 62.5
22 39.1 *2O4 22.6
* 23 57.8 *205 21.1
* 24 58.3 206 13.0
* 25 23.0 208 17.0
* 26 33.0 *209 0.0
* 27 9.3 •210 64.5
* 28 44.6 •211 33.0
* 30 22.7 212 0
* 31 56.7 *214 56.7
* 33 29.9 *215 10.4
* 34 56.7 216 8.6
* 36 0.0 217 0.0
* 39 1.7 218 2.8
* 40 1.5 *219 1.3
* 41 2.0 220 0.0
* 42 64.4 221 0.0
* 43 37.6 222 57.9
* 44 31.7 223 29.2
* 45 38.3 224 47.8
* 48 21.3 225 36.0
* 49 12.7 226 47.9
* 50 62.5 227 27.4
* 51 65.2 *228 42.2
52 36.9 *231 13.5
* 53 32.0 *232 16.5
* 54 33.0 233 15.2
* 55 60.6 234 38.9
* 56 11.2 235 28.1
* 57 16.8 236 21.0
58 38.2 237 21.7
* 59 44.5 *238 54.1
61 6.5 *240 33.0
* 62 62.5 • 244 24.2
* 63® 53.0 248 12.2
64 41.4 *l(l)R: tetbalO) of Rohrborn
65 20.5 discoverer: Rohrborn, 1955, 1956.
* 66 33.0 reference*: 1959, Z. Vererbungslehrts 90: 116-31.
124 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

other information: A series of 71 lethals including 3 discoverer: McQuate, 1951.


spontaneous ones, 55 from males treated with 1:4- references: Oster, 1952, Heredity 6: 403—7.
dimethanesulfonoxybutane (CB. 2041), and 13 from phenotype: Dies during third larval instar. Main
males treated with l:4-dimethanesulfonoxy-l:4- tracheal tubes greatly enlarged, sometimes lack
dimethylbutane (CB. 2348). All were located functional posterior spiracles. RK2.
genetically. cytology: Salivary chromosomes normal (Valencia
*l(l)rr: lethal(l) ring gland rudimentary and McQuate, 1951, Genetics 36: 580).
location: 1-0.3. *l(l)th lethal(l) tracheae lacking
origin: Ultraviolet induced. location: 1-59.
discoverer: McQuate, 1951. origin: Ultraviolet induced.
references: Oster, 1952, Heredity 6: 403-7. discoverer: McQuate, 1951.
phenotype: Dies during third larval instar. Larvae references: Oster, 1952, Heredity 6: 403—7.
live 15—30 days but do not become giant. Ring phenotype: Dies during first larval instar. Main
gland abnormally small, probably causing failure tracheal tubes absent, although small side
to undergo third molt. RK2. branches present. RK2.
cytology: Salivary chromosomes normal (Valencia cytology: Salivary chromosomes normal (Valencia
and McQuate, 1951, Genetics 36: 580). and McQuate, 1951, Genetics 36: 580).
*I(1)S: leihal(l) of Stark
origin: Spontaneous. *l(l)tr: lethal(l) tracheae ramified
discoverer: Stark, 1913, 1914. location: 1-56.
references: 1915, J. Exptl. Zool. 19: 531-58. origin: Ultraviolet induced.
Morgan and Bridges, 1916, Carnegie Inst. Wash. discoverer: McQuate, 1951.
Publ. No. 237: 64, 79. references: Oster, 1952, Heredity 6: 403-7.
other information: Four independently occurring phenotype: Dies during first larval instar. Main
lethals. tracheal tubes thick and have numerous side
*I(1)S9 branches. RK2.
location: 1- (to the right of car). cytology: Salivary chromosomes normal (Valencia
origin: Spontaneous. and McQuate, 1951, Genetics 36: 580).
discoverer: Auerbach.
references: Ede, 1956, Arch. Entwicklungsmech. *l(l)trs: lethal(l) tracheae stretched
Organ. 149: 256-66 (fig.). location: 1-8.0.
phenotype: Almost all embryos deformed at anterior origin: Ultraviolet induced.
end, where there is usually some undigested yolk. discoverer: McQuate, 1951.
Death occurs in embryonic, larval, and pupal synonym: l(l)ts (preoccupied).
stages. Primary abnormality is distribution of references: Oster, 1952, Heredity 6: 403—7.
cleavage nuclei, which causes blastoderm to be phenotype: Dies during first larval instar. Larvae
fragile at its anterior end. RK2. very large for this stage and all tracheal tubes very
l(l)sc: lethal(l) at scute thin, suggesting that they grow more slowly than
location: 1-0 (immediately to the right of sc). larvae and thus become stretched. RK2.
origin: Synthetic. cytology: Salivary gland chromosomes normal
discoverer: Muller. (Valencia and McQuate, 1951, Genetics 36: 580).
references: 1935, Genetica 17: 237—52. *l(l)ts: lethal(l) temperature sensitive
other information: Inferred from the inviability of location: 1-8.
In(l)nc*L&c9R = In(l)lB3-4;19F-20Cl^lB2- discoverer: Falbo and Re'.
3;18B8-9R [left break of In(l)sc9 in doubt] except references: 1945, DIS 19: 45, 57.
in the presence of Dpfi;2)sc* *\ phenotype: Inviable in cultures grown at 23°C but
shows more than 50 percent survival in cultures
*l(1)sd: lethal(l) schaiben defekt grown at 26.5°. RK3.
location: 1-17.9. *l(l)TS-45: lethal(l) no. 45 of T. Shiomi
origin: Induced by triethylenemelamine (CB. 1246). location: 1-5.8.
discoverer: M. J. Fahmy. origin: X ray induced.
references: Schnitter, 1961, Rev. Suisse Zool. 68: discoverer: Shiomi, 52f.
345-418 (fig.). references: 1954, DIS 28: 78.
phenotype: Dies during transition from larva to Imaizumi and Shiomi, 1955, Arch. Biol. (Liege) 66:
prepupa. Some larvae form puparia but do not dif- 483-87.
ferentiate further. Pattern of damage complex, most phenotype: Dies before hatching. No visible morpho-
severe defects being found in certain imaginal logical abnormality. Heterozygote of l(l)TS-45/
disks. Several larval organs abnormal, especially Base has average of 612 eye facets compared to
the salivary glands. RK2. only 402 in +/Bssc. Accumulation of urea or car-
*l(l)te: lethal(l) tracheae enlarged barnides in larvae of heterozygote; these compounds
location; 1-0.3.
presumably tend to normalize the Bar phenotype.
origin: Ultraviolet induced. RK2.
MUTATIONS 125

*l(l)TS-56 references: 1960, Genetics 45: 1649—70.


location: 1-1.5. phenotype: Viability of X/Y males 63 percent
origin: X ray induced. normal; further reduced in presence of E(var)7 and
discoverer: Shiomi, 52f. M(2)S2l°. X/Y males fertile. X/0 males lethal.
references: 1954, DIS 28: 78. Homozygous females viable, with fewer and smaller
phenotype: Lethal in late embryonic stage. Develop- bristles. RK2A as X/O male.
ment of tracheae, other chitinized parts, and body cytology: Associated with In(l)l-v59 = In(l)3-
segments abnormal. RK2. 4; 19-20.
I(1)v75
I(l)v3: lethal(l) variegated location: 1- (rearrangement).
location: 1- (rearrangement). origin: X ray induced.
origin: X ray induced. discoverer: Lindsley, Edington, and Von Halle.
discoverer: Lindsley, Edington, and Von Halle. references: 1960, Genetics 45: 1649—70.
references: 1960, Genetics 45: 1649-70. phenotype: Viability of X/Y males 26 percent
phenotype: Viability of X/Y males 74 percent normal; further reduced in presence of E(var)7 but
normal; recovery of X/Y males reduced by M(2)S210 not M(2)S210. X/Y males sterile, owing to failure
but not E(var)7. X/Y males sterile, owing to failure of sperm head to elongate. Viability of X/0 males
of sperm head to elongate. X/0 males lethal. less than 1 percent normal. RK2A as X/0 male.
RK2A as X/0 males. cytology: Associated with T(l;2)l-v75 - T(l;2)19-
cytology: Associated with T(l;3)l-v3 = T(1;3)4A;81, 20;41.
\(})vU9
location: 1- (rearrangement). location: 1- (rearrangement).
origin: X ray induced. origin: X ray induced.
discoverer: Lindsley, Edington, and Von Halle. discoverer Lindsley, Edington, and Von Halle.
references: 1960, Genetics 45: 1649-70. references: 1960, Genetics 45: 1649—70.
phenotype: Viability of X/Y males 78 percent phenotype: Semilethal. Viability of X/Y males 91
normal; further reduced in presence of M(2)S2*° but percent normal, of X/0 males 26 percent normal,
not E(var)7. X/Y males fertile. Viability of X/0 X/Y males sterile, owing to failure of sperm head
males 4 percent normal. Homozygous females sur- to elongate. RK3A.
vive and have blistered wings and duplicate an- cytology: Associated with T(l;2)l-vl29 =
terior scute liar and postalar bristles; addition of T(1;2)18B;41.
y+Y eliminates wing effect. RK3A.
cytology: Associated with T(l;4)l-vll = M)vU2
location: 1- (rearrangement).
I(l)v25 origin: X ray induced.
location: 1- (rearrangement). discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. references: 1960, Genetics 45: 1649-70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: Viability of X/Y males 83 percent
references: 1960, Genetics 45: 1649—70. normal. X/Y males fertile. Viability otX/0 males
phenotype: X/Y males of normal viability but less than 1 percent normal. Homozygous females
sterile, owing to failure of sperm head to elongate. viable. RK2A as X/0 male.
X/0 males lethal. RK2A as X/0 males. cytology: Associated with In(l)l-vl32 = ln(l)3-
cytology: Associated with T(l;2)l-v25 = T(l;2)19- 4,19-20.
20;40-41. I(l)vl35
location: 1- (rearrangement).
*l(J)v47 origin: X ray induced,
location: 1- (between cv and v). discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. references: 1960, Genetics 45: 1649—70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: Originally recovered as Y-suppressed
references: 1960, Genetics 45: 1649—70. lethal, but in subsequent tests both X/Y and X/0
phenotype: X/Y males have gg-like phenotype but males appeared to be lethal. Later tests by
with peripheral darkening of eye color; viability 41 Thompson show viability of X/Y males to be 40
percent normal; fertile. X/0 males lethal. RK2A. percent normal and X/0 males less than 1 percent
as X/0 male. normal. RK3A.
cytology: Associated with an insertion of an unspec- cytology: Associated with T(l;2)l-vl35 « T(l;2)18-
ified section of heterochromatin into 8F-9B. Link- 19;41. Induced simultaneously with T(2;3)135 =»
age tests suggest second chromosome origin of T(2;3)37;85A, from which it has since separated.
inserted material. T(l;2)l-v47 = T(1;2)8F-9B.
I(l)v59 I(l)vl39
location: 1- (rearrangement). location: 1-2 (between w and spj).
origin: X ray induced. origin: X ray induced.
discoverer. Lindsley, Edington, and Von Halle. discoverer: Lindsley, Edington, and Von Halle.
126 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

