Beruflich Dokumente
Kultur Dokumente
'if.;^
;'
.-vi'
GENET
1
1
ANlMTRODUCTION'ljjiiP
rHE
S TV D Y
f^
:|H|i!!l!i|
H E RE D T
I
Iji
Ijp
^. p.
pU lltirrarg
0H45I
W5
fwC.
ST,
S00141938
T
Date Due
3 j ieMay 58 !
29Feb'36
S^'S pt
2At.^
270cV?f
\
I
2 1 1965
<,!
JUL 5
1967
V968
^ gp fa
m,>?o,n
3 01^1^
IWay^S
22mar5|(t
44
3,-
^'
17061
GENETICS
BOSTON
CHICAGO
SAN FRANCISCO
MACMILLAN &
LONDON
CO., Limited
BOMBAY CALCUTTA
MELBOURNE
THE MACMILLAN
CO. OF
TORONTO
CANADA,
Ltd.
GENETICS
AN INTRODUCTION TO THE STUDY
OF HEREDITY
BY
BROWN UNIVERSITY
WITH
11 C.
72
COLLEGE OF
A.
& M. 1.
Dept. of Botany
COPYKIGHT, 1913,
up and electrotyped.
J. S.
Gushing Co.
Berwick & Smith Co.
Norwood, Mass., U.S.A.
*u, /ynr..
^ep^
THIS
IS
*.*
VOLUME
AFFECTIONATELY DEDICATED
TO
MY MOTHER
if.-.
??
?.*
PREFACE
The
had their
upon Heredity, given
following pages
of lectures
sity during
origin in a course
at
Brown Univer-
mer
biological
of Arts
summer
and
Long
Sciences.
An
hoped that
summary
It
is
will
heredity.
The
many
regarding
it
who wish
to
is
technical readers.
It
may
the jargon of
vii
PREFACE
viii
my
The
by
my
which were at
critics have
kindly
H. E. W.
Providence, R.
I.,
September, 1912.
CONTENTS
CHAPTER
I.
PAGE
Introduction.
1.
The
2.
3.
4.
II.
life
'
5
10
Introduction
2.
The
3.
4.
Mitosis
5.
Amitosis
cell
14
theory
14
....
....
15
typical cell
18
20
6.
Sexual reproduction
7.
Maturation
22
8.
Fertilization
24
20
Parthenogenesis
26
27
13.
14.
Conclusion
35
9.
10.
11.
12.
III.
triangle of
definition of heredity
28
29
33
Variation.
1.
2.
3.
36
37
their
a.
Nature
38
h.
Duplication
39
39
c.
Utility
d.
Direction in evolution
39
e.
Source
40
40
/.
Normality
g.
Degree
h.
Character
i.
j.
of continuity
40
41
41
Heritability
41
IX
CONTENTS
PAGE
CHAPTER
IV.
4.
Methods
5.
Biometry
6.
Fluctuating variation
7.
The
...
......
......
.......
....
......
......
......
....
......
.......
......
.......
.......
....
.....
....
a.
Relative variability
b.
Bimodal curves
c.
Skew polygons
43
47
47
47
50
8.
Graduated and
9.
The
causes of variation
52
a.
Darwin's attitude
52
b.
c.
Lamarck's attitude
Weismann's attitude
d.
Bateson's attitude
integral variations
Mutation.
52
53
55
55
1.
2.
57
3.
Freaks
58
4.
Kinds
5.
Species
6.
63
7.
64
8.
9.
10.
V.
42
42
of studying variation
of
mutation
and
A summary
varieties
of the
mutation theory
....
....
.....
...
.....
.....
......
3.
4.
An
5.
Confusion in definitions
6.
2.
7.
genetics
9.
10.
59
60
67
69
72
74
75
75
76
77
78
79
ters
8.
56
characters
Weismann's reasons
for
to vary or
.
to
.
.80
acquire
.
.81
.....
acquired characters
84
CONTENTS
XI
PAGE
CHAPTER
11.
12.
somatic characters
inconclusive
Environmental
c.
The
d.
Disease transmission
88
effects
91
14.
The
15.
Conclusion
92
sufficient to
94
....
opposition to
84
87
b.
13.
Weismann
95
96
Line.
1.
The
2.
3.
The
unit character
method
97
of attack
98
102
4.
103
5.
107
6.
Tower's potato-beetles
108
7.
Jennings'
work on Paramecium
Phenotypical and genotypical distinctions
The distinction between a population and a pure
Pure lines and natural selection
110
8.
9.
10.
VII.
86
Mutilations
The Pure
a.
facts of heredity
VI.
113
line
115
118
2.
120
a.
Blending inheritance
b.
Alternative inheritance
c.
Particulate inheritance
120
121
121
121
123
128
6.
7.
8.
132
Dihybrids
133
3.
4.
5.
9.
10.
124
130
131
132
CONTENTS
Xll
CHAPTER
11.
Vin.
The
PAGE
140
12.
Conclusion
143
13.
Summary
144
New
Ones.
1.
The
2.
False reversion
distinction
.....
146
149
....
a.
Arrested development
149
6.
Vestigial structures
149
c.
d.
Convergent variation
e.
Regression
.....
....
3.
Explanation of reversion
4.
Some methods
of
150
151
151
types
a.
b.
c.
d.
e.
/.
9-
5.
The
152
152
153
154
155
155
156
....
157
factor hypothesis
159
a.
160
b.
163
c.
6.
7.
The kinds
8.
Conclusion
164
165
d.
IX.
150
.....
......
166
169
of gray rabbits
171
173
Blending Inheritance.
1.
The
2.
Imperfect dominance
3.
Delayed dominance
" Reversed " dominance
4.
relative value of
......
......
174
175
177
178
CONTENTS
Xlll
CHAPTER
PAGE
5.
Potency
179
a.
Total potency
179
b.
Partial potency
180
c.
Failure of potency
180
6.
Blending inheritance
182
7.
183
8.
9.
186
to the
193
X.
Human
The Determination of
196
Sex.
modern
1.
2.
3.
The
The
4.
Monochorial twins
5.
Selectiv^e fertilization
6.
7.
197
nutrition theory
statistical
198
study of sex
200
201
202
205
of sex
Microscopical evidence
207
1.
207
2.
208
3.
210
b.
210
c.
Sex-limited inheritance
213
d.
XI.
skin color
1.
Color-blindness
214
2.
216
220
222
Conclusion
The
2.
Modifying factors
3.
Experiments
a.
6.
c.
application of genetics to
in
human
....
....
man
man
the case of
in
heredity
The Jukes
The descendants of Jonathan Edwards
The Kallikak family
human
5.
The
6.
Hereditarv defects
character of
traits
.
like
225
227
227
.
228
229
physical
.....
.
224
230
231
232
CONTENTS
XIV
CHAPTEB
7.
The
8.
Inbreeding
....
The
11.
4.
240
influence of proximity
Human Conservation.
1. How mankind may
3.
235
effects of inbreeding
10.
2.
PAGE
238
9.
XII.
control of defects
241
242
......
be improved
c.
An
d.
The
e.
Drastic measures
The
a.
b.
c.
6.
Who
educated sentiment
subsidizing the
247
248
250
252
254
By
By
By
245
251
segregation of defectives
244
255
256
fit
258
258
1.
Preventable death
2.
Social hindrances
shall sit in
judgment
258
259
260
XIII,
Bibliography
263
XIV.
Index
265
GENETICS
GENETICS
CHAPTER
INTRODUCTION
1.
The
nine-
teenth century was Darwin's century. His monumental work "On the Origin of Species by Means of
Natural Selection," which appeared in 1859, not only
dominated the biological sciences but also influenced
profoundly many other realms of thought, particularly those of philosophy and theology.
Now, at the beginning of the twentieth century,
a particular emphasis is being laid upon the study
of
heredity.
The
interpretation
of
investigations
cell,
vestigators, inspired
nOF&RTY UBRARY
CBe|i
C. State
n,
GENETICS
is
HER!
Fig.
genetics
is
1.
The
chiefly concerned,
acteristics of
triangle of
life.
the individual.
individual,
diagrammatically in Figure 1. It
may indeed be said that an individual is the result
of the interaction of these three factors since he
tage as expressed
may be
Heritage, or
INTRODUCTION
*'
vidual.
It
is
shall
It
is
his nature.
diagram
(Fig.
the triangle of
1),
as resting solidly
upon the
side
life is
represented
marked "heritage"
Environment and
is
sents
what he
is, is
far
more important
GENETICS
tion
may
proved blood
will better
2.
Ex-
Definition of Heredity
generation to another.
his
father's
does not
hair
mean
and
is
When
his
mother's complexion
it
quences.
is more comparable to the
handing dow^n from father to son of some valuable
patent right or manufacturing plant by means of
which the son, in due course of time, may develop an
independent fortune of his own, resembling in character and extent the parental fortune similarly derived
although not identical with it.
So it comes about that "organic resemblance"
Biological inheritance
INTRODUCTION
common
are
source.
To
unravel
the
lives.
golden
how
all
to
threads
of
inheritance
of the
animals and
and
new
organic designs of an
undreamed
of,
is
3.
or whether
life
itself
was the
original condition of
GENETICS
is
tell
us of the probable
evolution of worlds, the theory of spontaneous generation seems to be the last resort to which to turn.
generation presented
half appears
And rampant
("
Paradise Lost,"
Book VII,
line 543.)
INTRODUCTION
There are various methods
producing more
of
life,
even as
man
is
made
frequently
good,
When
if it is
growth
results,
Thus a fragment
is
not too
wound.
of the outgo,
before.
of repair,
for example,
is
in excess
of a
living sponge or a
it sprang.
When two daughter protozoans are thus formed, they are essentially orphans
because they have no parents, alive or dead. The
from which
over bodily into the next generation in the formation of the daughter-cells, leaving
continuous
it
life
is
does occur,
the natural
is,
as
order,
Weismann has
GENETICS
In these cases it is easy to see the reason for "organic resemblance" between successive generations.
Parent and offspring are successive manifestations
of the same thing, just as the begonia plant, restored
from a fragment of a begonia leaf, is simply an extension of the original plant.
Many
by
in the
parents.
grafting
are continuations
of
tected
INTRODUCTION
when springtime
The unity
maintained.
The bryozoan
mud.
as
may
from these spores, to the parents which produced them, is due to the essential material identity
of two generations.
These illustrations of heredity in its simplest manifestations give the key to "organic resemblance"
higher up in the scale.
Sexual reproduction is no
less plainly the direct continuation of life though in
this instance tivo sporelike fragments out of one
generation contribute to form the new individual of
the next generation instead of one fragment. In all
arise
cases there
generations.
is
of
GENETICS
10
4.
is
is,
is
fated in the
The germplasm, on
the contrary,
is
and
die.
the immortal
it
arise
it
is
individual
a continuous sub-
Although this
theory of the continuity of the germplasm has been
actually demonstrated in comparatively few instances,
all
the facts
many
of the
it.
of the
INTRODUCTION
of the individual,
or in reserve, like
and
11
remains undifferentiated,
a savings-bank account put by
this
day,
for a rainy
plasm
expended
in the immediate
is
demands
up the
the
of
tissues that
make
individual.
In one instance at
that of the
least,
worm
nematode
Ascaris,
con-
as
firmed by Boveri,
this
splitting
off
or isolation of the
germplasm occurs
with the very first
cleavage of the
fertilized
the
stage,
of
egg into
two-celled
when one
the two
cells
by
succes-
the animal
Fig. 2.
Scheme to
V Somatoplasm
germplasm.
itself.
Thus there
plasm,
Germplasm
results a continuous
receiving
contributions
GENETICS
12
are
this continuous
by triangles. From
germplasm there split off
represented
stream of
individuals," which go so far on the road of specialitissues that the power to be " born
zation into
again "
is
lost,
and so
somatoplasms are developments at different intervals from the same continuous stream of germplasm
instead of one somatoplasm being derived from a
preceding one. As a matter of fact the germplasm
from which the son arises is modified by the addition
of a maternal contribution, so that father and son in
reality hold the
same
brothers do.