references: 1960, Genetics 45: 1649-70. discoverer: Lindsley, Edington, and Von Halle.
phenotype: X/0 and X/Y males lethal. X/Y/Y references: 1960, Genetics 45: 1649—70.
males viable and fertile; show strong variegation phenotype: Viability of X/Y males normal, but
for w and rst. RK3A. reduced in combination with M(2)S210 but not
cytology: Associated with In(lLR)l-vl39 = E(var)7. X/Y males sterile, owing to failure of
In(lLR)3C6-7. sperm head to elongate. X/0 males lethal. RK2A
other information: Single recombinant carrying distal as X/0 male.
part of .X-centromere-bearing half of T(l;4)wmS = cytology: Associated with T(l;2)l-v219 =
T(l;4)3C3-4;101Fl-2 and proximal part of In(lLR)l- T(l;2)10A;40.
vl39 is variegated for w but not for rst and is *l(l)v223
viable. location: 1- (rearrangement).
*t(l)vl46 origin: X ray induced.
location: 1- (rearrangement). discoverer. Lind^sley, Edington, and Von Halle.
origin: X ray induced. references: I960, Genetics 45: 1649—70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: Viability of X/Y males 41 percent
references: 1960, Genetics 45: 1649-70. normal. X/Y males sterile, owing to variegation for
phenotype: Viability oiX/Y males 41 percent absence of external genitalia, especially in com-
normal; further reduced by M(2)S21 ° and E(var)7. bination with E(var)7. X/0 males lethal. RK2A as
X/Y males fertile. X/0 males lethal. X/0 male.
I(l)vl46/I(l)vl46/Y more viable than cytology: Associated with T(l;2)l-v223 =
I(l)vl46/l(l)vl46 females. Frequently have fewer T(l;2)UF;41;50E.
dorsocentrals. RK2A as X/0 male. I(1)v227
cytology: Associated with In(l)l-vl46 = In(l)5- location: 1- (rearrangement).
6;19-20. origin: X ray induced.
l(l)vl50 discoverer: Lindsley, Edington, and Von Halle.
location: 1- (rearrangement). references: 1960, Genetics 45: 1649-70.
origin: X ray induced. phenotype: Viability of X/Y males 48 percent nor-
discoverer: Lindsley, Edington, and Von Halle. mal; further reduced in combination with M(2)S21 °
references: 1960, Genetics 45: 1649-70. and E(var)7. X/Y males fertile. X/0 males lethal.
phenotype: Viability of X/Y males 15 percent RK2A as X/0 male.
normal. X/Y males sterile, owing to failure of cytology: Associated with In(l)l-v227 = In(l)l-
sperm head to elongate. X/0 males lethal. RK2A 2,19-20.
as X/0 male. I(l)v231
cytology: Associated with T(l;2)l-vl50 = T(l;2)16- location: 1- (rearrangement).
17;40. origin: X ray induced.
discoverer: Lindsley, Edington, and Von Halle.
IO)v163 references: 1960, Genetics 45: 1649—70.
location: 1* (rearrangement). phenotype: X/Y males viable and fertile. Viability
origin: X ray induced. of X/0 males less than 1 percent normal; the few
discoverer: Lindsley, Edington, and Von Halle. survivors have reduced rough eyes. Homozygous
references: 1960, Genetics 45: 1649—70. females normal. RK2A in X/0 male.
phenotype: Viability of X/Y males 17 percent cytology: Associated with In(l)l-v231 =In(l)lC-
normal and of X/0 males less than 1 percent D-,19-20.
normal. X/Y males sterile, owing to failure of *IO)v252
sperm head to elongate. RK2A in X/0 male. location: 1- (rearrangement).
cytology: Associated with T(l;3)l-vl63 » T(1;3)17A- origin: X ray Induced.
B;8O-81. discoverer: Lindsley, Edington, and Von Halle.
*}(l)v216 references: 1960, Genetics 45: 1649-70.
location: 1- (rearrangement). phenotype: Viability of X/Y males normal, of X/0
origin: X ray induced. males 2 percent normal. X/Y males sterile, owing
discoverer: Lindsley, Edington, and Von, Halle. to failure of sperm head to elongate. RK2A as X/0
references: 1960, Genetics 45: 1649-70. male.
phenotype: Viability of X/Y males 15 percent cytology: Associated with T(l;3)l-v252; determined
normal; reduced further in presence of M(2)S2i ° but genetically; cytology not done.
not B(var)7. X/Y males sterile, owing to failure of *l(l)v306
sperm head to elongate. X/0 males lethal. RK2A location: 1-0.
as X/0 male. origin: X ray induced.
cytology: Associated with T(l;2;3)l-v216; deter- discoverer: Lindsley, Edington, and Von Halle.
mined genetically; cytology not done. references: 1960, Genetics 45: 1649—70.
phenotype: Viability of X/Y males 78 percent
location: 1- (rearrangement), normal. X/Y males fertile. X/0 males lethal.
origin: X ray induced. Homozygous females viable. RK2A in X/0 males.
MUTATIONS 127

cytology: Salivary chromosomes show insertion of I(l)v459


material of unknown origin into 1B-E. location: 1- (rearrangement).
t(1)v361 origin: X ray induced.
location: 1- (rearrangement). discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. references: I960, Genetics 45: 1649—70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: X/Y males have rough eyes and deformed
references: 1960, Genetics 45: 1649—70. wings and wing veins; 78 percent normal viability;
phenotype: X/Y males show normal viability but are fertile. X/0 males lethal. RK2A as X/0 male.
sterile, owing to failure of sperm head to elongate. cytology: Associated with T(l;2;3)l-v459 =
X/0 males lethal. RK2A as X/0 male. T(l;2;3)3D-F;XR;50;80-81.
cytology: Associated with T(l;3)l-v361 = T(l;3)19- l(J)v463
20;80-8t. location: 1- (rearrangement).
I(l)v451 origin: X ray induced.
location: 1-56.7 (inseparable from f). discoverer: Lindsley, Edington, and Von Halle.
origin: X ray induced. references: 1960, Genetics 45: 1649—70.
discoverer: Lindsley, Edington, and Von Halle. phenotype: Viability of X/Y males 50 percent
references: 1960, Genetics 45: 1649-70. normal; further reduced in combination with E(var)7
phenotype: Viability of X/Y males 63 percent but not M(2)S2lO. x/Y males sterile, owing to
normal, of X/0 males 4 percent normal. X/Y males failure of sperm head to elongate. Viability of X/0
sterile; spermatogenesis appears normal but sperm males 18 percent normal. RK3A.
not motile. RK3. cytology: Associated with T(l;3)l-v463 = T(l;3)19-
cytology: Salivary chromosomes appear normal. 20-.81-82.
other information: No translocation detectable *l(l)w
genetically. Map distance between v and i reduced location: 1-66.
from standard 23.7 to 12. discoverer: Schubel, 1934.
references: 1934, Am. Naturalist 68: 278—82.
I(l)v453 phenotype: Males survive; homozygous females die.
location: 1- (rearrangement). RK3.
origin: X ray induced. other information: Probably a lethal allele of bb.
discoverer: Lindsley, Edington, and Von Halle.
references: 1960, Genetics 45: 1649—70. *I(1)X2: lethol(l) X ray induced
phenotype: Viability of X/Y males normal; reduced location: 1- (near forked).
in combination with M(2)S21 ° but not E(var)7. X/Y origin: X ray induced.
males sterile, owing to failure of sperm head to discoverer: Auerbach.
elongate. X/0 males lethal. RK2A as X/0 male. references: Ede, 1956, Arch. Entwicklungsmech.
cytology: Associated with T(l;3)l-v453 = Organ. 148: 437-51 (fig.).
T(1;3)12D; 80-81. phenotype: Embryos die in advanced stage of devel-
I(l)v454 opment. They live beyond normal hatching time,
location: 1- (rearrangement). move actively, but do not hatch. Embryo distorted;
origin: X ray induced. head material not involuted and pharyngeal material
discoverer: Lindsley, Edington, and Von Halle. external; body wall has disarranged segmentation
references: 1960, Genetics 45: 1649-70. in medial region. Mutant disrupts mechanism con-
phenotype: Viability of X/Y males 50 percent nor- trolling mitosis in early stages of gastrulation,
mal; further reduced in combination with M(2)S210 occasionally as early as blastoderm formation.
and E(var)7. X/Y males sterile, owing to failure of RK2.
sperm head to elongate. X/0 males lethal. RK2A cytology: Salivary chromosomes normal.
in X/0 males.
cytology: Associated with T(l;2;3;4)l-v454 = *IO)X10
T(1;2;3)12B;22-23;81 + T(2;4)44F;t01F. location: 1-0.0 (near <sc).
origin: X ray induced.
f(1)v455 discoverer: Auerbach.
location: 1-(rearrangement). references; Ede, 1956, Arch. Entwicklungsmech.
origin: X ray induced. Organ. 149: 247-58 (fig.).
discoverer: Lindsley, Edington, and Von Halle. phenotype: Variation in expression of factors dis-
references: I960, Genetics 45: 1649-70. continuous. There are three types of lethal em-
phenotype: Viability in X/Y males low; further bryos; some may survive into larval stage. Type 1
reduced in presence of both M(2)S210 and E(var)7. stops development after formation of a cap of undif-
X/Y males sterile, owing to failure of sperm head ferentiated cells. Type 2 has limited differentia-
to elongate. X/0 males invisible. Eye color varie- tion, often the nervous tissue exclusively, but no
gated in I(l)v455/w females. RK2A in X/0 males. organ formation. Type 3 survives beyond normal
cytology: Associated with T(l;3)l-v455 « hatching time, has no .gross abnormalities, but does
T(1;3)3C;81. not hatch. RK2.
128 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