From
all
is
so marvelously
we
call
INTRODUCTION
is
13
of each
of the hereditary
processes.
may
when
that in
posite
may
crossed
soms.
produce
is
The new
heredity,
embodied
in
the science of
genetics,
much
It is con-
with somatoplasm, which represents what the germplasm has done in the past, as
with the germplasm and what it can do in the future.
cerned not so
CHAPTER n
THE CARRIERS OF THE HERITAGE
Introduction
1.
chapter,
may be direct,
as
when
This
a mother protozoan
common stream of
germplasm.
It is the purpose of the present chapter to consider
this material continuity between succeeding generaare somatoplasms derived from a
tions
and to
discover,
To
this
to take
end
it
up what
2.
will
is
meant by the
"cell theory."
had
made up
its birth.
14
of cellular units,
15
little
cell,
often
3.
typical
Typical Cell
undifferentiated
the
cell
the
nucleus
is
cell
is
represented
diagrammatically in Figure
3.
wall
Cefl
C^to plasm
Centrosome
Nuclear membrane
--Nucleus
.-Chromatin network
Fig. 3.
nuclear membrane.
the
enveloping
Diagram
The
of a typical
cell.
common with
made up of living
nucleus, in
cytoplasm,
is
GENETICS
16
wall or
membrane which
from another.
in the
The
ment
new
Stentors,
ment
The
nucleus,
it
should be said,
is
is
made up
easily
of
more
demonstrated
by
material,"
is
is
"colored
all
the various
any one
cells
that
make up
17
the
chromosomes
number with some
species these
in
for
any
may
as the centrosome.
At
cell
often be identified
a tiny
body known
of a cell the
GENETICS
18
it
is
apparently impossible, as
In
such instances the outfit of cells provided in the embryo determines the numerical limit of the cells
When this supply is exavailable throughout life.
hausted no more cells appear to replace those w^hich
out.
4.
The ordinary
out of one
is
process
termed
Mitosis
by which two
cells
are
made
It occurs constantly,
mitosis.
The resting
cell (Fig.
4)
is
characterized
by the
6)
moved
farther
is
of the nuclear
19
membrane and the more compact form of the chromosomes. At the end of the prophase (Fig. 8) the chromosomes have come to lie at the equator of the cell.
Fg. 4.
TFie
resting cell
n^.7 Prophase
Pig. 10.
Fl J.6. Earlij
ng.8. End
nj.9. Metaphase
Beginning Anaphase
R^.12
Figs. 4-13.
Prophase
Rg.ll.
Beginning Telophase
Diagrams
of
Prophase
Anaphase
illustrating mitosis.
After Boveri.
being connected by the mantle fibers with the centrosomes, each of which has now come to occupy a
polar position. In the metaphase (Fig. 9) the chromo-
somes
split lengthwise,
of the
GENETICS
20
cell
telophase (Fig.
chromosomes grow
the half
completed.
is
12)
the
In
half
and
the division of the cell body into two parts becomes
complete. The mantle fibers have disappeared and
the nuclear membrane begins to re-form around the
chromosomes. Finally, at the end of the telophase
(Fig. 13) the nuclear membrane becomes complete,
the chromosomes break up into a chromatin network,
and two resting cells take the place of the single one
with which the process began (Fig. 4).
until they attain full size
5.
Amitosis
to
degeneration.
of the
When
amitosis
on the road
takes
of mitosis
the place
it
is
fre-
cell.
Sexual Reproduction
by which one
21
cell is trans-
typically the
nucleus, a fact
Beneden
The
in 1883.
egg-cell
is
more
which
it
is
They
are,
effect fertilization.
fertilization.
GENETICS
22
is
now known
7.
Maturation
Certain
nature,
tion.
wise be double.
When
number
of
in
23
tions.
Primordial
Manvj similar
Sex Ceils
divisions
cell
Maturation=\^
^
Sperinatocyte
Spermatids
1- OocijU
2.-Oocijte
'^bortiva
Fig. 14.
Scheme to
illustrate
E^*
maturation of germ-cells.
GENETICS
24
ing
cell),
is
formed by
cell),
characteristic
zygote (yoked
cell).
in the
diagram) remains constant in each germ-cell respectively until the division of spermatids into spermatozoa, and of the second oocytes into mature eggs and
second polar cells, when it is reduced to one half the
normal number. As spermatozoan and mature egg
unite in fertilization, the original number of chromosomes is restored in the fertilized egg (zygote)
of
8.
The
tion,
Fertilization
15 to 23.
fJWPEMT UBRARt
Pig.lS. Entn^ of
fiig.18.
Sperm
Approach of Sperm
Splitting of
of
Sperm
Chromosomes
Division of Cenlrosome
Fi^.ZO. Formation of
Chromosomes
Nucleus
Nucleus
Fi*g.2.I.
R^.IZ
H^. 2. Anaphase
n^.SS.Two-celled Sla^e
Figs.
15-23. Diagrams
illustrating fertilization.
25
After Boveri.
GENETICS
26
is
sperm nucleus,
matin network
equal
is
fibers, in
the
In Figure
22,
pull apart
Parthenogenesis
9.
Fertilization
is
by no means an
the formation of a
new
essential process in
27
thenogenetically.
and
filial
Moreover,
generation.
it
is
the
07ily
bridge.
In the majority of animals the egg develops enand independent of the mother,
thus limiting to the egg-cell itself all possible materAlthough there
nal contributions to the offspring.
tirely outside of
is
abundant evidence that half of the filial charcome from the male parent, the only
acteristics
new
individual
is
even
is
fertilization.
The
entire
factor
of
heritage
in
all
probability,
is
re-
into the
GENETICS
28
variably
individual developing
cells
To
that in-
new
the
fertilized
size as
and, furthermore,
when one
make
ways
in
ever they
may
be,
what-
of the heritage,
more conven-
It
is
im-
portant at present, however, to emphasize the obvious conclusion that determiners of heredity
must
What
Do
::
entities,
basis
is
heredity
for
29
as
German embryologist
the
"It
is
by the
aid of microscopes
or are they
it.
It
is
something
which controls the unfolding of the developing organism with respect to both quantity and quality,
and which also governs the time and rate of appearance of
its
"The mechanism
of heredity
is
the mechanism of
differentiation."
12.
Certain investigators,
who
of
the
seek a morphological
chromosomes
as the car-
few
of the
indicated below.
First:
GENETICS
30
size
between the egg-cell and the sperm-cell, in heredtwo are practically equivalent, as has been
ity the
repeatedly shown by making reciprocal crosses between the two sexes. The only features that are
apparently alike in both the germ-cells are the
chromosomes.
The
inference
is,
The
existence of an extra
chromosome
out
later,
is,
in probable
as will be pointed
chromosome material
of the
is
during
when two
or
more
nucleus.
cially
sperm-cells, instead of
urchins' eggs.
fertilized
31
The
fact that
chromosomes
in the
may
be predicted,
and, conversely, that from the appearance of sexually
mature hybrids the character of certain chromosomes
in their germ-cells may be predicted.
Again, the correlation of a particular chromosome
in the germ-cells with a definite adult character,
namely sex, has been repeatedly demonstrated in
connection with the so-called "extra chromosome"
to which reference has already been made.
Fifth: Finally, excellent evidence of a definite
causal connection between certain chromosomes of
the germ-cells and particular somatic characters has
characters in the adult hybrids
GENETICS
32
experiments upon
Boveri found that he was
the eggs of sea-urchins.
able in some instances to shake out the nuclei bodily,
chromosomes and all, from the mature eggs of the
sea-urchin, Splicer echinus,
critical
us
the
carriers of
heredity.
Several
biologists,
however,
raise
protesting
an objecting
against the
mo-
In
sexual
amount
reproduction,
moreover,
some small
form
33
may have
It
lem
is
not unlikely that the key to this whole probbe furnished by the biochemists and that the
will
final analysis of
will
logical in nature.
possible a million
com-
The
limited
we take
seem to be un-
an astronomer dealing
GENETICS
34
may
and maintain
As
said before,
it is
various enzymes initiate, retard, or accelerate successive chemical combinations occurring in the pro-
known
as heredity.
of cell activity
makes
it
possible for
is
indicated
by
him
"The
to
es-
cytological
stances play
some
is,
life.
that
it is
is
The
tain
35
Conclusion
that
of
it
and
I think
some
of the
of genetics."
CHAPTER
III
VARIATION
1.
Nature
in
The
is
like
them.
It
is
On
it
difficulties
is
in
common
and animals
getting or-
VARIATION
37
of
variation
is
2.
Much
of genetics.
it
many minor
differences.
A flock of sheep may all look alike to a passing stranger, but not to the man who tends them.
A dozen
blue violet plants from
The key
is
is
much more
commonly
wide-
believed.
by Dr.
GENETICS
38
whom
he strives to
brief
Kinds of Variation
With
may
be
Under
and
size,
number
Many
free,
less fertile
than when
known
to
VARIATION
39
ample
of a single variation or
clovers,
*'
sport."
Four-leaved
for the reason
common, neverthe-
occurs frequently.
less
may be
harmful to the organism possessing them.
Useful variations are of the kind emphasized by Darwin as being effectively made use of in
natural selection. Indifferent variations, on the
other hand, are those which apparently do not play
an important part in the welfare of their possessor,
as, for example, the color of the eyes or of the hair.
Finally, the degree of degeneration in certain organs
may be cited as an illustration of harmful variations.
The amount of closure of the opening from the intestine into the vermiform appendix in man is an example of a harmful variation, since the larger the
With
c.
useful, indifferent, or
tions
With
is
(orthogenetic) or indefinite
(fortuitous)
"A
Peculiar Legless
Lamb."
Stockard.
Biol. Bull,
xiii,
p. 288.
GENETICS
40
for
him
makes
it
easy
that
furnish
directions
the
repertory
of
opportunity,
With
e.
somatic or germinal.
arise as modifications
They
Somatic, or
body
due to environmental
may be
variations,
may
factors.
be quite
may
without regard to the environment, are deepseated, and of racial rather than of individual sigarise
nificance.
With
/.
fall
may
normal, or they
tations
tions
With
may form
a continuous
series,
tinuous
in
character.
An
VARIATION
41
encyclopedic volume
"On
by Bateson
in his
of Variation."
h.
With
quantitative
or
qualitative.
six-rayed
may be
starfish
number
With
an average stand-
may
all
from the
center about a new
parental standard and consequently come under the
head of mutations,
j. Finally, and most important in the present
average quite distinct
to heritability,
variations
may
GENETICS
42
4.
Roughly
ways of studying
variations first, Darwin's method of observation
and the description of more or less isolated cases
second, Galton's biometric method of statistical
inquir}^
and third, Mendel's experimental method.
The second of these methods will be considered in
stated, there are three
this chapter.
5.
The new
Biometry
is,
the applica-
school of biometricians.
may
which
individual case,
VARIATION
offer
43
generalizations.
6.
Fluctuating Variation
are
dition.
According to the mathematical laws of
chance these deviations sometimes are plus and
sometimes minus, and consequently they may be
termed fluctuating variations.
Pearson gives as a simple illustration of fluctuating
variation the
beech-leaves,
Number of Ribs
trees,
GENETICS
44
either tree
first
is
number
instance the
Number
of ribs fluctuates
In the
around
oi
Individuals
UsTor
35 p.
Arithmetical
Constants
MeanCA.M)=49
Mode CM) -5
30
CoeKicient
oi
(o-)
= .7e4
VariabilitY (C.V.)=
1^1
Formulae
zo
S(x.f)
A.D. =
cr
rzcx^.f)
=T
15
c.\/
A.M.
2 = sum
x=
from A.M.
W
f
Number
Fig. 25.
number
total
of
in the class
number
Rays
starfish rays.
From data by
Goldschmidt.
eighteen as the
around
fifteen.
commonest kind
in the
second case,
easily
VARIATION
45
all
of
which
is
number
4.915, a
number
starfishes selected at
random.
is
namely,
polygon.
is .52.
In other words,
.52
is
the average
(5
4.915) of a ray,
GENETICS
46
and consequently the average of the entire fortyseven amounts to .52 of a ray. In another collection of starfishes where either more seven-rayed or
two-rayed specimens might be present, the average
deviation would probably be greater.
By computing the average deviation, therefore,
and using it as the criterion of variation, a comparison of the variability of organisms that have been
taken from different localities or subjected to different conditions can be definitely expressed.
measure
of variability
by biometricians,
is more accurate,
more commonly
in use
it
This
is
is
sum
of all
the deviations
in
the arithmetical
in
mean
n, the total
number
/,
2,
of
number of individuals
the sum of the classes and
the
of individuals.^
VARIATION
mean
47
making up the
pol^^gon, the
it.
7.
a.
The
Relative Variability
ability of
two
lots of
Number
Woods
Hole
A.M.
West Shore
1,001
58.928
2.339
Penzance Point
1,002
61.718
2.737
Nobska Point
Nobska Point
Nobska Point
Barnacle Beach
Big Wepecket
Mid-Wepecket
1,002
61.737
2.152
1,001
61.944
2.234
496
66.944
2.366
998
63.932
2.604
1,006
57.426
2.052
500
57.606
2.098
.0352
.0412
.0324
.0337
.0507
.0393
.0308
.0447
61.066
2.335
.0386
1,008
59.051
3.023
520
60.892
3.361
.0454
.0703
59.664
3.138
.0538
Average
Cali-
fornia
f
'[
for
Mass
Belmont Beds
San Francisco Bay
Average for Cal.
Difference
The obvious
table
Probable
OF Shells
Locality
is
Error
.803
GENETICS
48
new
more variation
in
Bimodal Curves
Sometimes two conspicuous modes make their appearance in a frequency polygon, as Jennings found,
f|Jonibr
of
individuals
25
VARIATION
49
this
Paramecium
species,
aurelia
in each instance.
q\
leaves
\z
w
;o
9
8
7
6
5
4
3
2
1
Number ot
ribs
15
16
17
18
19
10
EI
The ribs of leaves from two beech trees. When put together
they form a polygon which does not reveal its double origin. From
data by Pearson.
Fig. 27.
is
more
which fluctuates
GENETICS
50
c.
Shew Polygons
The
may
Among
number
of ray
florets varies
themselves,
fluctuates
VARIATION
51
85
80
15
70
65
60
55
50
45
40
35
30
25
20
;5
10
Kv!''
12
13
Fig, 28.
.
15
16
17
18
Variation
19
EO
Z1
It
23
24
25
26
27
28
29
30
31
32
33
34
JS
36
37
38
39
mum leucanthemum)
in
this
particular
instance
of the
to
invasion
colony.
resulting
GENETICS
52
8.
is
It is
11 mm., 11 to 12 mm., 12 to 13 mm., etc.
somewhat more difficult to classify variations when
color or pattern
is
it
9.
With
An axiom
is
self-evident, requiring
as
axiomatic.
no explanation.
VARIATION
The absence
53
than
from this point of
view, the phenomenon requiring an explanation.
Although Darwin himself spent some time in pointthe occurrence of variations
is,
accepted
it
as a
it
its
causes.
b.
Lamarck and
his followers
refer-
is,
of
efforts
M,
Af.
oO
-L.
-L.
50
35
<
'^0
45
Ratio
50
of
70
75
80
85
60
65
55
height of head to length of shell
90
95
100
o/
/o
>
cording to the Lamarckian school, in the "environment" and "training" sides of the triangle of life
rather than in the "heritage" side (Fig. l)c
For example, Woltereck, by controlling the single
GENETICS
54
manner indicated
Number
in
When
Figure 29.
poor food
of
Flowers
Number
cf
oUaens
L
lo
10
10
Fig. 30.
Variations in the number of stamens in the flowers of the " livefor-ever" (Sedum spectabile) under various controlled conditions.
For detailed description, see text. After Klebs.
of the
head height to
it
Sedum
spectabile,
are
VARIATION
55
that
is
it is
a device
by the mingling
By
phimixis).
of
two
germplasm {am-
strains of
number
of observa-
Tower found among his potato-beetles, for examthat two strains reared in the same environment
ple,
showed
differences
striking
necessarily due to
Similar cases
factors.
d.
Lastly,
for the
Study
in
variation,
intrinsic rather
Bateson,
may
fact
than to extrinsic
whose work
"On
it
is
is
Materials
classic,
rather futile
He
attempts by saying
my
judgment, premature."
In conclusion, the words of Darwin written half
"Our ignorance of the laws of
a century ago
may still be appropriately
variation is profound"
quoted, notwithstanding the fact that in biometry
we have at least an excellent analytical method by
means of which considerable insight into variation
variation
is,
in
is
being gained.
CHAPTER
IV
MUTATION
1.
Among
fluctuating
variations to
which
Darwin was
tions or
day
in a
to analyze the
nected.
56
MUTATION
2.
51
it is
variation, but
difference
that frequently
it is diflicult
to ascertain
by
inspec-
due to a mutation
comes in breeding, for the
progeny of a fluctuation will vary around the old
average of the parental generation, while the progeny
or a fluctuation.
The
is
test
GENETICS
58
eye.
They
altogether.
To
3.
Freaks
MUTATION
4.
De
59
Kinds of Mutation
according
to
tion of a
acters
new
character to the
making up the
sum
individual.
of
If
complex char-
rumor may be
said to
as heritable mutations
is presented in Davenport's
Heredity in Relation to Eugenics."
Regressive mutations are characterized by the
dropping out of something. Thus albinism is caused
by the absence of pigment or color. Albinic mutants
which breed true are well known, particularly among
mammals, such as rats, mice, rabbits, cats, guineapigs, and even man himself.
Degressive mutations include cases of the return
of a character which was formerly present in the
past history of the race, but which has for generations
been absent or latent. Castle's four-toed race of
guinea-pigs furnishes an example of this class of
mutations. In 1906 Professor Castle discovered a
newly born guinea-pig in one of his pens with four
toes on each hind foot, from which he has successfully
*'
The
hypothetical an-
GENETICS
60
foot,
5.
The
definition of species.
What
Since
of the divine
it
law
to be annihilated!
it
deserved
for
MUTATION
61
attempt to bring order out of chaos by classifying the living world, and it served its purpose well
until Darwin's idea of half a century ago, that the
origin of all species is from preceding species, put an
entirely new face upon the whole matter.
Organisms of different species were found to be related to
one another, and even man could no longer escape
acknowledging his poor animal relations. As a
cessful
consequence, likenesses rather than differences thereafter claimed the most attention.
made common
had hitherto
it
was shown
may
of
individuals
arbitrarily
called
species.
The
upon analysis
dif-
GENETICS
62
fering
Fig. 31.
Diagram to illustrate various ideas about "species." Under
Species A are represented two groups of individuals which are near
enough alike to be placed within a single species, but which are sufficiently unlike each other to constitute the " sub-species " or "varieties " of Darwin.
Under Species B are various groups of individuals
distinguished from each other by the addition or loss of one or more
characters. These groups represent the "elementary species" and
"varieties" of de Vries. 'The "barrier of Linnaeus" attempted to
separate species absolutely from each other. Darwin sought to find
loopholes in this barrier. To-day attention is directed rather to the
relation between individuals than to the boundaries between species.
almost
An
have become
limitless.
progressive mutation
differing
is
MUTATION
by the addition
63
muta-
6.
The
known
oldest
mutation
is
in
Nature
*'
fringed
made
its
appearance in the garden of the Heidelberg apothecary Sprengel in 1590 among plants of the "greater
The fringed celcelandine," Chelidonium majus.
andine bred true at once and is now a widespread
and well-known
species.
always given
rise
it
has
to a constant progeny.
found
in English cornfields.
Instances are
known
of
which
in turn
of
GENETICS
64
7.
progeny
It
is
plant
of
believed
though
by botanists
is
it is
now, so far as
is
knowm, extinct
al-
as a
or
mutants, 0.
brevistylis,
styled flowers,
and
These
tw^o
leaves.
unknown
characterized
by shortsmooth
Paris, Leyden,
and the
Kew
Gardens.
as
de Vries
felt
by
safe in regard-
MUTATION
65
Amsterdam, where
in
He
Generation
GENETICS
66
lamarckiana.
Finally, scintillans
cause
it
only in
its
inconstancy be-
The 1895
plant gave 53
cent
lata.
One
lamarckiana,
39 per
cent
and
per
scintillans,
1
per cent
scintillans, 21
MUTATION
67
essential points.
The
ana
is
as unmistakable
it is
in
Holland.
A parallel
rise to
mutability of 0. lamarcki-
and
as diverse in
America as
mutations
is
is
probably
Mutations are doubtless much more comthan has been generally supposed, and it is likely
that they will receive more attention in the future
than they have in the past. De Vries rather pointedly
says: "The theory of mutations is a starting-point
selection.
mon
slow changes has held back science from such investigations during half a century."
8.
GENETICS
68
long, sagging
back and
somewhat
German
much
be
bow-
it
of a
Later on this breed gave place in public favor to another mutant, the Merino, which produces a superior
grade of wool.
Hornless cattle suffer fewer injuries from one another than horned cattle.
racy^
Hereford
well-established race
a bovine aristocwith registry books and blue blood all their own.
of polled
cattle, constituting
in
cattle,
Fowl."
in
Domestic
MUTATION
69
a toe
with
out oil-gland or
tail
with one
with two pairs of spurs withwith neck devoid of feathers
five, six,
shaft
with
barbs
twisted
with feather
and
dichoto-
the
first
and
refused to blend
Bateson (1894), in
his
dissimilar form."
continuous variations"
among
animals,
many of which
al-
in the non-inherit-
is
GENETICS
70
an
ably of
primroses
Evening
display
the
in
Holland or America, in a wild state or under cultiMutations, like poets, are born, not made.
vation.
It has been suggested by Standfuss that species
may go through the same kind of a life-cycle that individuals do, only
'^
taking
infinitely
more time to do
it.
As shown in
Figure
they
32,
and enter
the prodigious
growth period of
infancy and youth,
both of which are
Fig. 32.
Diagram
characterized
much
by
fluctuation.
of old age,
it
approaches in
and end
many features
is
in death or extinction.
edly illustrated
no
recall,
although
by phylogenetic
This cycle
is
repeat-
lines of fossil
forms
MUTATION
71
an excessive number
fantastic shapes,
spines
of
the
ammonites,
gigantic size
as
theromorphs.
cycle in
and
belemnites,
in
trilobites,
or
among
or
of
The
when mutants
an individual, may extend over a considerable period of the whole cycle,
or it may be confined to a relatively small segment.
reproductive period of a species
It
is
off,
as of
may have
period of species-reproduction.
When
it is
with
this
in
little,
has
sible sources of
mutation
is
that mutations
may
be
As a matter
of fact, Spron-
gel's
GENETICS
72
however,
what
is
made
clear.
is
thus produced.
is
A Summary
10.
new
The main
may
be indicated as follows
a. New species arise abruptly regardless of environment without transitional forms, and at present they
Vries
known
New
though
it
may
repeatedly give
tions.
c.
is,
New
mutations
are,
from the
like.
first,
constant, that
by the addition
elementary
species,
of
type,
something new,
as well as forms
many
/.
arise simultaneously in
''
sport."
mean with
is
the case
MUTATION
73
Mutations
may
occur in
all
directions, that
is,
species.
j.
must
sift
The bearing
heredity
new
of the
lies in
belief, it is
that
out
species.
make up
heritable variations.
If this supposition
proves to be true, mutations furnish the essential
in the study of heredity.
Consequently,
whatever knowledge we may gain of them has a direct
relation to the entire problem of genetics.
material
CHAPTER V
THE INHERITANCE OF ACQUIRED CHARACTERS
Summary of Preceding Chapters
1.
Hereditary resemblance
offspring
plasm.