*l(J)X20
location: 1- (near sc). origin: X ray induced.
origin: X ray induced. discoverer: Abrahamson, 64f5.
discoverer: Auerbach.
references: Ede, 1956, Arch. Entwicklungsraech. origin: X ray induced.
Organ. 149: 101-14 (fig.)- discoverer: Judd, 63g9.
phenotype: Four types of defective embryos pro-
duced. Types 1 and 2 reach late stage of develop- origin: X ray induced,
ment and are alive at time larvae normally hatch. discoverer: Judd, 62gl7.
Type 1 has a complete nervous system, but incom- cytology: Salivary chromosomes appear normal
plete hypoderm. Type 2 has hypoderm but a defi- (Judd).
cient nervous system. Types 3 and 4 stop devel-
oping at early stages. Type 3 has no development origin: X ray induced.
beyond gastrulation, and type 4 forms no blasto- discoverer: Judd, 63gl9.
derm. RK2.
*I(1)X27
location: 1-63.4. origin: X ray induced.
origin: X ray induced. discoverer: Judd, 62g26.
discoverer: Auerbach.
references: Ede, 1956, Arch. Entwicklungsmech. origin: X ray induced.
Organ. 149: 88-100 (fig.). discoverer: Judd, 62g31.
phenotype: Embryos alive and in a late stage of de- cytology: Associated with In(l)Uzwl&31 =In(l)3A;6
velopment at normal hatching time but do not hatch. (Judd).
Degeneration begins at about 25 hr. Germ band
irregular at beginning of gastrulation, apparently origin: X ray induced.
the result of defective ventral furrow formation. discoverer: Judd, 62k5.
Consequently, hind-gut is open dorsally, nervous cytology: Salivary chromosomes normal (Judd).
system irregularly developed, and ventral nerve
cord interrupted in region of mid-gut. Other abnor-
malities from different causes are: (1) gut remains origin: X ray induced.
sac like; (2) ectoderm remains unsegmented; and (3) discoverer: Judd, 62k6.
musculature of body wall is underdeveloped. RK2. cytology: Salivary chromosomes normal (Judd).

l(1)zwl°h lethal(l) reste to white origin: X ray induced in z-bearing X chromosome.


location: 1-1.1. discoverer: Judd, 63k26.
origin: X ray induced. I(l)zw2a3
discoverer: Abrahams on, 62al. location: 1-1.2.
cytology: Salivary chromosomes appear normal; origin: X ray induced.
placed in region 3A5-7, on the basis of its inclu- discoverer: Abrahamson, 62a3.
sion in Df(l)64c4 = Dt(l)3A4~6;3C3-5 but not in phenotype: Lethal homozygous and when heterozy-
= Df(l)3A6-8;3Cl-3 (Judd), gous with all other alleles of I(l)zw2 except
\(l)zw2&6. RK2.
origin: X ray induced. cytology: Salivary chromosomes appear normal.
discoverer: Abrahams on, 62a2. Placed in region 3A7-B1, on the basis of its inclu-
sion in Df(l)64j4 = Df(l)3A6-8t*3Bl-2 (Judd).
origin: X ray induced.
discoverer: Judd, 62b22. I(l)zw2b26
cytology: Salivary chromosomes normal (Judd). origin: X ray induced.
discoverer: Judd, 62b26.
cytology: Salivary chromosomes appear normal
origin: X ray induced in z-bearing X chromosome. (Judd).
discoverer: Judd, 64d8. I(1)zw2c2i
origin: X ray induced.
origin: X ray induced in z-bearing X chromosome. discoverer: Judd, 62c21.
discoverer: Judd, 64dl3. cytology: Salivary chromosomes appear normal
CJudd).
origin: X ray induced, I(1)zw2c28
discoverer: Judd, 63e6. origin: X ray induced,
cytology: Salivary chromosomes normal (Judd). discoverer: Judd, 62c28.

origin: X ray induced, origin: X ray induced,


discoverer: A bra hams on, 64 f2. discoverer: Abrahamson, 64f3.
MUTATIONS 129

I(1)zw2g4 I(l)zw6»2
origin: X ray induced, origin: X ray induced.
discoverer: Lefevre, 62g4. discoverer: Judd, 6212.
cytology: Salivary chromosomes appear normal phenotype: Lethal homozygous and when heterozy-
(Judd). gous with other alleles of t(l)zw6\ only allele of
I(l)zw6 that survives when heterozygous with
origin: X ray induced. Dt(l)62dl8 =Df(l)3B2'Cl;3C3-5 (Judd). RK2.
discoverer: Lefevre, 62g6. cytology: Salivary chromosomes normal (Judd).
phenotype: Lethal homozygous and when heterozy- I(l)zw7*3
gous with all alleles of I(l)zw2 except I(l)zw2a3. location: 1-1.4.
RK2 origin: X ray induced.
discoverer Judd, 63e3.
location: 1-1.3 [based on position of I(l)zw3h22]. cytology: Salivary chromosomes normal. Placed in
origin: X ray induced- region 3B3-C2, on basis of its inclusion in both
discoverer: Judd, 62bl2. Df(l)62dl8 =Df(l)3B2-Cl;3C3-5 and Df(l)wrJ2 =
cytology: Associated with In(l)l-zw3bl2 = ln(l)3A8- Df(l)3A6'8;3Cl'3 (Judd).
Bl;13; placed in region 3A7-B1, on the basis of its
inclusion in Dt(l)64j4 = Df(l)3A6-8;3Bl-2 (Judd). origin: X ray induced.
discoverer: Judd, 63g20.
I(l)zw3h22
origin: X ray induced. location: 1-1.1 [between l(l)zwl and I(l)zw2\.
discoverer: Judd, 62h22. origin: X ray induced.
cytology: Salivary chromosomes normal Qudd). discoverer: Judd, 63glO.
I(l)zw4d28 cytology: Placed in salivary chromosome region
location: 1-1.1 [between l(l)zwl and I(l)zw2\. 3A5-7, on the basis of its inclusion in Df(l)6464 *=
origin: X ray induced. Di(l)3A4-6;3C3-5 but not in Dffijw*"/2 =Df(l)3A6-
discoverer: Judd, 62d28. 8;3Cl-3 (Judd).
cytology: Placed in salivary chromosome region I(l)zw9f4
3A5-7, on the basis of its inclusion in Df(l)64c4 = location: 1-1.4.
Df(l)3A4-6;3C3-5 but not in Df(l)wtJ2 = Df(l)3A6- origin: X ray induced.
8;3Cl-3 (Judd). discoverer: Abrahams on, 64f4.
cytology: Placed in salivary chromosome region
origin: X ray induced. 3B3-C2, on the basis of its inclusion in both
discoverer: Judd, 63e4. Dt(l)62dl8 = Df(l)3B2-Cl;3C3-5 and Df(l)w'J2 =
cytology: Salivary chromosomes normal (Judd). D/jfl^d-S^Cl-;? (Judd).
l(2)39a
I(1)zw4s24 location: 2-50 (right of Bl ?).
origin: X ray induced. origin: Spontaneous,
discoverer: Judd, 62g24. discoverer: Curry, 39a.
references: 1939, DIS 12: 45.
location: 1-1.4. l(2)55i
origin: X ray induced. location: 2-55.0 (probably to the left of the centro-
discoverer: Judd, 62jl. mere).
cytology: Placed in salivary chromosome region origin: Spontaneous.
3B3-C2, on the basis of its inclusion in both discoverer. Burdick, 55L
Dt(l)62dl8 =*Df(l)3B2-Cl;3C3-5 and references: 1956, DIS 30: 69.
Df(l)3A6-8;3C1.3 (Judd). Mukai and Burdick, 1959, Genetics 44: 211-32.
1960, Genetics 45: 1581-93.
) Schnick, Mukai, and Burdick, 1960, Genetics 45:
location: 1-1.3. 315-29.
origin: X ray induced, Mukai and Burdick, 1961, Japan. J. Genetics 36:
discoverer: Judd, 62b23. 97-104.
cytology: Salivary chromosomes appear normal phenotype: Larvae hatch but die before pupation.
(Judd). Females heterozygous for l(2)55i have higher
65 fecundity than homorygous wild-type females. The
origin: X ray induced, lethal is therefore not eliminated from laboratory
discoverer: Judd, 63e5. populations. RK3.
l(2)$6a
origin: X ray induced, tocotion: 2-90.
discoverer: Judd, 63el3. origin: Spontaneous.
cytology: Salivary chromosomes apparently normal discoverer: Burdick, 56a.
(Judd). references: 1956, D3S 30: 69.
130 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

phenotype: Homozygous lethal; heterozygote shows opening puparium. Occasionally, a fly sponta-
normal viability. RK3. neously escapes puparium without serious injury.
other information: Crossing over normal. Differences in content of free amino acids and pep-
*l(2)57 tides between l(2)bl and wild type can be distin-
origin: Spontaneous. guished in third instar larvae, prepupae, and early
discoverer: Paik. pupae. RK3,
references: I960, Evolution 14: 293-303. l(2)Bld: lethal(2) from Blond
other information: A series of 11 let ha Is selected location: 2-53.1.
from Korean wild populations. origin: Spontaneous.
discoverer: Bridges.
*l(2)1076 l(2)bw: lethald) with brown
location: 2-15 (a,bout 40 units from Bl). location: 2-104.
origin: Spontaneous. origin: Spontaneous in 6w 2b mr chromosome.
discoverer: Ives, 49h. discoverer: Curry, 36i.
references: 1951, DIS 25: 70. cytology: Salivary chromosomes seem to show slight
phenotype: Lethal homozygous and in combination deficiency or disturbance in 59C and D (Bridges).
with ln(2L)Cy. RK3. I(2)C: lethal(2) o/ Curry
*l(2)1323 location: 2-67.1.
location: 2-55 (0/162 crossovers with Bl). origin: Spontaneous.
origin: Spontaneous. discoverer: Curry, 34a21.
discoverer: Ives, 51 g. phenotype: Lethal before pupation. RK3.
references: 1951, DIS 25: 70. cytology: Placed in salivary region 49D4 through
phenotype: Lethal homozygous and in combination 49E5 on the basis of its inclusion in Df(2R)v£B =
with In(2L)Cy + In(2R)Cy. RK3. Df(2R)49D3-4;50A2-3 and in Di(2R)v^ =
K2)a Df(2R)49Cl-2;49E2-6 (Morgan, Bridges, and
location: 2-64.7. Schultz, 1938, Carnegie Inst. Wash. Year Book 37:
origin: Spontaneous. 306).
discoverer Bridges, 16a 15. l(2)cg: lethal(2) with comb gap
'references: Bridges and Morgan, 1919, Carnegie location: 2-15 (between dp and of).
Inst. Wash. Publ. No. 278: 286, 302. origin: Spontaneous.
phenotype: Almost completely lethal; body color of discoverer: Nichols-Skoog, 33dl9.
rare survivor pale. RK3. references: Curry, 1939, DIS 12: 46.
K2)ax 1(2) cn bwco-3a: s e e l(2)S3a
locotion: 2-106.9. 1(2) cn bwco-7; S ee 1(2)S7
origin: Spontaneous. l(2)crc: lethal(2) cryptocephal
discoverer Bridges, 19b28. location: 2-55.
references: 1937, Cytologia (Tokyo), Fujii Jub. Vol. origin: Spontaneous.
2: 745-55. discoverer: Hadorn, 1942.
phenotype: Lethal in very early larval stage. RK3. synonym: crc.
cytology: Located in 60B on salivary chromosome by references: Hadorn and Gloor, 1943, Rev. Suisse
Bridges, but not included in Df(2R)Px = Zool. 50: 256-61.
D£(2R)60B8-10;60Dl-2. Gloor, 1945, Arch. Julius Klaus-Stift.
Way Vererbungsforsch. Sozialanthropol. Rassenhyg.
location: 2-8.3. 20: 209-56.
origin: Spontaneous. Fristom, 1965, Genetics 52: 297-318.
discoverer: Bridges, 30d5. phenotype: Homozygotes undergo pupation but rarely
K2)B eclose from puparia. Imaginal head is not
location: 2- [in 2L of In(2L)t]. everted from thorax. Except for slightly reduced
discoverer Bridges, 1930. eyes and shortened legs, wings, and thoracic
l(2)bh tethaK2) bluter bristles, the head and thorax are fully differen-
location: 2-43.8. tiated. Head eversion is inhibited by integument
origin: X ray induced. being more rigid than normal. Mutant integument
discoverer: Kafer, 50b. contains more glucosamine than normal. Feeding
references: Benz, 1953, DIS 27: 55. glue osa mine to wild-type larvae produces a pheno-
1957, Z. Induktive Abstammungs- Vererbungslehre copy very similar to l(2)crc. Abdomen often shows
88: 78-114 (fig.). no differentiation and internal organ development
phenotype: Lethal at end of pupal stage. Homozy- arrested at pupal stage. RK3.
gotes make emerging movements, but puparia have l(2)gh lethal{2) giant /crvoe
abnormally thick protein layer so that iraaginal location: 2-0.0.
hypodermis is punctured in attempt to eclose. origin: Spontaneous.
Hemoljmph is lost and flies die. Apparently normal discoverer Bridges, 33e9.
hocnoxygotes may be obtained by artificially Synonym: lgl.
MUTATIONS 131

references: Hadorn, 1937, Proc. Soc. Exptl. Biol. origin: Spontaneous.


Med. 36: 632-34. discoverer: Redfield, 23b.
1937, Proc. Natl. Acad. Sci. U.S. 23: 478-84. references: 1924, Am. Naturalist 58: 566—69.
1938, Rev. Suisse Zool. 45: 425-29. 1926, Genetics 11: 482-502.
Vogt, 1947, Z. Naturforsch. 26: 292-94. phenotype: Homozygous females produce one
phenotype: Homozygotes undergo embryogenesis and daughter to 5.5 sons. Abnormal sex ratio caused
three larval ins tars. Larvae reach normal maximum by inviability of females. l(2)mat does not seem to
size but fail to pupate; they then become bloated to be allelic to da, which has a similar effect. RK3.
giant size. Pseudopupae are sometimes formed but l(2)me: lethal(2) meander
no morphogenesis occurs, and imaginal disks location: 2-72 (71-73).
degenerate. Ring gland small and appears immature origin: Spontaneous.
in third-instar larva (Scharrer and Hadorn, 1938, discoverer: Hadorn, 44g20.
Proc. Natl. Acad. Sci. U.S. 24: 236-42). A third- synonym: Itno.
instar l(2)gl host transplanted with a normal ring references: 1947, Exptl. Biol. Symp. Vol. 2:
gland pupates but does not metamorphose. Thus a 177—95, Cambridge Univ. Press.
deficiency of hormones from the ring gland is 1947, DIS 21: 68.
probably one result of l(2)gl but not the only one. Schmid, 1949, Z. Induktive Abstammungs-
Faulhaber (1959, Z. Vererbungslehre 90: 299-334) Vererbungslehre 83: 220-53 (fig.).
finds that the abnormal development affects the Chen and Hadorn, 1954, Rev. Suisse Zool. 61:
quantities of the different amino acids, peptides, 437-51.
and proteins. Welch (1957, Genetics 42: 544—59) 1955, Rev. Suisse Zool. 62: 338—47.
finds that DNA of nuclei in several tissues, espe- phenotype: Larvae do not grow normally; die while
cially salivary glands, is markedly reduced. RK3. small. Body length remains relatively shorter than
cytology: Locus lies between 21A1 and 21C1 tracheal stems, which become convoluted in a
(Lewis, 1945, Genetics 30: 137-66). meandering manner. Salivary glands reach 30 per-
other information: The order of 1(2)0 and net cent normal size; pharyngeal development normal.
unknown. Intestines lack proteolytic enzymes. RK3.
K2)gl2 I(2)mr2: lethal(2) with morula
origin: Ultraviolet induced. location: 2-70.
discoverer: Meyer, 51a. origin: Spontaneous,
references: Meyer and Edmonds on, 1951, DIS 25: 72. discoverer: Bridges, 25k24.
phenotype: Like l(2)gl. RK3. *I(2)NS: lethal(2) Nova Scotia
*I(2)9I3 location: 2-107.0 [to the right of l(2)ax and to the
origin: Spontaneous in In(2L)Cy + In(2R)Cy. left of sp].
discoverer: Meyer, 51a. discoverer: Bridges, 23j31.
references: Meyer and Edmondson, 1951, DIS 25: 73. references: 1937, Cytologia (Tokyo), Fujii Jub. Vol.
phenotype: Larvae heterozygous with l(2)gl2 are like 2: 745-55.
1(2)0. RK3A. phenotype: Lethal when larvae are about 2 mm long.
I(2)H: lethal(2) of Humphrey Development of tracheae and other chitinized parts
location: 2-50. abnormal. RK3A.
origin: Spontaneous. cytology: Exists only as In(2R)NS, px 1(2)NS sp.
discoverer: Humphrey, 32k. Salivary chromosome locus in 60B10-12 on the
references: Dunn, 1934, DIS 1: 30. basis of its inclusion in Df(2R)Px » D%2R)60B8'
1935, DIS 4: 9. 10;60Dl-2 but not in the 2RPXD element of
phenotype: Usually dies as pupa; 10—15 percent of T(l;2)Bld= T(1;2)1C3-4;6OB12-13 (Bridges).
flies survive, look normal but are weak. Homozy- l(2)pm: lethal(2) polymorph
gote usually sterile when inbred but fertile in out- location: 2-30,3.
crosses. RK3. origin: X ray induced,
l(2)hst: lethal(2) histolytic discoverer: Kafer, 50b.
location: 2-56. references: Benz, 1953, DIS 27: 55.
origin: X ray induced. 1957, Z. Induktive Abstammungs- Vererbungslehre
discoverer: Thompson, 59k. 88: 78-114 (fig.).
phenotype: Homozygote dies in early pupal stage. phenotype: Flies die throughout larval and pupal
Heterozygous viability good. RK3. stages. Larvae do not contract before pupation;
I(2)M: lethal(2) from Moftr hence, pupae are long and thin. Imagos often
location: 2-(b«tween dp and 6). cryptocephalic. Chief characteristic is a severe
origin: Spontaneous. muscular dystrophy. Protein metabolism extremely
discoverer: Bridges, 33118. disturbed. In larval stage, free amino acids and
l(2)Mad51"": see 1(2)S one peptidc are in abnormally high concentration.
l(2)Mass38*-: see 1(2)S Prepupae only slightly different from normal in this
t(2)maf: tethal(2) maternal respect. Occasional survivors viable and fertile.
location: 2- (near pr). RK3.
132 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