Perfect inheritance, or uniformity of generations,
does not
It
is
of evolutionary advance.
Some
character and
statistical
discontinuous
differences
may
variations,
representing
qualitative
statistical analysis.
74
The Bearing of
this
in succeeding
Hence
Can
75
it
be-
of varia-
of
Can
any way
changes wrought
somatoplasm be so impressed upon the germplasm as to change it in such a way that it, in turn,
will give rise to similarly modified somatoplasm in the
Can nurture as well as nature be
next generation ?
in the
transmitted
In answering these questions we are of course concerned solely with biological inheritance and not at
all with those extra-biological accumulations in the
and the
and which make
way
like
which constitute
civilization
upon us much
3.
in the
76
GENETICS
several viewpoints.
pound
of cure
who have at
human conditions
pists,
for sociologists
for educators,
may
and philanthro-
who
cherish hopes
mind
sitory;
religious workers,
for
strengthened
that
conquests
who want
in
their faith
character-building
may
may
life,
to
for
know
all
these and
characters be inherited
4.
many
more,
An
it is
:
important
Can
acquired
nature
is
Lamarck with
respect to
77
how such
first
to ex-
but the
man
who,
in a
masterly manner,
who
fo-
professor
in
Weismann made
Baden.
who
Darwinians
deny, such
inheritance.
If
the
question could be decided by a vote or by an expression of opinion, the result would be doubtful, since
ing the
inherited.
But
just
The
Confusion in Definitions
source of
much
of the lack of
agreement
in
what con-
stitutes
of
GENETICS
78
it was red,
was black. So they hacked away
at each other, as all good knights should do in the
defense of the truth, until they both fell down dead
beside the shield which was black on one side and
red on the other.
Of course actual characters are never inherited, but
only the determiners or potentialities which regulate
the way in which the organism reacts to its environ-
ment
it
It
is
obvious that
dis-
cussion
accepted.
6.
Weismann
defines
germplasm.
its
origin in the
79
Examples of acquired characters in the Weismannian sense are mutilations, the effects of environment, the results of function as in the use or disuse of
certain organs, and such diseases as may be due either
to invading bacteria or to the neglect or abuse of
the bodily mechanism.
7.
vidual
"
make up
familiar lines
the indi-
"Born"
their
characters which
by quoting the
classi-
characters
origin
in
the
are
constitutional,
germplasm
itself.
having
They
are
illustrated
features.
gies
may have
GENETICS
80
we
them
in this connection,
it
should be noticed
itself,
if
is
an acquired character
of
They
results of environ-
on the part
parental
gonorrhoea
or
tuberculosis
arising
by
from
as
What
their
appearance in later
Variations reappear
of
acquired
characters
are
life.
"WTiat
inherited.^"
it
81
have
individual
arises,
and
germplasm.
Any
character that
is
is
no transfer
of characters
between
Thus
it is
Such temporary
9.
appearance
is
acquisi-
do not
in-
What may
The
bridization.
GENETICS
82
characters in amphimixis and hybridization will receive further attention in a later chapter.
The
tions, in
may show a
of course occur,
than sexually
produced generations, indicates that amphimixis in
itself is by no means sufficient to account for all kinds
larger degree of variability
of variations.
The assumption
of
It
is
acting
it
admit, simply
an unknown quantity,
consequently doomed to remain so.
is
at present
alternative classes
first,
two
upon
into
somatoplasm
sarily at the
first
true
the
affect the
The
so
that
is
acquired
characters
according
to
in-
our
stimuli
upon the
83
germplasm, regardless of any simultaneous modification of the somatoplasm, must be excluded as irrelevant to a discussion of the heritability of acquired
Weismannian
characters in the
Many
instances
of
direct
all.
influence
of
external
MacDougall,
for example,
in-
duced character.
Sitkowski,
also, fed
moth
moths that
and these
in turn
of,
is
the somatoplasm.
GENETICS
84
10.
Weismann's reasons
may
cells.
does occur
is
plasm
characters.
little
more
closely.
11.
No Known Mechanism
is
more
somatic
cells
more
to return to their
cell
85
complexes too
hard to see
how
It
It
is
tire
all
others,
nately there
is
no tangible basis
all
parental somatic
It
cells
is
take
GENETICS
86
new
individual, hence
cell
structure
favors
this
hypothesis, while
many
the germ-cells.
We
must
Morgan
acknowledge,
however,
with
Lloyd
12.
The evidence
for
the
inheritance
of
acquired
to go around the
it
does not.
began to
sift
much
like the
influences
h.
Environmental
effects;
It
falls chiefly
a.
c.
of warts
toads.
quired characters
d.
tliat
fact,
by handling
87
is
Mutilations
classes have
been mutilated into deformity by bandaging, without the mutilation in any way becoming
an inherited character. The same result is also
true of circumcision, a mutilation practised from
ancient times by the Jews and certain other Eastern
for centuries
peoples.
The
man
GENETICS
88
same
point.
What Weismann
tails
off
this decaudalization
One may
new-born mice.
see in the
away on shelves,
catacombs of
filed carefully
as a "
by saying: "Wooden
b.
Trees
legs are
not in-
like
the
are
not
when
known
their
to
produce a modified
adverse environmental
progeny
conditions
are
removed. Similarly, the persistent sunburn of Englishmen long resident in India does not reappear in
their children born in England.
89
and
feet
grew noticeably
larger than in
mammals which
an authentic instance
quired characters,
of the Alps
is it
in the course
GENETICS
90
warm
valleys
at a lower altitude.
amount
species.
More
recently
Kammerer, by
similarly reducing
mandra maculosa, a salamander that normally produces about seventy eggs which, when hatched in
water, become gill-breathing tadpoles, into a salamander producing only two to seven young which
are born alive without gills and are able to live out
91
on
c.
The
generations.
It has
GENETICS
92
mud
generation just as
and
write.
Many
Disease Transmission
have
their im-
pathogenic
bacteria.
invading
mediate
cause
Bacteria
reason that
it is
in
for
the
93
that is, a
lack of resistance to bacterial invasion due to defectiveness in physical or physiological equipment,
may
disease, may "acquire" a generally weakened germplasm and so produce a progeny exhibiting general
liability to disease
but it is doubtful if such a condition can properly be termed the inheritance of
an acquired character, since the particular definite
disease in question is not demonstrably heritable.
When alcoholism "runs in a family," its reappearance in the son is probably due to the fact that he
is derived from the same weak strain of germplasm
as his father.
The fact that the father succumbed
;
is
is
it is
may
some cases
in
is
arise
through
germplasm has
less resistance to
ceeding generation.
more
likely to
The
succumb
alcoholism in a suc-
to the disease.
This, how-
GENETICS
94
ever,
is
When a man of the present generation has rheumatic gout, it is a severe stretch both of patriotism
and of the powers of heredity to trace the origin of
the affliction back to a revolutionary ancestor
acquired
sciatic
many
who
is
quite as direct as
sufficient to ac-
Weismann
chapter,
is
at least difficult,
if
not
To prove
according to
firsts
Weismann
95
particular
third,
the same
succeeding
The Opposition
The opponents
of
to Weismann
Weismann
point out, as a
weak
There
is
GENETICS
96
affect the
as, for
Conclusion
is,
that the somatoplasm affects the germ-cells, the inheritance of acquired characters
is
by no means
thereby established.
In order to do
in question,
this,
its
influence
a general way,
which
is
It will
characters.
CHAPTER
VI
of Attack
of facts to a science,
it
is
made
up.
Chemistry was alchemy until the chemical elements were identified and isolated. Histology was
terra ohscura until the cell theory brought forward
"cells" as the units of tissues. In the same way
there could be no science of genetics until the conception was developed that the individual is a bundle
of unit characters rather than a unit in itself.
So it
now
speak of inheritance as
has come about that we
applied to unit characters rather than to individuals
as a whole.
bination of
many
units
makes
it
is
a com-
sible
Thus
97
"
GENETICS
98
fourth unit
is
more
six
that
is,
many unit
as
made up
are
many
which the
of
very
new
addition of one
2.
Galton's
Law
of Regression
and
offspring
by means
of treating statistically a
What Galton
human
stature as a
human
stature
is
normally
less
is
is,
than male
1.00
"midparent"
99
in
each
formula
$
1.08
Inches
The
results
inches are
of
shown
in
measurements expressed in
the following table, in which the
his
100
GENETICS
respective midparents.
Midparental
101
GENETICS
102
It
means
of
In his mind arose the question whether it would not be possible by continually
breeding from plus parents, granting that plus parcontrolling heredity.
for
To
group of plants
measurable.
since
is self-fertilizing, prolific,
Somewhat
Phaseolus,
and
this
easily
much
as expected.
That
beans little offspring, according to the expectation, although they each produced offspring
nor
little
manner
of fluctuating variability
line.
In this way
of
of the
the
known
nineteen
"pure
Johanssen's
lines," a further
some
103
original
most
recent years.
4.
pure
pure
line
in
line is
question.
That
is,
selection icithin
question.
This
is
illustrated in Figure 34 in
which the
is
shown
results
for four
GENETICS
104
beans weighing 60
VII, and
XV,
eg.
lines IT,
Cent,-
Weight of
parent eed
10 20 30 40
Pure
line
60 70
number
10
20 30 10 50 60 70
10
20 SO 40 50 60 70
10
20 }0 40 SO 60 70
;0
20
50 40
SO SO 70
The result of selection in four pure lines of beans. The vertical columns, representing the average progeny from different sized
Fig. 34.
all derived from the same pure line, contain groups nearer
aUke than the horizontal columns, representing progeny from the
same sized parents, but different pure lines. All the numbers indicate
centigrams. Data from Johanssen.
parents
vertical
fluctuating parent.
As a
TTa'rvitqt' Vitat?
105
shown
in
GENETICS
106
TO
02
a;
03
OS
>
o
HI
a
CO
O
^-5
a^
O.
-H
oo
o o S
Q,
lO
a "^M
W
1-1
t.
!<
05
>J
CI
03
(?
\(5>\ c
CI
O O
>>
'o
a
o
'O
I
"a;
'oj
10
OB
C3
ff3
a
a
o
q
^ o:
O
^ os
3 ^
2 a c3
O
a
a;
6^
03
^^
l6>
bO
pH
a
03
4^
o
W)
c3
;h
o
>
to
03
"5
03
CO 7^
o
5h
03
so far as inheritance
is
107
concerned, and
it
makes no
from which
is a
with
no
mediocrity
or
"intype
complete return to
As a
heritance" of the parental modification.
far-reaching importance
played
in
by natural
heredity,
5.
*'the
it is
may
be
obtained.
First,
when two
similar
organisms identical in
truly as
in a single in-
arising
GENETICS
108
changes to the sexual method of reproduction, segregation of characters may occur and different
combinations
6.
As an
result.
Tower's Potato-beetles
Among
the
numerous
cultures
of
this
Fig, 36.
Diagram showing the ineffectiveness of selection through twelve
generations within a homozygous strain in the case of the Colorado
potato-beetle (Leptinotarsa). In each generation extreme dark specimens were selected as the parents of the succeeding generation but the
progeny always swung back to the type. After Tower.
GENETICS
no
way
Thus
in a
pure
line
of
two
An
Jennings'
effect in
Work on Paramecium
furnished
206
zoo
194
176
142
US
100
The
actual
mean
length of
Magni-
Fig. 37.
is
without
effect.
differ
from each
size, structure,
of niultiphcation
111
With
was able to
average
drawn to
size,
scale, is
mean length he
pure
shown
lines,
MICRA
256 <
whose
in Figure 37.
>
Fig. 38.
Diagram of a single race (D) showing the variation in the size
Magnified about 230 diameters. After Jennings.
of the individuals.
arranged in a
are
series.
shown in Figure
Each horizontal row represents a
bred by Jennings
is
micron
is
pure lines
39.
single race or
jo^o^th of a millimeter.