*l(2)pup: lethai(2) pupal *l(2)S3a


location: 2-47. location: 2- (between dp and Sp).
origin: Spontaneous. origin: Gamma ray induced.
discoverer: Ives, 38J25. discoverer: Seto, 1953.
references: 1945, Genetics 30: 175. synonym: l(2)cn bwc°-3a; Co-3A.
1945, DIS 19: 46. references: 1956, J. Heredity 47: 21-27.
phenotype: Dies in middle or late pupal stage. Ex- 1958, DIS 32: 157-58.
ternal anatomy appears normal except for heavy 1961, DIS 35: 94-95.
melanization of wings and legs. RK3. 1963, DIS 37: 128-29.
phenotype: Die predominantly in pupal stage; infre-
I(2)R: lethal(2) of RecHield quent weak sterile survivors with normal pheno-
location: 2- [in 2L with In(2L)t]. type. Death occurs earlier in crowded cultures
discoverer: Redfield, 1933. (Seto, 1957, DIS 31: 160-62). Pupation delayed
*I(2)S1: lethal(2) of Seto 1—2 days beyond normal. Respiration rate during
location: 2- (not located). pupation 50 percent normal (Seto, 1959, DIS 33:
origin: Spontaneous. 159-60). RK3.
discoverer: Seto, 1951. *I(2)S4
synonym: itfyMadSln-1; N-l. location: 2- (near pr).
references: 1954, J. Exptl. Zool. 126: 17-32. origin: Spontaneous.
1954, Am. Naturalist 88: 373-78. discoverer: Seto, 1951.
1956, J. Heredity, 47: 21-27 (fig.). synonym: l(2)Mad51"-4; N-4.
1961, DIS 35: 94-95. references: 1954, J. Exptl. Zool. 126: 17-32.
phenotype: Dies as larva or prepupa. Puparium 1954, Am. Naturalist 88: 373-78.
elongated, often with larval segmentation, en- 1956, J. Heredity 47: 21-27 (fig.).
larged, resembling 1(2)0.. Cephalic complex un- 1958, DIS 32: 157-58.
everted. Eye rudiment often pigmented. RK3. 1963, DIS 37: 128-29.
*I(2)S1A phenotype: Dies in prepupal and pupal stages.
location: 2- (between Sp and 6). Puparium small and slender with fragile exoskel-
origin: Spontaneous. eion. Appearance like l(2)crc, with uneverted
discoverer: Seto, 1953. cephalic complex. Development of hypodermal
synonym: l(2)Mad53n-tA', N-1A. derivatives and pigment variable. Pupation delayed
references: 1956, J. Heredity 47: 21-27 (fig.). 1—2 days. Crowding suppresses expression (Seto,
1958, DIS 32: 157-58. 1957, DIS 31: 160-62). RK3.
1961, DB 35: 94-95. *I(2)S7
1963, DIS 37: 128-29. location: 2- (between Sp and b).
phenotype: Dies in late pupa; differentiation of ex- origin: Gamma ray induced.
ternal structures almost complete. Color of pupa discoverer: Seto, 1953.
darker, changing to deep brown after a few days; synonym: l(2)cn bwc°-7; Co-7.
heavy melanization in thicker parts of exoskeleton; references: 1956, J. Heredity, 47: 21-27.
nephrocytes deeply pigmented. Respiration rate 80 1957, DIS 31: 160-62.
percent normal (Seto, 1959, DIS 33: 159—60). 1958, DIS 32: 157-58.
Slight delay in pupation. Development ceases 1961, DIS 35: 94-95.
earlier under crowded conditions (Seto, 1957, DIS phenotype: Die predominantly in pupal stage. Occa-
31: 160-62). RK3. sional sterile adults produced. Time of pupation
*l(2)$3 delayed. Respiration rate of prepupae and pupae 50
location: 2-(between c and px). percent normal (Seto, 1959, DIS 33: 159-60). RK3.
origin: X ray induced. *K2)sn
discoverer: Seto, 1951. location: 2- (not located).
synonym: l(2)MasB38x-3; X-3. origin: X ray induced.
references: 1954, J. Exptl. ZooL 126: 17-32. discoverer: Seto, 1951.
1956, J. Heredity 47: 21-27 (fig.). synonym: itfytiassSS*-11; X-ll.
1957, DIS 31: 160-62. references: 1954, J. Exptl. Zool. 126: 17-32.
1958, DIS 32: 157-58. 1956, J. Heredity 47: 21-27.
1961, DIS 35: 94-95.
1961, DIS 35: 94-95.
1963, DJS 37: 128-29.
phenotype: Dies as pupa. Puparium normal. Devel-
phenotype: Dies in prepupal stage. Puparium small opment may cease before or shortly after eversion
and dumpy. Deformed prepupa only partly fills
of frontal sac. Usually has melanized patches on
pupariswj; anterior structures rudimentary; rest of
dorsum. Wings and legs variably developed. RK3.
body saclike. Scattered small pigmented or
meiamijwd art as along tracrteal trunks; appendages location: 2- (not located).
and cephalic complex reduced or absent. Some origin: Spontaneous,
delay in pupation. RK3. discoverer. Seto, 1955.
MUTATIONS 133

synonym: l(2)Wau55n-l3; N-13. condition. Most flies complete but only rarely able
references: 1957, DIS 31: 160-62. to eclose; rare adults are weak and unproductive.
1961, DIS 35: 94-95. Time of pupation delayed about one-half day. RK3.
phenotype: Dies as late pupa or as adult. Puparium *l(2)S50
normal. RK3. location: 2- (rearrangement).
*I(2)S32 origin: Spontaneous.
location: 2- (between dp and Sp), discoverer: Seto, 1951.
origin: Spontaneous. synonym: l(2)Mad51n"S0; N-50.
discoverer: Seto, 1951. references: 1954, J. Exptl. Zool. 126: 17-32.
synonym: l(2)Mad51n-32; pj-32. 1954, Am. Naturalist 88: 373-78.
references: 1954, J. Exptl. Zool. 126: 17-32. 1956, J. Heredity 47: 21-27 (fig.).
1954, Am. Naturalist 88: 373-78. 1957, DIS 31: 160-62.
1956, J. Heredity 47: 21-27 (fig.). 1958, DIS 32: 157-58.
1958, DIS 32: 157-58. 1961, DIS 35: 94-95.
1961, DIS 35: 94-95. phenotype: Dies as pupa. Puparium normal. Devel-
1963, DIS 37: 128-29. opment ceases in early pupa; body generally unpig-
phenotype: Dies in early pupa. Usually no pigment mented; eyes rarely pigmented; hypodermal deriva-
or bristle formation; leg and wing sacs adhere to tives underdeveloped; melanotic degeneration of
pupa case, resulting in appendages developing in hypodermis in region of eye, external genitalia, and
cramped position; some melanization at extremi- appendage extremities. Pupation delayed. RK3A.
ties. Pupation delayed 1 day; pupa badly shrunken. cytology: Associated with In(2L) and In(2R) with
Crowding suppresses expression (Seto, 1957, DIS unknown break points.
31: 160—62). Respiration rate 50 percent normal *I(2)S51
(Seto, 1959, DIS 33: 159-60). RK3. location: 2- (near pr).
*I(2)S42 origin: Spontaneous.
location: 2- (between Bl and L). discoverer: Seto, 1951.
origin: Spontaneous. synonym: l(2)Mad51n-sl; N-Sl.
discoverer: Seto, 1951. references: 1954, J. Exptl. Zool. 126: 17-32.
synonym: l(2)Mad51n-42; N-42. 1954, Am. Naturalist 88: 373-78.
references: 1954, J. Exptl. Zool. 126: 17-32. 1956, J. Heredity 47: 21-27.
1956, J. Heredity 47: 21-27. 1957, DIS 31: 160-62.
1958, DIS 32: 157-58. 1958, DIS 32: 157-58.
1961, DIS 35: 94-95. 1961, DIS 35: 94-95.
phenotype: Dies in late pupa. Appears well differen- phenotype: Dies in prepupal stage. Puparium
tiated externally; eyes with little or no pigment; normal. Development similar to that of 1(2)S61, but
internal head structures poorly developed. RK3. with certain tissues further developed. Pupation
*I(2)S42* delayed 1—2 days. Respiration less than 50 per-
origin: Spontaneous derivative of 1(2)S42. cent normal (Seto, 1959, DIS 33: 159-60). RK3.
discoverer: Seto, 1954. *I(2)SSS
synonym: l(2)Mad51n-<2a; N-42A. location: 2- (between dp and Sp).
references: 1956, J. Heredity 47: 21-27 (fig.). origin: Spontaneous.
1957, DIS 31: 160-62. discoverer: Seto, 1955.
1961, DIS 35: 94-95. synonym: l(2)Wau55nS*; N-55.
phenotype: Ceases development in early pupa; references: 1958, DIS 32: 157-58.
puparium enlarged, elongated, and larva like; exo- 1961, DIS 35: 94-95.
skeleton thin and fragile; pupa adheres to anterior phenotype: Dies in late pupa; occasional weak adult
end of puparium, res-t of pupa contracted anteriorly. survivors. Phenotype similar to 1(2)S45. Crowding
Space between pupa and puparium filled with light, results in more and earlier mortality (Seto, 1957,
oily fluid. Time of pupation delayed 2—3 days. DIS 31: 160—62). Pupation delayed about one-half
Imaginal disks poorly developed. RK3. day. RK3.
*I(2)U5 *I(2)$S9
location: 2- (between Sp and o). location: 2- (not located).
origin: Spontaneous. origin: Spontaneous.
discoverer: Seto, 1953. discoverer: Seto, 1951.
synonym: l(2)Mad53n-*3; N-45. synonym: l(2)Mad51n'^9; N-59.
references: 1956, J. Heredity 47: 21-27 (fig.). references: 1954, J. Exptl. Zool. 126: 17-32.
1957, DIS 31: 160-62. 1956, J, Heredity 47: 21-27.
1958, DBS 32: 157-58. 1961, D^ 35: 94-95.
1961, DIS 35: 94-95. phenotype: Dies in late larval or prepupal stage.
phenotype: Dies as late pupa just before eclosion. Puparium normal; frontal sacs uneverted; leg and
Slightly smaller than normal; leg sacs do not wing sacs incompletely developed; body sac like.
elongate, end legs develop in cramped and stunted RK3.
134 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