GENETICS
112
pure
line,
sign
The mean
is
indicated
lot,
as
by the
shown
55
DOOHa
UUUU
aO'OHHa
8()898eH08M
gyflHH
98
gyjHMM 89
MM
Fig. 39.
Diagram
different races
of the species
shown
45
in Figure 37.
The individual showing the mean size in each race is indicated by a cross placed above it. The mean for the entire lot is at
the horizontal line. The magnification is about 24 diameters. After
Jennings.
single race.
by the
vertical line,
is
155 micra.
of individuals belonging to
The
each size
line their
is
total
number
not indicated,
number
is
more
113
From
the foregoing
it
will
be determined by an inspection of
its
somatic char-
acters alone.
alike, regardless of
to be identical phenotypically
by Johanssen
or to belong to the
same phenotype.
On
same
genotype.
GENETICS
114
it
in a different sense.
Natural history and common usage deal principally with phenotypes, that is, with organisms as
they appear. The older theories of heredity were
likewise concerned wdth phenotypes, but we are now
coming to see more clearly than before that heredity
must always be a case of similarity in origin, that is,
in germinal composition, and that similarity in appearance by no means always indicates similarity
in origin or true relationship.
The assumption
many
made
conclusions in comparative
the foundation
anatomy and
who
upon
115
of view to
The
9.
mixture of pure
lines
tion.
line,
when they
factor of environ-
permanent hereditary
differences
due to germinal
which the
composed.
Johanssen has made the distinction between pure
population
is
lines
(Fig. 40), in
The beans
test tubes.
one
Tubes
size.
in
any
vertically
Thus
bean
in
it
is
one
inverted
may
size.
be a rare
size of
GENETICS
116
tube of
jpure line 3,
monest
size in
may
another
The
as jpure line 2.
pure
five
Pure: Line
in
40
are
Figure
combined
ji
-^
in
population at the
n
^ igji^^ i
lines
represented
^^
bottom
of
figure^
making
-j^jji
the
phenotype
phenotypes
five
five
geno-
In the
population, howtypes.
ever,
the
five
are
genotypes
within
hidden
one phenotype.
Hence, while
selection within
no hereditary
Population
fluence,
HI
Diagrams
Fig.
40.
it is
dent that
lines
The
beans of each pure line are represented as assorted into inverted test tubes making a curve
Test tubes containof fluctuating variability.
ing beans of the same weight are placed in the
same vertical row. After Johanssen.
in-
evi-
selec-
tion within a
population
may
shift
or
move
over
the
type
of
the progeny
117
by
but
upon a
partial
population.
From
lations
this distinction
it
is
clear
between pure
why
lines
and popu-
on
tain standard.
As soon
a "reversion to type"
there
is
line ceases.
in dealing
however, a pure
bers of
it,
line
is
all
the
mem-
GENETICS
118
10.
From
means
it
appears that by
normally in nature,
it is
to
words,
strengthen the frontier posts that are already esIt cannot break into the wilderness and
tablished.
create
new
centers.
members
of a pure line,
having
tend to
critical step
indispensable
species,
is
How
can the
the
ever occur
evolutionary
That
it
derivation
of
Ameba
to man.
At present the only loophole
of escape
seems to
lie
acters, or in
is
119
way
by natural
selection,
line,
of
CHAPTER
VII
Modern
first,
by microscopical
plants.
The
first
two
of these
methods
of
approach have
In the
present chapter attention will be directed to a consideration of experimental breeding with reference
is, breeding from unlike parwhich Jennings characterizes by the
expressive phrase, "the melting-pot of cross-breedmg.
to hybridization, that
ents, a process
2.
The Melting-pot
of Cross-breeding
Hybridization, or cross-breeding,
by Galton
as
formulated
120
or par-
these,
blending inheritance
may
121
be called the
from
Galton
parent.
inheritance of
process
by
the
stature in which a
tall
and
illustrated
human
this
By
Chapter IX.
rental contributions
In inheritance of human eye-color, for example, the offspring usually have eyes colored like
offspring.
is
eye-
when the
GENETICS
122
Alternative
Particulate
Characteristics
of parental
jermplasm as
Shovun
in
the
Somaplasm
Double germplasm
termed from
Contributions
from both parents
Possible kind*
of apparent
dfsprinj this
jermplasm
maif produce
Fig. 41.
123
Vienna.
At the time
results
appeared
in 18G6).
in Briinn.
GENETICS
124
of Charles
Darwin.
sight.
What Mendel
125
normally
self-fertilized
flowers
it
is
it
from
ripe pollen
another flower.
Mendel's
cross-breeding
experiments
on
peas
formulated as follows
When parents that are unlike with respect to any
character are crossed, the progeny of the first gen:
The
manner
is
its
off-
When,
An
make
plain the
man-
artificially crossed
of a tall variety
with those of a dwarf variety,
parent.
It
made no
difference
tall
like
the
first
GENETICS
126
When
these
tall
sex.
quently crossed with each other, or allowed to produce offspring by self-fertilization which amounts
to the same thing, 787 plants of the tall variety and
277 of the dwarf kind were obtained, making approximately the proportion of 3 to
On
1.
Tall, r,
That
dwarf, U
tall, T(t).
tallness
When now
follows
the hybrids,
T(t),
gether,
:
-{- 1
(all
T-\-t (a ll possible
TT+
Tt
Tt
tt
TT+2T(t)+tt
sperm characters)
was
as
127
T (t).
The same thing may
character latent,
by the checkerboard
plan.
Male Gametes
42).
Each square
to
^.T
z
<
la
The
<
possible
offspring.
of the parents
is
shown
while
the
arrows repre-
Fig. 42.
Diagram to illustrate
theoretically the formation of the
four possible zygotes in the second
filial generation of a monohybrid.
their heredi-
tary composition.
The
this:
essential
hereditary
upon
usually
is
briefly
independent
crossing, regardless of
temporary dominance.
Mendel
He obtained
practically
GENETICS
128
1.
Character
1^29
in a year,
while zoologists
rabbits
of
progeny
is
much
GENETICS
130
phenotypic characters
is
technical difficulty.
What
the
modern experimenter
an organism,
first,
in genetics desires
is
6.
The
was the
The
essential
Principle of Segregation
thing which
Mendel demonstrated
may
common
stream of germplasm
from which, in subsequent generations, they may
segregate out apparently unmodified by having been
intimately associated with each other. This "law of
sources
unite in a
It
1
'^
"Mendel's Principles
of Heredity," 1909.
" Einf iihrung in die experimentelle Vererbungslehre," 1911.
the
of
131
separate
blossoms.
The
all
general
that
character,
single
monohybrids,
is,
is
given in
Figure 43.
K (Recesiln)
(Dominant)
D(R)
1
DD
DD
2D(R)
D(R)
RR
i
OD
RR
RR
^
'
Fig.
uf
43. General Mendelian formula for a monohybrid.
7.
DD
character which
is
it
said
character
is
In contrast to the homozygous condition, an organism is said to be heterozygous when it derives the
determiner of a character from one parent only.
Such an organism is described as a heterozygote with
respect to the character in question.
may
A homozygous
appear
alike.
GENETICS
132
that
so,
is,
they
may have
The
always
is
different.
Identification of a Heterozygote
although
pointed out
it
may
later.
not always
is
it
to identify
and observing
both look
character, T,
is
example,
T{t), for
alike, since
sufficient to
produce complete
of
tall
tall
tallness.
peas are
tt,
Case
Case
That
(Case
r
T
I.
II.
is, if
I),
+rX +
+ tXt +
<
gous (Case
+ 50 per cent
if
is
ti.
homozygous
II),
is
heterozy-
is
= oOper
character, but
9.
will
in question.
133
or
is
not, present.
When
it is
When
it
it
follows that
When
it will
it
not
all,
it
is
DiHYBRIDS
been made exclusively to monohybrids, any two of which are supposed to be similar
except with respect to a single unit character. ]\Ionohybrids are comparatively simple, but when two
So
GENETICS
134
of dihybrids
by
crossing
smooth-yellow
crossed,
all
the
SY
and wrinkled-green
offspring
are
WG
are
but
smooth-yellow,
may
and wrinkled-green
WG, which
SY
union
will
be
number
sixteen
of zygotes
(4x4 =
16).
formed by
That
is,
their
the
is
squared, (3
of 3 to 1 in tlihybrid
+ 1)- =
135
com-
16.
the
number
The
1)-.
ofispring ol)tained
of chance,
come to
fall-
The
sixteen
possible
zygotes
resulting
from
which in turn, as
will be demonstrated
phenotypic and nine
SG+
WY+
SY+
WG = all the possible egg gametes
SY+
SG+
WY+
WG = all the possible sperm gametes
SGSG+ SGWY+ SGSY+ SGWG
SGWY
+SYSY+ SYWG
+ WYSY
SGSY
SGWG
WYWG
+ SYWG+WGWG
alike
cross.
WYWG+SYSY+WGWG
different combinations
By
acters in parentheses,
is
any dihybrid
dominant
are
:
GENETICS
136
may
genotypes
follows
Phenotypes
137
in
Each Class
GENETICS
138
is
is
with respect to
human
of this factor,
hair.
HA/f^
'
r;^
T^'
^/
KEY TO Symbols
= Dark
O =
=
=
J
Li^ht
CurlLj
Straight
'^s DARK
Fig. 45.
B. and
The heredity of human hair according to data by
G. C. Davenport. The arcs represent the somatoplasms of four individuals. Within the arcs are the gametes formed by these individuals.
The dominant character is placed on the outside of the arc where it
will
be
visible.
139
The dark
all
compo-
dark
phenotypes
dark
curly,
light
Nu.-nber
in
6AcK
or
light
Nuii<br
straight,
curly
Phenotype:
GENOTrPE
DdrA
cur/ij
notypes
in
present
nine
will
different
Ddrk
combina-
genotypic
shown
sible cases, as
Light cur/ij
in
Figure 46.
Figure
strdight
45
further-
;6
le
Fig. 46.
sible
tion
of germ-cells
third,
Light strdight
binations resulting
when two
hetero-
the
The
cells of
GENETICS
140
making up the
arcs within
which
number
soma-
tic cells.
or
these are the characters that are visible or phenotypic, while the non-apparent recessives are placed
on the
is
Cross-breds of the
have
first
generation
will, therefore,
interbred
may form
and when
which corresponds to a
different zygotes,
8)
?i
RSP
RsP
RSb
Rsp
141
SP
i
l'
rsP
i
Sp
I
(3
1)^
eight
in
p
I
RSP
RSP
RSP-
RsP-
RSp-
Rsp-
rSP-
rsPr
Sp
sp
Fig. 47.
Diagram showing the possible combinations in a guinea-pig
trihybrid of the F2 generation. R, rosetted coat r, non-rosetted coat
(absence of R) S, short hair s, angora hair (absence of S) P, pigmented p, albino (absence of pigment). The eight possible triple
gametes of each parent are placed in the upper and left hand margins.
Each of the sixty-four squares represents a possible zygote or fertilized egg, having received a triple gamete from each parent.
;
different
phenotypes
and
twenty-seven
different
genotypes.
The
its
resulting combinations
work on guinea-pigs
which confirms the Mendelian hypothesis on an
is
well illustrated
by
Castle's
GENETICS
142
Number
in
each class
Genotype
SS
2
64
PP RR
extensive scale.
are represented
by
When
(8),
all
a smooth, or non-rosetted
rosetted
(i?),
143
long-haired
(s),
is
(r),
letters.
short-haired
crossed
with a
rise
to sixty-four
(8
8)
Conclusion
GENETICS
144
hybrids derived from such an ancestry would include 25Q or (3 + 1) ^ possibilities instead of 64 or
(3
1)
^,
differing characters
combined
are
in
When
ten
the parental
kinds of offspring
generation, (3
From
+ 1)
among
^^
1,048,576.
character, a third,
Thus
in
it is
and
so on.
obtained at
13.
Summary
Independent
145
number
become
The
may
visible.
come manifest
in the
may
be intimately associated together in the individual, during the complicated process of maturation
that always precedes the formation of a new individual, separate or segregate out as if independent
of each other and thus are enabled to unite into
new combinations.