*I(2)S61 *l(2)SP9c
location: 2- (near pr). location: 2-55.1 (between rl and stw).
origin: Spontaneous. origin: Spontaneous.
discoverer: Seto, 1951. discoverer: Spiess.
synonym: l(2)Mad51n-6l; N-61. references: Spiess, Helling, and Capenos, 1963,
references: 1954, J. Exptl. Zool. 126: 17-32. Genetics 48: 1377-88.
1954, Am. Naturalist 88: 373-78. phenotype: Lethal homozygous and in combination
1956, J. Heredity 47: 21-27 (fig.). with M(2)S2; viable in combination with l(2)Spll
1957, DIS 31: 160-62. and 1(2)SP15. RK3.
1958, DIS 32: 157-58. cytology: Placed in salivary region 41A based on
1961, DIS 35: 94-95. its inclusion in D{(2R)M'S21 ° = Df(2R)41A
1963, DIS 37: 128-29. (Burdick).
phenotype: Dies in prepupal stage. Puparium color *l(2)SP9d
darker than normal. Prepupa incompletely de- location: 2-55.1 (to the right of stw).
veloped; frontal sacs uneverted; wing and leg sacs origin: Spontaneous.
everted but development curtailed; free-floating fat discoverer: Spiess.
body fragments may fill fluid space between pupa- references: Spiess, Helling, and Capenos, 1963,
rium and prepupa. Pupation delayed 1—2 days. Genetics 48: 1377-88.
Respiration rate normal for first day and a half *l(2)SP10
then ceases (Seto,1959, DIS 33: 159-60). RK3. location: 2-37.5.
*l(2)Sph leihol(2) of Spiess origin: Spontaneous.
location: 2-35.0. discoverer: Spiess.
origin: Spontaneous. references: Spiess, Helling, and Capenos, 1963,
discoverer: Spiess. Genetics 48: 1377-88.
references: Spiess, Helling, and Capenos, 1963,
Genetics 48: 1377-88.
K2)sPn
location: 2-55.1 [between rl and stw; to the left of
*l(2)SP2b
1(2)SP15 (Burdick)].
location: 2-49.
origin: Spontaneous.
origin: Spontaneous,
discoverer: Spiess.
discoverer: Spiess.
references: Spiess, Helling, and Capenos, 1963,
references: Spiess, Helling, and Capenos, 1963, Genetics 48: 1377-88.
Genetics 48: 1377-88. phenotype: Lethal homozygous and in combination
*l(2)SP6b with M(2)S2; viable in combination with l(2)Sp9c
location: 2-50.0. and I(2)apl5. RK3.
origin: Spontaneous. cytology: Placed in salivary region 41A based on
discoverer. Spiess. its inclusion in Df(2RyM-S2^ 0 = Df(2R)41A
references: Spiess, Helling, and Capenos, 1963, (Burdick).
Genetics 48: 1377-88. *l(2)SpU
I(2)SP7 location: 2-61.5.
location: 2-3.2. origin: Spontaneous.
origin: Spontaneous, discoverer: Spiess.
discoverer: Spiess. references: Spiess, Helling, and Capenos, 1963,
references: Spiess, Helling, and Capenos, 1963, Genetics 48: 1377-88.
Genetics 48: 1377-88. *l(2)SpU
*l(2)Sp8 location: 2-32.0.
location: 2-61.5. origin: Spontaneous.
origin: Spontaneous. discoverer: Spiess.
discoverer: Spiess. references: Spiess, Helling, and Capenos, 1963,
references: Spiess, Helling, and Capenos, 1963, Genetics 48: 1377-88.
Genetics 48: 1377-88.
*!(2)Sp9a i(2)$P15
location: 2-1.9. location: 2-55.1 (between rl and stw).
origin: Spontaneous. origin: Spontaneous.
discoverer Spiess. discoverer: Spiess.
references: Spiess, Helling, and Capenos, 1963, references: Spiess, Helling, and Capenos, 1963,
Genetics 48: 1377-88. Genetics 48: 1377-88.
l(2)Sp% phenotype: Lethal homozygous and in combination
location: 2-49. with M(2)S2; viable in combination with l(2)Sp9c
origin: Spontaneous. and l(2)Spll. RK3.
dl*cov»r#r. Spiess. cytology: Placed in salivary region 41A based on
r*f*r«nc«s: Spiess, Helling, and Capeno«, 1963, its inclusion in Df(2R.)M-S210 « Dt(2R)41A
Getseiics 48: 1377-8S. (Burdick).
MUTATIONS 135

*I(2)SP18 other information: Results of Bridges interpreted to


location: 2-65.3. show allelism to M(3)wB (3-79.7); may have been
origin: Spontaneous. related to maternal effect described by Schultz.
discoverer: Spiess. Position based on crosses by Muller (1918), in
references: Spiess, Helling, and Capenos, 1963, which he used l(3)a separated from In(3R)C.
Genetics 48: 1377-88. l(3)ac: lethal(3) accessory
l(2)Su(H): lethal(2) from Suppressor of Hairless location: 3- (midregion).
location: 2-99. discoverer: Schultz, 25g.
origin: Spontaneous. phenotype: Enhances maternal effect of In(3R)C,
discoverer: Bridges, 3717. l(3)a on recovery of M(3)w daughters from
cytology: Located in salivary region 58A1 through M(3)w/In(3R)C, l(3)a mothers (Schultz). RK3.
58F8 on basis of its inclusion in Df(2R)M-l = *l(3)blo-l: lethal(3) bloated larvae
Df(2R)57Fll-58Al;58F8-S9Al. location: 3- (to the left of p).
*I(2)T: lethal(2) of Thompson discoverer Bridges, 25k7.
origin: Spontaneous in normal chromosome of SM1/+ references: Chen, 1929, J. Morphol. 47: 135-99.
heterozygote. phenotype: Larvae become very large and trans-
discoverer: Thompson, 1956, 1957. parent; die in the prepupal stage. Growth of
synonym: I(2)56i24 through l(2)57hlO. imaginal disks irregular. RK2.
other information: A series of 13 independently 1(3)D1: see DP
occurring and genetically located lethals. *l(3)e: lethal(3) with ebony
l(2)Wau55n-: see 1(2)S location: 3- (not located).
1(3)1: see l(3)a origin: Spontaneous in In(3R)C, e.
1(3)26 discoverer: Schultz.
location: 3-52.2 [between l(3)S6 and 1(3)S7\. phenotype: Dies as fully developed normal-appearing
origin: X ray induced in a cu kat chromosome simul- imago unable to eclose. RK3A.
taneously with ry26.
discoverer: Schalet. *l(3)hd: lethal(3) head defect
synonym: 1(3)52.5.2. location: 3- (not located).
references: Schalet, Kernaghan, and Chovnick, discoverer: Bridges, 1924.
1964, Genetics 50: 1261-68. references: Morgan, Bridges, and Sturtevant, 1925,
other information: Placed between 1(3)S6 and 1(3)S7', Bibliog. Genet. 2: 230.
on the basis of its inclusion in Df(3R)ryK, phenotype: Dies in pupal stage with black tumorlike
Dt(3R)tvSi, and Df(3R)ry66 but not in Df(3R)ry2*', growth in head. RK3.
Df(3R)ry33, Df(3R)ry*2, or Df(3R)ry70f a l l of w hich I(3)PL
include 1(3)S6 but none of which include 1(3)S7. location: 3- (left arm).
l(3)36d!0 origin: Spontaneous in 3L carrying Itv(3L)P.
location: 3- (close to D, or rearrangement). I(3)PR
origin: Spontaneous, location: 3-90.2.
discoverer: Bridges, 36dlO. origin: Spontaneous inIn(3R)P.
*l(3)36d24
phenotype: Homozygous lethal; lethal in combination
location: 3- (near centromere),
with M(3)j. RK3.
origin: Spontaneous,
K3)S1: lethal(3) of Schalet
discoverer: Bridges, 36d24.
location: 3-51 (to the left of kar),
references: 1937, DIS 7: 13.
origin: X ray induced in a kar3 chromosome-
Bridges and Bridges, 1938, Genetics 23: 111-14.
discoverer Schalet.
1(3)52.52: see 1(3)26
other information: Placed to the left of l(3)S2, on
*l(3)62g
the basis of its exclusion from Dt(3R)ry76, which
origin: Spontaneous.
is deficient for 1(3)S2 and loci to the right.
discoverer: Paik.
references: 1963, Proc. Intern. Congr. Genet., 11th.
l(3)Slo
origin: X ray induced in a kar2 chromosome.
Vol. 1: 163-64.
discoverer: Schalet.
other information: A series of 65 lethals recovered
other information: Allelism with 1(3)S1 tentative and
from Korean wild populations.
K3)a based on similarity in interaction with D£(3R)ry7*.
location: 3-81.6. I(3)S2
origin: Spontaneous in In(3R)C location: 3-51.5 [between l(3)Sl and kar].
discoverer: Morgan, 111. origin: X ray induced in a kar2 chromosome.
synonym: 1(3)1. discoverer: Schalet.
references: Muller, 1918, Genetics 3: 422-99. references: Schalet, Kernaghan, and Chovnick,
phenotype: Lethal homozygou®. Reduces recovery of 1964, Genetics 50: 1261-68.
M(3)w-be&ring daughters from In(3R}C, l(3)s/M(3)w other information: Placed between l(3)St and kmr, on
females (Schultx). RK3. the basis of its inclusion in Df(3R)ty76 but not
D%3R)ry29, D%3R)ry33, or Df(3R)ry36. None of
136 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