CHAPTER
VIII
There
two ways
in
atavism.
Atavism, or "grandparentism,"
may
be defined as
skipping a generation with the result that a particular character in the offspring is
thetical,
ancestor."
146
147
becomes
If
there
clearer
by
illustration.
heterozygous
is
Grandmother
grandfather
HorflOZYGOTE
HOMOZYGOTE
Duplex
Duplex'j
.t*oMOr,
c^;S^^>9
%lu}\^
/
I
Heterozygote
'
Simplex
148
GENETICS
shown diagrammatically
brown-eyed character.
This figure also illustrates what typically occurs in
the formation of Mendelian monohybrids of the first
and second filial generations. The squares are
symbols for the dominant characters, while the circles
When the
are symbols for the recessive characters.
the
circle
recedes
from
superimposed,
are
view,
two
The large outside figures indicate the somatoplasm,
therefore the phenotype.
The
what
it is
The
that deter-
To
149
after a lapse of
2.
many
generations.
False Reversion
selves."
Arrested Developjnent
Vestigial Structures
mouth
of
the
many
GENETICS
150
babies and
mouth
ridges
many
quences to man these and scores of similar characters, which, taken together, make man in the eyes
of the comparative anatomist a veritable old curiosity shop of ancestral relics, are the last traces of
characters which formerly had a significance in some
;
of
these structures
still
reversion.
It
is
reacquisition
life.
Convergent Variation
Erinaceus, an insecti-
mal
all
1.51
no reason, however, for regarding this character as due to descent from a common
spiny ancestor. It is not reversion to an ancestral
type, but rather a case of convergent variation.
Similarity does not always indicate genetic continuity.
In the case of birds albinism, melanism and flavism are modifications of ordinary pigmentation which
appear irregularly among many different species as
pathological " sports," but no one of these conditions
can be regarded as reversions to ancestral white,
spines.
is
Regression
Galton's "law of regression" refers to the w^idespread phenomenon already explained of a constant
swinging back to mediocrity which the breeder must
reversion.
3.
Explanation of Reversion
GENETICS
152
of a
is
now known
is
bination of letters."
4.
make a more
be maintained
153
made by them
in their offspring.
We
by a modified environment,
it is
increasingly difficult
which
will
stock.
That
this
common
is
This
method
is
blind and
is
not
now
fail
full of error.
Its
this
The
procedure
now find
them,
is
who experimented
GENETICS
154
for
among
stedt barley.
Rimpau's method
conditions with a
amount
is
to
minimum
of fertilizer
rather than a
and then to
maximum
select individuals,
where there is little crowding, but from situations where the environmental conditions are ordinary or even unfavorable. Individuals making a
good showing under such usual, or even adverse,
conditions are worthy by nature rather than by nurture and are consequently most desirable as progeniBy this method the attempt is
tors of future stock.
not to keep the progeny of single individuals separate, but to mass together the best as they appear
under ordinary normal environment.
This again is an indirect method of procedure,
although the character of the germplasm is more
nearly hit upon in this way than by Hallet's method,
since the mask of temporary accessory modifications
is stripped so far as possible from the somatoplasm,
and the phenotype made to approximate the genothe
field
typical constitution.
c.
The method
been in part
depends upon the preservation and exploitation of the mutations occurring
in nature.
It recognizes clearly the fact that change
of type is dependent upon a germplasmal variation
of de Vries has already
It
if
155
furnishes to
})reeder
attempting to force
in
nature into producing something new. These mutations, when isolated, may become the progenitors of
desirable
new
lines.
d.
This
is
an
isolation
success-
The
seeds
selection
an improved strain
be maintained.
e.
The method
may
of isolating
pure
lines or
homozygotes
mutations
so, too, in
is
it is
recog-
germplasm.
The method
of
Johanssen
Rimpau
is
GENETICS
156
not sought
zation
is
among
genotypes.
This
is
is
selected.
is
/.
variants
for
It
eliminating
all
combination.
of
of
makes
The
characters themselves
is
produced
of existing
characters.
The
spectacular
"creating"
new
success
plant forms
of
is
Luther Burbank in
due largely to his very
skill in
detecting
among
do not quite
fill
his requirement.
The
in
by
this procedure.
The
agation
It will
is
by asexual prop-
first
two,
is
157
who wish
breeders
It
is
after all
possible
by the determina-
like
recessive,
then
all
that
question,
is
to breed
two
is
the
necessary to
character
in
GENETICS
158
homozygous and,
regardless of
On
if
in other words,
whether
it
is
homozygous or
in
it
graph
of
Chapter
VII.
If,
it
produces
according to what
it
may
be crossed with.
When
dominant
be perpetuated.
on the contrary, the dominant character to be
tested proves to be simplex or heterozygous, as determined by the fact that, when crossed with a recessive, 50 per cent of the progeny are recessive, then
it requires more than a single generation to establish a
homozygous dominant strain.
in question will
If,
population
is
is
if the re-
appear,
composed
159
5.
It
decade
last
compound determiners
may
is
also
control
true,
The
may
The
of heredity.
idea of
be
con-
determiners
For instance,
also produces a
character.
on the belly.
Mendel, whose experiments led him to believe that
each character depends upon only a single determiner for the reason that he worked on characters
lighter color
compound determiners.
These compound factors may be arranged
cases at least, of
rious categories.
(1)
may
is
in va-
be,
added to a
may
appear
(2) A supplementary factor which is added to a
dissimilar factor with the result that a character is
modified in some
(3)
way
cumulative factor
which,
when added
to
GENETICS
160
that a character
(4)
of
An
inhibitory factor,
some other
factor,
and
so on.
hypothe-
some detail, since it helps to explain both reverand the formation of new types.
sis in
sion
a.
were
artificially
had purple
type of
all
progeny
all
Sicilian
161
Diagram to illustrate the possible progeny from two heterozygous purple sweet peas according to data from Bateson. C, color
P, pigment facfactor (large circles) c, absence of C (small circles)
In the zygotes
tor (large crosses)
p, absence of P (small crosses).
within the checkerboard squares the gametic symbols are superimposed.
Fig. 49.
two
essential
factors
for
purple color.
When
the
GENETICS
162
two are
crossed, however,
all
sixteen
ways.
different
two
may
Number of
Zygote in
Figure 49
2
3
Among
types
6- 8-14
11 -12
(large circle)
15
16
163
Number op
Zygote in
Figure 49
Duplex for both color (large circle) and pigment (large cross)
Duplex for color (large circle) but simplex for pigment
2-5
3-9
(large cross)
circle)
(large cross)
circle)
and pigment
(large
4-7-1013
cross)
b.
Castle's
Agouti Guinea-pigs
The
tain guinea-pigs
is
due to the
fact that
pigment
is
band of
is more
The
yellow, while
most
or less concealed
leaden color.
The
is
black followed by a
by overlapping
distribution of
pigment
which
hairs
in
is
such a
when taken
as a whole.
when he discovered
that
it is
GENETICS
164
some
of the
necessary for
its
c.
expression.
Another instance of the interaction of supplementary factors is seen in the spotting of piebald
Cuenot discovered that such spotting is due
mice.
to the absence of a uniformity factor which if present
causes color to be uniformly distributed over the
entire coat.
Both
of these
may
is
Miss Durham
Miss Durham,
in
s Intensified
105
Mice
name
its
The
implies, intensity
of
color,
and
The
factors of intensity
an entirely different
nature, as they have been proven to be independently transmissible through albinos where a color
character could not appear because of the absence of
dilution of
color are of
pigment.
The following illustration of this kind of supplementary factors taken from Miss Durham's
experiments will serve to make the case clear. The
symbols employed are
:
B =
h
I
i
P
BICP
BiCP
hICP
hiCP
=
=
=
=
=
=
=
=
=
black
GENETICS
166
The
made
are represented in
Mendelian dihybrid
ratios
is
given in paren-
Black
{BICP)
X Silver-fawn {hiCP)
X Chocolate {hICP)
X Silver-fawn {hiCP)
Black {BICP)
Blue {BiCP)
Blue {BiCP)
these hybrid
unknown
before
viz.,
and brown-eyed
yellow,
167
Castle's
experiment in breeding
was made.
For the sake of clearness the formation
brown-eyed yellow is shown below in Figure
cT
Y^
S^
BE
BE BE Be
V
V
Be
be
BE bE BE be
Black
Black
Black
Black
Be Be Be bE B e be
Bfack-eijed Yellow
Black
Black -eHedYellovj
I]
12
JO
bE| BE
bE
be
Black
Be|]BE
50.
\y
Be
of the
bE B bEllbE bE be
Black
Black
Chocolate
Chocolate
13
14
15
16
b G
BE be B
Black
Black-eHed Yellow
be
bE be
Chocolate
be
Brown-e\jed Yellow
Fig. 50.
Diagram to illustrate the origin of a brown-eyed yellow guineapig from two heterozygous black parents based upon Castle's experiments. The factor for yellow (F) is present in every gamete and is
consequently duplex in every zygote but is hidden whenever the fach,
tor B is present. B, black pigment hiding brown or chocolate
chocolate (absence of B)
E, extension of B over the entire body
hiding Y e, restriction of B to eyes alone thus exposing Y over the
;
entire body.
GENETICS
168
Symbols
B =
=
Y=
E =
e =
absence of B, or chocolate.
brown or
chocolate.
restriction of
over
C = complementary
P to produce
color.
P =
P may
First Cross
(BeY)
Second Cross
black-eyed yellow
(BEY), three
(bEY),
The actual
and one brown-eyed yellow (beY).
results in Castle's experiments gave all four kinds
in close numerical agreement with this expectation.
The action of extension and restriction factors is,
therefore, plainly a case of Mendelian dihybridism
in which two independent pairs of alternative characters are concerned.
1G9
Rabbit Phenotypes
Perhaps no better application of the factor hymay be found than the case of the color
pothesis
of rabbits.
concerned, particularly
These
varieties are
now
among domesticated
quite explainable
races.
by the
The
have
GENETICS
170
many
breeders as well as
would double
Explanation of Symbols in
Br =
B =
F =
the
Foregoing Table
C to produce hroivn
C to produce black
on C to produce yellow
a factor acting on
pigmentation.
a factor acting on
pigmentation.
a factor acting
pigmentation.
The
rabbit gamete and up to date have not been separable as independent unit characters, although they
Yellow rabbits
result, as
C =
common
of
e.
production
any pigment.
factor.
It
was discovered
in 1903
by
Cuenot.
c
the absence of
of
E=
e
Si
and
visible.
Demonstrated by Castle
in 1909.
171
U=
SL
u = the absence
of uniformity
which
results in spotting
with white.
A =
Si
to be ex-
gray coat.
When
it
on
was
7.
It
The
is
differ-
phenotypically a gray
two
will
way from
all
the
it
rabbits
1)
17^
GENETICS
The Kinds of Gray Rabbits
(Color only)
173
to a gray
1,
types of young
may
list
1, 3, 6, 10, 13,
20,
Conclusion
That a relatively small number of factors may produce an extensive array of combinations is evident
from this data.
is
Finally,
It
is
now
acter
may remain
come
to
is
its
only
activity
also clear
of reversion
how
how
is
super-
a particular char-
expression
necessary to
It
easy to see
and at
hybridization, in w^hich
is
last
many
more
thought, be
to furnish far
at first
expected.
^
tions,
lenix
W.
CHAPTER IX
BLENDING INHERITANCE
1.
way
in the face of
many
is
beginning to give
steadily accumulating.
Even Mendel
BLENDING INHERITANCE
that segregation, not dominance,
is
175
Mendehan scheme.
factor in the
Imperfect Dominance
2.
dominance
plained.
is
The
is
test
necessary wlienever
double dose.
flowering plants.
all
first filial
genera-
When, however,
GENETICS
176
characterized
Andalusian
itself is
Andalusian
Andalusian
Anda usian
i~"
Black
T
Black
BlacK
T
Andalusian
Andalusian
1
Splashed White
T
Sp<- White
SpL White
Andalusian
Fig. 51.
The heredity
of
lusian.
There
is
lusian
is
dominance
is
crossed.
If
furnished
when red
BLENDING INHERITANCE
177
produce red, roan, and white offspring in the proportion of 1:2:1, thus showing that roan is a heterozygous character in which the dominance of red is
imperfect.
Even
Delayed Dominance
character which
is
really
dominant
is
sometimes
it
may
properly be termed a
delayed dominant.
fowl
is
dominant
N
GENETICS
178
These black
however, disappear at the time of the first
The complete dominance of white is, there-
molt.
fore,
simply delayed.
4.
"Reversed" Dominance
is
apparently
It
is difficult
to see
how
the presence of
expression.
to
BLENDING INHERITANCE
it
fails,
some
for
reason, to
become
179
This
effective.
by
Potency
5.
He
character
may
That
reason,
is,
is
"The potency
its
of
germi-
entire ontogeny."
if
insufficient, there
may
be either an incom-
question, or
The
it
may
fail
entirely to develop.
variations of potency
may
be grouped into
manifestation
further
word
of these
Total Potency
that
is,
by
homozygotes produced
GENETICS
180
of the
by a duplex dose
same
character.
It
is
as
if
an opaque
fluid.
h.
Partial
potency
Partial Potency
covers
all
cases
incomplete
of
produces cross-breds in
the
first
snails
also,
when
Numerous
further instances
nance could be
of
incomplete domi-
cited.
c.
Failure of Potency
for
BLENDING INHERITANCE
181
all
question.
It
heterozygotes and
duce a family in
is
all
the
in extracted
all
inhibited."
Here were two birds of the same blood, phenoand presumably genotypically alike,
which because of an individual difference in the
potency of the determiner for rumplessness produced
typically alike
same array
of hens.
GENETICS
182
Blending Inheritance
6.
segregation
into the
complete dominance.
In poultry, for example, when Cochins, which are
fashion
when
together.
It
is
portant factor
is
segregation.
in
the
of hered-
second
filial
generation
where
does
not
.^
BLENDING INHERITANCE
183
what was once beheved to be the rule has now become the exception. He goes on to say: *'One clear
may mention.
exception I
more
closely.
7.
As a
rabbit ears
may
of 118
mm.
ear-length of 210
lengths
had
is
164
ear-lengths,
when
ear-
pair
mm.
in
168
mm.
These
results
are represented
and
diagram-
This illustration is typical of many other breeding experiments made by the same investigators
^
of Inheritance
in
Rabbits."
and Cobb.
"Studies
GENETICS
184
....b'
;t
99666
<S
e
e
34 56
e
e
e
e
Offspring ofZsnd
to >
(B lo
oo
o
-
o
o o
r- t-
CM
'
c~-
6669 dd
9e
E
e
(S)
e
X)
^Q
E-
to o
iO
iO
E E
w
p r*
t*
'-
Offsprin^of land?
Fig. 52.
of
BLENDING INHERITANCE
second
filial
185
it
may
is, a
or
the
other
of
the
one
parental types, was
return to
much less apparent than evidence of blending.
Furthermore, crosses were made in which lop ears
of various fractional lengths were obtained as desired,
including
these
|, |, f , J, f , J,
fractional
lengths
and |
lengths.
apparently
Not one
segregated
of
in
were obtained in three ways first, by crossing fulllength lops with short-eared rabbits as indicated in
the first cross of the case cited above; second, by
crossing one half lop lengths together, demonstrated
by the second cross in the illustrative case given, and
Theoretically,
third, by mating J and f lop lengths.
I and I as well as f and f lop lengths would also produce I lop lengths, for in all of the crosses that were
made the length of ear behaved in a blending fashion.
These results were based, not upon a single measurement of each specimen, which might be open to
considerable error, but upon daily measurements
from the time the rabbits were two weeks old until
The
their ears ceased to grow at about twenty weeks.
growth curves drawn from these daily measurements
showed continually an intermediate or blending condition in progeny derived from diverse parents.
A Mendelian explanation of this apparently exceptional case of blending inheritance has been suggested
:
result
of
Nilsson-Ehle's
GENETICS
186
Experiment Station
of Svalof in
Sweden.
8.
first
These
generation.
drawn from
same
filial
generation,
of
red-
kerneled strain.
The explanation
is this.
of these
In the case of
BLENDING INHERITANCE
187
Taken
together,
For example, if a brown-chaffed wheat with the formula BB\ in which B and B' each represent a brownchaffed factor, is crossed w^ith a white-chaffed wheat of
the formula bb\ in which b and b' each represent the
absence
of
and B^ respecthen
tively,
all
the progeny of
this cross will be
brown-chaffed,
having the zygotic
formula
BBW.
When
upon
hybrids,
the
following
four
combina-
tions
are
and
sible,
others
fore,
first
pos-
no
bb\
filial
generation, as
shown
in Figure 53.
GENETICS
188
The numbers
how many
and
this
one
will
n
n
n
u+
n+
mn+
f
Number
determiner
two
The brown-chaff
factor,
moreover,
is
present in
BLENDING INHERITANCE
varying doses
among
these fifteen
possil)ilities, as indi-
d<
189
It
brown
is
will
evident,
be rep-
GENETICS
190
In
this
instance
out of sixteen individuals of the second generation theoretically present a perfect "blend" besix
tween the original brown- and white-chaffed grandparents, although complete segregation has actually
occurred.
Figure
for
white-kerneled
the
trihybrid
wheats
in
which
of
only
red-
and
one white-
filial
of red determiners in
figure at the
bottom
generation.
each zygote
of
is
each square.
The number
indicated
by the
vertical, horizontal
and diagonal
When
kernel.
is
the
first
generation
Pure
Fig. 56.
The
somewhat
is
all
red -h cuhite
result of
a heterozygous red of a
Hybrid red
lighter
strain.
When
together,
The
BLENDING INHERITANCE
191
greatest
m
m
m
m
m
second
filial
theoretically displayed
Figure
55,
#
m #
#
a#
generation, as
really
in
contain
produced families
third
filial
#
m#
m#
m#
m#
m#
m#
#
m#
m#
Nilsson-Ehle
red-kernel,
the
of
generation by
second generation.
was
It
be expected that, if
these hybrid wheats of the
to
+
+
+
# +
would
progeny
distributed with
to the
number
be
reference
and
of red-
white-kerneled individuals,
in the following ratios
+
+
'
r[G. 57.
Thedistributionof the
sixty-four possibilitifs in tin- F-i
generation when three similar
determiners act together as a
trihybrid.
GENI^TICS
192
3 red to
white when
is
present.
Among
white.
all
red to no white.
heredity;
(3)
segregate; and
(4)
The Nilsson-Ehle
has been confirmed in America by East in a masterly series of breeding experiments upon maize.
ers
number
it
of intergrades
number
with
of
duplicate
six duplicate
determiners
increases.
Thus
second
filial
The reappearance
1.
among 4095
BLENDING INHERITANCE
193
9.
planation
TO
the Case
of Rabbit
Ear-
length
The
so-called
blending rabbit
as pointed out
ears,
now be made
to
along
fall
with
into line,
of
segregation.
If we assume that the long ear of the lop rabbit
has only three independent but equal determiners for
excess length, the case becomes one of Mendelian
out
Nilsson-Ehle's
like
following manner:
red-kerneled wheat
in
the
52.
GENETICS
194
If all of these
220
is,
mm. -
100
mm. =
20
mm.
very
if
common phenomenon,
it
is
true that
now
If
first
gen-
ear-length,
in their
kerneled wheat.
These
possibilities
lowing f reqviencies
would be arranged
in the fol-
BLENDING INHERIJANCE
Since the average
five,
litter
among
rabbits
195
is
about
is
20 out of 64,
parent
is
filial
gen-
The
fact,
furthermore, that
the
fractional ear-
bred approximately true in the second and subsequent filial generations, may also be explained by
the Nilsson-Ehle hypothesis.
this
{^ = ^V
GENETICS
196
if
Finally,
wanting,
kinds
of
it
Human
as
Skin Color
is
crosses
between
Mendelian segregation.
In
rather of true
is
all
shown
offspring
human
families included
hundreds
of offspring
man
number,
could doubtless
large
itance.
CHAPTER X
THE DETERMINATION OF SEX
Modern
1.
From
made
Likewise
among
do-
who with
of
Sex"
of
of speculations concerning
sex.
J.
Arthur Thomson,
"The Evolution
in 1889, says
"The number
of speculations
since
Dreylincourt,
in
the
eighteenth
century,
book
roll.
GENETICS
198
of
is
Galen,
left
who
ovary
did the
mankind,
the body, "being
side
of
Modern
theories
earlier speculations,
of
may
namely, those which depend upon controllable external or environmental factors such as food, climate,
chemical dosage and will power, and those which de-
Of external
influence
control.
is
190
possibly
cells
stage of the
life
tilized egg.
been suggested that since the egg is characterized by possibly a more advanced metaboHc
condition than the sperm due to the presence of the
nutritive yolk, consequently the more yolk or nutriIt has
tion there
correct predictions.
of sex
is
bound
to turn
''evidence" to sub-
latter
have been.
unusual hardship,
GENETICS
200
series of nutrition
Cuenot
They concluded,
From
The
statistical
sources
numbers
is
of the
it
produced a
two sexes.
practical equality
an average
human
births
201
for
fe-
males.
According to various authorities, the relative number of males per 100 females is given for horses as 99,
pigeons,
of males
is
in all sorts of
MONOCHORIAL TwiNS
same
are of the
difference in
GENETICS
202
embryos
is
sl
chain of
same sex.
These facts point toward the conclusion that the
determination of sex takes place at the time of ferof the
tilization.
5.
Selective Fertilization
is
Mendelian
character.
Mendel
himself,
without elaborating this idea into a definite hypothesis, suggested the probability that sex is a heritable
character behaving in the same way as other heritable characters.
Castle,
W.
E.,
"The Heredity
4,
1903.
of Sex." Bull.
Mus. Comp.
Zool.,
203
MALC
FEMALE.
Gametes
ZYGOTES
M(r) M(M)
(DF
r(M)
Fig. 58.
ried
Diagram
by
to
show
it
question
theory,
may
mice, that
GENETICS
204
is
since a
and thus,
its
unknown.
This supposition
is
further supported
by the
fact
of mice,
if
{DD)
fail
to produce offspring.
old concep-
It
In certain parthenogenetic
present in the female, since two kinds of easily distinguishable eggs are produced, one of which develops
into males
and the other into females without ferany kind of a union with a sperm-cell.
tilization or
205
and
this sup(F.
is
(Female!
99
(Male)
9cf
probable as evidence
accumulates.
Ac-
:)
Male)
-h
W
9
QcT
(Male)
(Female)
Fig. 59.
Diagram to show tho neo-Mendelian theory of the heredity of sex, using
sex symbols.
Type
in
female
determiner
in the heterozygous
I, and
XX the female de-
case of type
The
formulae
may
be
still
who
ob-
206
GENETICS
all cases,
as
shown
positive,
by assuming
is
present in
in Figure 61.
Type
female,
since
a male.
All of these three
theoretical
agree
in
schemes
assuming
is hetero-
is
the result of
The evidence
first,
second,
hermaphrodites in heredity.
a.
A,
of
The
1.
207
"X"
Chromosome
and
McClung
in
studying the
discovered a
corisy
similar
later
phenomenon with
of its spermatozoa.
Diagram
when one parent
Fig. 62.
other
is
to
is
(1901),
respect to the
Soon
after,
chromosomes
208
shown in Figure 62
and 61.
GENETICS
or in type II of Figures 59, 60
is
invariably
number
of inde-
A,
work
2.
in university courses.
Various Forms of
X Chromosomes
its
It
may
209
be paired
female
sex.
The reason
germinal heritage.