these deficiencies include 1(3)S1 and all include other information: Produces local reduction in
/car and genes to its right. crossing over.
I(3)S3 I(3)S7
location: 3-51.7 (between kar and mes). location: 3-53 [to the right of 1(3)26].
origin: X ray induced in a kar2 chromosome. origin: X ray induced in a kar2 chromosome.
discoverer: Schalet. discoverer: Schalet.
other information: Placed between /car and mes, on references: Schalet, Kernaghan, and Chovnick,
the basis of its exclusion from Df(3R)ry78 and its 1964, Genetics 50: 1261-68.
inclusion in Df(3R)ry27, Df(3R)ryS2, Df(3R)ry7S, other information: Placed to the right of 1(3)26* on
and Df(3R)ry77. None of these deficiencies in- the basis of its exclusion from Df(3R)ry66, which
clude /car and all include mes and loci to its right. includes 1(3)26 and loci to its left.
I(3)S4 l(3)S7o
location: 3-52.1 (to the right of pic). origin: X ray induced in a kar2 chromosome.
origin: X ray induced in a kar2 chromosome. discoverer: Schalet.
discoverer: Schalet. references: Schalet, Kernaghan, and Chovnick,
references: Schalet, Kernaghan, and Chovnick, 1964, Genetics 50: 1261-68.
1964, Genetics 50: 1261-68. other information: Extends farther to the right than
phenotype: Homozygous lethal, but there a few rela- l(3)S7 since it is lethal in combination with an
tively normal-appearing survivors that are mostly undescribed deficiency for red with which l(3)S7
females. RK3. survives.
other information: Placed to the right of ry by re- l(3)S7b
combination and to the right of pic on basis of its origin: X ray induced in a kar2 chromosome.
survival in combination with ry3$ which behaves discoverer: Schalet.
as though it were deficient for ry and pic. Placed references: Schalet, Kernaghan, and Chovnick,
to the left of 1(3)S5 by recombination. 1964, Genetics 50: 1261-68.
I(3)S5 other information: Extends farther to the right than
location: 3-52.1 [between 1(3)S4 and 1(3)S6]. 1(3)S7, by same criterion as l(3)S7a.
origin: X ray induced in a kar2 chromosome. *l(3)S7c
discoverer: Schalet. origin: X ray induced in a kar2 chromosome.
references: Schalet, Kernaghan, and Chovnick, discoverer: Schalet.
1964, Genetics 50: 1261-68. references: Schalet, Kernaghan, arid Chovnick,
other information: Placed to the right of 1(3)S4 by 1964, Genetics 50: 1261-68.
recombination analysis and to the left of l(3)5>6, on *l(3)S7d
the basis of its inclusion in Df(3R)ry75 and origin: X ray induced in kar2 chromosome.
Dt(3R)ry76, which do not include 1(3)S6. discoverer: Schalet.
l(3)S5o references: Schalet, Kernaghan, and Chovnick,
origin: X ray induced in a kar2 chromosome. 1964, Genetics 50: 1261-68.
discoverer: Schalet.
references: Schalet, Kernaghan, and Chovnick, *l(3)Sph !ethal(3) of Spiess
1964, Genetics 50: 1261-68. location: 3-33.8.
I(3)S6 origin: Spontaneous.
location: 3-52.2 [between 1(3)S5 and 1(3)26]. discoverer: Spiess.
origin: X ray induced in a kar3 chromosome. references: Spiess, Helling, and Capenos, 1963,
discoverer: Schalet. Genetics 48: 1377-88,
references: Schalet, Kernaghan, and Chovnick, *I(3)SP2
1964, Genetics 50: 1261-68. location: 3-79.3.
other information: Placed to the right of 1(3)S5, on origin: Spontaneous.
the basis of its exclusion from Df(3R)ry7S and discoverer: Spiess.
D%3R)ry76, both of which include I(3)S5 and loci references: Spiess, Helling, and Capenos, 1963,
to the left. Genetics 48: 1377-88.
I(3)S6° *K3)$p5
origin: X ray induced in a kar2 chromosome. location: 3-41.0.
discoverer: Schalet. origin: Spontaneous,
references: Schalet, Kernaghan, and Chovnick, discoverer: Spiess.
1964, Genetics 50: 1261-68. references: Spiess, Helling, and Capenos, 1963,
other information: Causes local reduction in Genetics 48: 1377-88.
crossing over. *I(3)SP6
location: 3-40.4.
origin: X ray induced in a kar2 chromosome, origin: Spontaneous.
discoverer: Schalet. discoverer: Spiess.
references: Schalet, Kernaghan, and Chovnick, references: Spiess, Helling, and Capenos, 1963,
1964, Genetics 50: 1261-68. Genetics 48: 1377-88.
MUTATIONS 137

*I(3)SP9 phenotype: Lethal homozygous and in combination


location: 3-101.1. with Df(4)M. No interaction with other genes in
origin: Spontaneous. Df(4)M. Dies as larva. RK3.
discoverer: Spiess. cytology: Placed in region 101E through 102B16, on
references: Spiess, Helling, and Capenos, 1963, basis of its inclusion in Df(4)M =
Genetics 48: 1377-88. Df(4)101E-F;102B6-17.
*l(3)SP10 I (4) Jo
location: 3-41.7. origin: X ray induced.
origin: Spontaneous, discoverer: Gloor and Green, 1957.
discoverer: Spiess. synonym: 1(4)27.
references: Spiess, Helling, and Capenos, 1963, references: Hochman, Gloor, and Green, 1964,
Genetics 48: 1377-88. Genetica 35: 109—26.
*I(3)SP17 phenotype: Like 1(4)1. RK3.
location: 3-38.4. i(4)n
origin: Spontaneous. origin: X ray induced.
discoverer: Spiess. discoverer: Gloor and Green, 1957.
references: Spiess, Helling, and Capenos, 1963, synonym: 1(4)28.
Genetics 48: 1377-88. references: Hochman, Gloor, and Green, 1964,
*I(3)SP19 Genetica 35: 109-26.
location: 3-100.9. phenotype: Like 1(4)1. RK3.
origin: Spontaneous. 1(4)1 c
discoverer. Spiess. origin: X ray induced.
references: Spiess, Helling, and Capenos, 1963, discoverer: Gloor and Green, 1957.
Genetics 48: 1377-88. synonym: 1(4)32.
l(3)tr: lethal(3) translucida references: Hochman, Gloor, and Green, 1964,
location: 3-20 (18.1 to 22.0). Genetica 35: 109-26.
origin: Spontaneous. phenotype: Like 1(4)1. RK3.
discoverer: Hadorn, 40116. 1(4)1 d
references: 1947, Exptl. Biol. Symp. Vol. 2: origin: X ray induced.
177-95, Cambridge Univ. Press. discoverer: Gloor and Green, 1957.
1947, DIS 21: 68. synonym: 1(4)30.
1956, Cold Spring Harbor Symp. Quant. Biol. 21: references: Hochman, Gloor, and Green, 1964,
363-73 (fig.). Genetica 35: 109-26.
phenotype: Larvae become bloated and transparent phenotype: Like 1(4)1. RK3.
from accumulation of abnormal amount of hemo- other information: Incorrectly reported as an allele
lymph. Concentration of amino acids in hemolymph of 1(4)25 by Hochman, Gloor, and Green (1964).
higher than normal; concentration of proteins 1(4)1 -JFC: see 1(4)9°
reduced. Pupation delayed one day (25°C); dwarfed 1(4)2
pupae formed in inflated puparia; death follows location: 4-(not located),
pupation or completion of imaginal differentiation origin: X ray induced.
of head and thorax; abdomen never metamorphoses. discoverer: Gloor and Green, 1957.
After transplantation into normal hosts, imaginal references: Hochman, Gloor, and Green, 1964,
disks develop normally; ovaries also develop Genetica 35: 109—26.
normally and are fully capable of producing viable phenotype: Lethal in embryo. Lethal in combination
eggs [Sobels, 1950, Experientia 6: 139-40 (fig.)]. with the so-called btD fourth chromosome. RK3.
In pure oxygen, frequency and extent of imaginal 1(4)2"
differentiation strongly increased [Sobels and origin: X ray induced.
Nijenhuis, 1953, Z. Induktive Abstammungs- discoverer: Gloor and Green, 1957.
Vererbungslehre 85: 579-92 (fig.)]. RK3. synonym: 1(4)21.
cytology: Salivary chromosomes normal (Rosin). references: Hochman, Gloor, and Green, 1964,
Genetica 35: 109-26.
location: 3- (right.arm). phenotype: Like 1(4)2. RK3.
origin: Spontaneous in 3R carrying In(3R)P. l(4)2b
l(3)XaR origin: X ray induced.
location: 3-91.8. discoverer: Gloor and Green, 1957.
other information: Used to balance T(2;3)®px*. synonym: 1(4)23.
1(4)1 references: Hochman, Gloor, and Green, 1964,
location: 4- [within Df(4)Ml Genetica 35: 109-26.
origin: X ray induced. phenotype: Unlike other alleles, dies as larva.
discoverer: Gloor and Green, 1957. l(4)2b/I(4)2h heterozygotes survive and have
references: Hochman, Gloor, and Green, 1964, narrow bodies both as pupae and imagos. Viability
Genetica 35: 109-26. reduced; fertile. RK3.
GENETIC VARIATIONS OF OROSOPHILA MELANOGASTEfi
138

I(4)2'>*D: leihal(4) 2 in bent Dominant 1(4)4"


origin: Spontaneous. Associated with supposedly origin: X ray induced.
btD chromosomes. discoverer. Gloor and Green, 1957.
references: Fox, 1947, DIS 21: 85. synonym: 1(4)18.
Hochman, Gloor, and Green, 1964, Genetics 35: references: Hochman, Gloor, and Green, 1964,
109-26. Genetica 35: 109-26.
phenotype: Lethal homozygous and in combination phenotype: Like 1(4)4. RK3.
with other 1(4)2 alleles. RK3. l(4)4b
1(4)2* origin: X ray induced.
origin: X ray induced. discoverer: Gloor and Green, 1957.
discoverer: Gloor and Green, 1957. synonym: 1(4)20.
synonym: 1(4)29. references: Hochman, Gloor, and Green, 1964,
references: Hochman, Gloor, and Green, 1964, Genetica 35: 109-26.
Genetica 35: 109-26. phenotype: Like 1(4)4. RK3.
phenotype: Like 1(4)2. RK3. l(4)4c
l(4)2d origin: Spontaneous.
origin: X ray induced. discoverer: Hochman, 61a.
discoverer. Gloor and Green, 1957. synonym: l(4)PT-3: lethal(4) Powell, Tennessee-3.
synonym: 1(4)37. references: 1961, Am. Naturalist 95: 375-82.
references: Hochman, Gloor, and Green, 1964, 1963, DIS 37: 45.
Genetica 35: 109-26. Hochman, Gloor, and Green, 1964, Genetica 35:
phenotype: Like 1(4)2. RK3. 109-26.
1(4)2* phenotype: Like 1(4)4. RK3.
origin: X ray induced. l(4)4d
discoverer. Gloor and Green, 1957. origin: Spontaneous.
synonym: 1(4)38. discoverer Lipe, 62k.
references: Hochman, Gloor, and Green, 1964, synonym: l(4)AM-2: lethal(4) Amherst, Massachu-
Genetica 35: 109-26. setts-2.
phenotype: Like 1(4)2. RK3. references: Hochman, Gloor, and Green, 1964,
1(4)2* Genetica 35: 109-26.
origin: Spontaneous. phenotype: Like 1(4)4. RK3.
discoverer Hochman, 61a.
synonym: l(4)PT-2: IeOial(4) Powell, Tennessee-2. 1(4)5
references: 1961, Am. Naturalist 95: 375-82. location: 4- (not located).
1963, DIS 37: 48. origin: X ray induced.
Hochman, Gloor, and Green, 1964, Genetica 35: discoverer: Gloor and Green, 1957.
109-26. references: Hochman, Gloor, and Green, 1964,
phenotype: Like 1(4)2. RK3. Genetica 35: 109-26.
1(4)2* phenotype: Homozygotes die as embryos. RK3.
origin: Spontaneous. 1(4)6
discoverer; Hochman, 63kl9. location: 4- (not located).
pti«rtofyp«: Like 1(4)2. RK3. origin: X ray induced.
discoverer: Gloor and Green, 1957.
<w!§i«s Spontaneous. references: Hochman, Gloor, and Green, 1964,
discoverer: Hochman, 63122. Genetica 35: 109-26.
•Imnetyp'e: Lethal &a embryo. In combination with phenotype: Homozygotes die as larvae. RK3.
l(4)2b, produce* a few survivors that have narrow 1(4)6°
bodies both as pupae ami imagos and are fertile. origin: X ray induced.
discoverer: Gloor and Green, 1957.
synonym: 1(4)36.
oriigin: 1 ray induced. references: Hochman, Gloor, and Green, 1964,
l i e o v m r : Gloor an-d Gr®en, 1947.
Genetica 35: 109—26.
sytKMtym: t{4)34,
phenotype: Homozygotes die as pupae. RK3.
references: Hochm&n, Gloor, aad Green, 1964,
l(4}6t>
Ge»@tfca 35: 109-26.
origin: Spontaneous.
toe«fi«n: 4- (sot located). discoverer: Hochman, 61a.
•riffn: X ny induced, synonym: 1(4)PT-1; lethal(4) Powell, Tennessee-1.
disc»v««*r: Gloor tad Green, 1957. references: 1961, Am. Naturalist 95: 375—82.
1963, DIS 37: 48.
<wf«nmc«s: Hodman, Gloor, aed Green, 1964,
Hochman, Gloor, and Green, 1964, Genetica 35:
GHMKIC* 35: 109-26.
109-26.
it#« di® as larvae. RK3.
phenotype: Homozygotes die as pupae. RK3.
MUTATIONS 139

*1(4)6 c phenotype. Thus, whereas the at locus is within


origin: X ray induced, Df(4)M, 1(4)9 b is not.
discoverer: Gloor and Green, 1957. 1(4)9-
synonym: 1(4)15. origin: Spontaneous.
references: Hochman, Gloor, and Green, 1964, discoverer Hochman, 64a4.
Genetica 35: 109-26. phenotype: Like 1(4)9. RK3.
1(4)7 1(4)10
location: 4- (not located). location: 4- (not located).
origin: X ray induced. origin: X ray induced.
discoverer: Gloor and Green, 1957. discoverer: Gloor and Green, 1957.
references: Hochman, Gloor, and Green, 1964, synonym: 1(4)33.
Genetica 35: 109-26. references: Hochman, Gloor, and Green, 1964,
phenotype: Homozygotes die as embryos. RK3. Genetica 35: 109—26.
1(4)8 1(4)10: see 1(4)8*
location: 4- (not located). 1(4)11: see Di(4)ll
origin: X ray induced. 1(4)12: see Df(4)12
discoverer: Gloor and Green, 1957. 1(4)13
references: Hochman, Gloor, and Green, 1964, location: 4- [within Df(4)M\.
Genetica 35: 109—26* origin: X ray induced.
phenotype: Homozygotes die as pupae. RK3. discoverer: Gloor and Green, 1957.
*l(4)8° references: Hochman, Gloor, and Green, 1964,
origin: X ray induced. Genetica 35: 109-26.
discoverer: Gloor and Green, 1957. phenotype: Homozygous lethal. Lethal in combina-
synonym: 1(4)10. tion with Df(4)M and ciD but not with Df(4)17,
references: Hochman, Gloor, and Green, 1964, Df(4)34, or 1(4)18. 1(4)13/+ is normal. RK3.
Genetica 35: 109—26. cytology: Placed in region 101E through 102B16, on
phenotype: Homozygous lethal. RK3. basis of its inclusion in Dt(4)M =
Df(4)101E-F;102B6-17.
origin: X ray induced. 1(4)14
discoverer: Gloor and Green, 1957. location: 4- (not located).
synonym: 1(4)19. origin: X ray induced.
references: Hochman, Gloor, and Green, 1964, discoverer: Gloor and Green, 1957.
Genetica 35: 109-26. references: Hochman, Gloor, and Green, 1964,
phenotype: Like 1(4)8. RK3. Genetica 35: 109-26.
1(4)9 phenotype: Homozygotes die as larvae. RK3.
location: 4- [within Df(4)ti\. 1(4)14"
origin: Spontaneous. origin: X ray induced.
discoverer: Stowell, 62k. discoverer. Gloor and Green, 1957.
synonym: 1(4)BU-1: lethal(4) Bountiful, Utah-1. synonym: 1(4)26.
references: Hochman, Gloor, and Green, 1964, references: Hochman, Gloor, and Green, 1964,
Genetica 35: 109-26. Genetica 35: 109-26.
phenotype: Lethal homozygous and when heterozy- phenotype: Like 1(4)14. RK3.
gous with D£(4)3, Di(4)ll, Df(4)12, Dt(4)24, 1(4)14*
Df(4)34, Df(4)G, and spa c «<. RK3. origin: Spontaneous.
* 1(4)9° discoverer: Wrathall, 611.
origin: Spontaneous. synonym: l(4)ST-2: lethal(4) Solway, Tennessee-2.
discoverer: Crow. references: Hochman, 1963, DIS 37: 48.
synonym: 1(4)1-JFC: lethal(4) of /. F. Crow. Hochman, Gloor, and Green, 1964, Genetica 35:
references: Hochman, Gloor, and Green, 1964, 109-26.
Genetica 35: 109-26. phenotype: Like 1(4)14. RK3.
phenotype: Like 1(4)9. RK3. 1(4)15
1(4)9* location: 4- (not located).
origin: Spontaneous. origin: Spontaneous.
discoverer: Hochman, 61 e. discoverer: Grandmann, 62b.
synonym: l(4)ar: lethal(4) in chromosome containing synonym: l(4)ST-4: Mhal(4) Solway, Tmn@&Mee-4.
abdomen rota Cum. references: Hochman, 1963, DIS 37: 49.
references: Hochman, Gloor, and Green, 1964, Hochman, Gloor, and Green, 1964, Genetica 35:
Genetica 35: 109-26. 109-26.
ph*notype: Like 1(4)9. RK3. phenotype: Most homozygotes die as pupae. A few
other information: Proof that the lethal is not at the survive, especially in uncrowded cultures. Sur-
ar locus comes from the observation that vivors have spread wings and minor vein abnor-
D%4)M/ar 1(4)9*> survives and exhibits the ar malities such as crossveins between L2 and L3;
140 GENETIC VARIATIONS OF DROSOPHILA MELANOGASTER

males lack external genitalia but produce motile synonym: l(4)AM-3: lethal(4) Amherst, Massachu-
sperm; both sexes sterile. RK3. setts-3.
1(4)15: see 1(4)6^ references: Hochman, Gloor, and Green, 1964,
1(4)15" Genetica 35: 109—26.
origin: Spontaneous. phenotype: Homozygotes die as embryos. RK3.
discoverer: Wrathall, 62a. 1(4)23: see 1(4)21>
synonym: 1(4)SLC-1: lethal(4) Salt Lake City-1. 1(4)24: see Df(4)24
references: Hochman, 1963, DIS 37: 49. 1(4)25
Hochman,, Gloor, and Green, 1964, Genetica 35: location: 4- [within Df(4)M\.
109-26. origin: Spontaneous.
phenotype: Like 1(4)15. RK3. discoverer: Hochman, 62a.
synonym: l(4)ST-3: lethal(4) Solway, Tennessee-3.
l(4)15t> references: 1963, DIS 37: 48-49.
origin: Spontaneous. Hochman, Gloor, and Green, 1964, Genetica 35:
discoverer: Lipe, 621. 109-26.
synonym: 1(4)MW-1: lethal(4) Madison, Wisconsin-1. phenotype: Homozygotes die as larvae. RK3.
references: Hochman, Gloor, and Green, 1964, cytology: Placed in salivary chromosome region
Genetica 35: 109-26. 101E through 102B16, on basis of its inclusion in
phenotype: Like 1(4)15. RK3. Df(4)M = Df(4)101E-F;102B6-17.
1(4)17: see Df(4)17 other information: Incorrectly reported as an allele
1(4)18 of 1(4)1* by Hochman, Gloor, and Green (1964).
location: 4- [within Df(4)M63*]. 1(4)26: see 1(4)14*
origin: X ray induced. 1(4)27: see 1(4)1*
discoverer: Gloor and Green, 1957. 1(4)28: see 1(4)1 *>
synonym: 1(4)35. 1(4)29
references: Hochman, Gloor, and Green, 1964, location: 4- Lwithin Dt(4)G\.
Genetica 35: 109-26. origin: Spontaneous.
phenotype: Homozygous lethal. 1(4)18/ ci is cf; discoverer: Hochman, 62k.
l(4)18/M(4)63a is lethal; l(4)l8/ciD is viable. synonym: l(4)BU-2: lethal(4) Bountiful, Utah-2.
Abo