In the microscopical evidence on this point there
is one apparent exception to the rule that females
are homozygous and males heterozygous with respect
Baltzer (1910) found that in one of the seato sex.
urchins an extra sex chromosome is associated with
the female sex, so that two kinds of mature eggs are
produced upon maturation and only one kind of
sperm-cells.
In other words, in this case the female
is heterozygous for sex and the male homozygous,
of the
is
true for
all
other
Such cases as
above in Figures
59, 60
and
61.
under type
given
GENETICS
210
Ay
3.
chromosomes
parthenogenesis, where the union of an
The behavior
of
the
in
cases
egg-cell
of
and
male
is
condition.
The
female in
all
duplex
(2
chromatin content."
b.
Castration
known
as "secondary
plumage
male
birds, the
dis-
211
(Queen)
LAR BODY
OCYTC
Polar body
^ PcLAR
BODY
Mature cgo
5perm cells
fERTlUZEO
EGG
Fig. 63.
Diagram of the heredity of sex in bees, ants and wasps. The
outline chromosomes represent sample somatic chromosomes.
The
solid black chromosomes stand for sex.
The female has two sex
chromosomes while the male has but one.
GENETICS
212
ondary sexual characters, and also that the determiners for the opposite sex are often present in a latent
condition, or, in
either
words,
other
male or female,
is
sex.
If
example,
is
Male
cut in half,
it is
worm
Ophiotrocha, for
PardSiliwllij castrated
male
Female
Fig.
64.
The
by the
cirripede, Sacculina.
Evidence that a Mendelian sex determiner is correlated with "secondary sexual characters " and that the male is heterozygous for sex
while the female is homozygous. After Smith.
now
instead of
ovaries,
proving that
it
was
originally
is
213
performed
large
is
heterozygous.
When
maleness
is
only femaleness
is
make
sex.
c.
Sex-limited Inheritance
is
a character depend-
ing
fashion
GENETICS
214
sex or the other.
by the term
*'
They
sex-limited."
(1)
This phenomenon
Color-blindness
may
Gametes
f:
is
(x
90 9@
Fig. 65.
symbol
The underscored
for color-blindness
215
is
Hnked
Sex-Hmited inheritance,
as shown in this case, may be ilhistrated l>y the (hagram on the opposite page (Fig. 65) in which, for the
Under-
2<
to a sex
From
chromosome.
diagram,
of color-blindness, that
is,
will
will, therefore,
carry
it
in a latent condition.
who
normal
of the sons,
and none
is
mate with
be color-
is,
in
color-blindness according to the sex-limited interpretation are indicated in the following table
GENETICS
216
Parents
217
Key to Symbols
Conslilulion with
respect
Phenotype
to ihe
GROSSULARIATA
factor
Gross.c? Dupl
ex
Genotype Gametes
(joi
Ooi
Gross, c?
Simplex
Iqi
Ooi
Lact.c?
fSlulliplex
()0I
Jfll
GR0SS.9
Simplex
Oof
Lact.^
Nulliplex
Ooi
flo
01
Do
Fig. 67.
for
variety grossulariata.
will
make
GENETICS
218
cross
is
numbers
45 males
(2)
to 50 females (4).
(2)
and
was confined
only
when
back to
(5).
It
was
recessive
females
(5)
in
Cross
3 (Fig.
Cro
ss
219
(0
Gross
cf
GAMETE5
Zygotes
1-5
Gross,
Gross. ^
d*
(2)
Fig. 68.
The formation
(4)
of heterozygous grossulariata individuals,
both
lacticolor
females.
Cro
5s
(2)
Gross. cf
Fig. 69.
GRoss.d
GRoss-d*
(/)
(Z)
The cross-breeding
Lact.
(5)
of
Gross,
(f
(4)
GENETICS
220
Finally,
when
these newly
made
lacticolor
males
(3)
and
limited condition.
all
satisfactory
tw^o
when present
is
determiner.
This case
is
is,
is,
as
with
Cross
221
3
(2)
Gross d
(Ooioi)
7(X
Gross,
d*
Lact. 5
(3)
(5)
(2)
Fig. 70.
Lact. c?
6Z-'
Gross ^
lacticolor female.
known
to occur in
Cross 4
GROSS
tACT.
d"
(5)
(2.^
Fig. 71.
Back cross of lacticolor male with grossulariata female producing the original sex-limited condition in which all the females are of
Raynor.
Data
for Figures
GENETICS
22
Among
such dimorphic
by
may
ing progeny indicate plainly that one sex is homozygous while the other is heterozygous with respect to
the sex character. This confirmatory evidence is
quite in line with that already brought forward that
is
Conclusion
7.
far obtainable
is
is
fertilized,
way
from
all
sources
unalterably fixed
by
definite deter-
as other
Mendelian
"Nearly
statement:
is
the
all
recent
investigations
while
it
of the individual."
it
is
man
it
shows
to predetermine the
may
do.
The
Castle
"Negative as are
sex control, they are perhaps
223
ments."
and devote
it
to
more
profitable employ-
CHAPTER XI
THE APPLICATION TO MAN
1.
Human
hand
civilization goes
in
man
exerts
Primitive
by his
Any knowledge
of
the
a distinct contribution to
human
and
plants,
progress.
fied
224
content,
rust,
225
Such an immediate
result
heredity in nature.
of civilization
man.
ferent story to
2.
tell
will
have a
dif-
Man
Thus with
of natural selection
man
is
partially under
"grace"
GENETICS
226
with
artificial
means
of self-preservation,
We do not
life
of usefulness.
modern
civili-
some
and universal "natural laws"
the case of man, and it is profitable
most
inflexible
are ineffective in
to bear this in
netics to
The
of ge-
man.
laboratory for
world, but
it is
human
heredity
is
the wide
227
consisting in an analysis of experiments already performed rather than in initiating new experiments.
Such
institutions
sanitariums,
insane
as
and homes
asylums,
prisons,
the
unfortunate are
excellent foci for studying certain phases of human
heredity, because they are simply convenient places
for
3.
in genetics
The Jukes
A classic example of an experiment in human heredwhich has been partially analyzed by the statistical method is that furnished by Dugdale in 1877 in
the case of "Max Jukes" and his descendants. It
includes over one thousand individuals, the origin of
all of whom has been traced back to a shiftless, illiterate, and intemperate backwoodsman who started
his experiment in heredity in western New York
when it was yet an unsettled wilderness.
In 1877 the histories of 540 of this man's progeny
were known, and that of most of the others was
About one third of this degenerate
partly known.
strain died in infancy, 310 individuals were paupers
who all together spent a total of 2300 years in almsIn addition
houses, while 440 were physical wrecks.
were
descendants
female
the
to this, over one half of
crimprostitutes, and 130 individuals were convicted
Not one of the entire
inals, including 7 murderers.
ity
family had a
common
school education,
although
GENETICS
228
and 10
Only 20
of these did so in
state's prison.
It
is
human
dollars,
is
many
professor of law;
governors,
Members
229
constitutions,
Steamship Company 15 railroads, many banks, insurance companies, and large industrial enterprises
have been indebted to their management. Almost
if not
every department of social progress and of
public weal has felt the impulse of this healthy,
long-lived family.
It is not known that any one of
;
of crime."
Quaker
girl
forefathers.
woman
1
of
a respectable
live
"
From
life
after
this legal
H. H. Goddard.
All of
GENETICS
230
been the
determining factor in the formation of their respective characters."
4.
These instances of human breeding show unmistakably that "blood counts" in human inheritance,
even though the hereditary unit characters that lead
to these general results have not yet been analyzed
laws of heredity.
For instance, as an
231
Max
moral and
trait."
5.
and character
traits of eye-color
of hair,
may
the mass,
be regarded as indifferent
is
concerned,
*'
long
list
of
blood" may,
handicap
social
traditions of
in a general
way
to be subject
GENETICS
232
yet undergone
the careful
analysis
demanded by
modern
than
lump
in
inheritance.
6.
Hereditary Defects
and
also in
man in one
(according to Elderton),
ment
233
human
hereditary
defects,
although
in
most
and incomplete.
Deafness, for example,
is
a defect which
it is
is
heredi-
at present
Condition op Parents
statistically.
GENETICS
234
is
happen
to
recombine after segregation to form the new individDeafness will be produced in the offspring only
ual.
when matings occur in which the proper factors are
combined. Such an undesirable result is much more
likely to happen if both parents come from the
same, or related, hereditary strains than if they are
derived from families in no way connected by blood.
Herein lies the biological objection to cousin
marriage which tends to bring together, and thus
Outcrossing, on the
to perpetuate, like defects.
contrary, through the law of dominance, tends to
conceal defects and to prevent their expression.
Many other cases of human defects, such as imbecility or insanity, are extremely difficult of analysis
from the standpoint of heredity because, in the first
place, the defective conditions descriptively included
factors.
and to
learn,
if
possible,
in
^285
(Davenport)
7.
The method
of possible control of
human
defects
GENETICS
236
two cases
only, namely, in 3
fective parent
is
of unaffected offspring,
defect
is
5,
is
and even
way
dominant
defect
is
recessive individuals.
time to shake
off
It
is
ob-
germplasm of a
by continued mating with
By this method it is possible
in
and
heterozygous,
to
rid
the defect.
it
When
it
once
dis-
thereafter
is
and
will
an entire absence
it
is
of the character
since
something
result
quite
is
different,
normal homozygote, as
If
in case 4 of the
237
be normal
accompanying
in
appearance,
when
EkO
[Nh<N)[rU^^
N]-r{N)@[r]LN
c
Fig. 72.
A, alcoholic C, criminalistic
After Goddard.
;
d,
died
T, tubercular.
GENETICS
238
ably
is
if
highly unfit,
to
Weakness
fit.
in
with strength in
may
Inbreeding
is
is
injurious
Davenport.
239
any Hne.
the Jews,
On
known human
living
ancestors.
If
this
theoretical
supposition
The majority
of
plants are probably self-fertihzed while hermaphroditic animals, which sometimes at least are selffertilized particularly
among
very common.
Nature has secured, on the other hand, often by
GENETICS
240
among
insects
9.
Numerous experiments
or-
ganisms.
Darwin,
for
instance,
planted
morning-glories,
two beds which were laid out side by side, that is,
in an environment as nearly the same as possible,
but with half of the beds screened from insects
which usually transfer pollen from flower to flower.
In the screened half where all insects were excluded
the flowers were of necessity self-fertilized, while in
The
by the
insects
which had
free access to
them.
two beds were kept separate and the experiment was continued for ten years,
so that at the end of that time two lots of morningglories, one self-fertilized for ten generations and the
other presumably cross-pollinated for the same length
seeds produced in the
were obtained for comparison. The criteDarwin used was the vigor of the plants as
shown by the length of the vine. He found that the
of time,
rion
241
is
detrimental.
it
fell
to 3.2.
and obtained a
for
twenty-nine genera-
For the
parallel result.
number per
litter
first
was
many
self-fertilization,
and
On
Drosophila,
Castle
inbred
brother
and
sister
fly,
for
the
line.
No
all
will
fertility
apply equally
cases.
10.
Inbreeding
is
In-
asylums
of various
GENETICS
242
to insure
human
inbreeding.
Similarly, localities
by migrations
of the
The
ness.
body
spell of proximity,
The
effects of stagnation
11.
Inbreeding in
ous.
itself
may
The consequence
of inbreeding as
is
shown by
that latent or
traits
which
recessive characters
243
inefTec-
tive.
Cousin-marriages, although producing a high percentage of defects, do not necessarily reproduce unThey simply bring out latent or
desirable traits.
recessive characters for the reason that under these
and cause
it
not to appear.
is properly regarded as the
it
more
defective strain.
Outcrossing always increases heterozygous combinations in the germplasm and covers up undesirable
recessive traits
tional
dominant
to the surface.
traits.
CHAPTER
XII
HUMAN CONSERVATION
1.
be Improved
method
consists in
The
The
first is
of
born."
of
Any
human
betterment, however,
The
biologist
who
is
equally at
who
245
fails sufficiently
to real-
of
More
Facts Needed
be largely statistical,
collect
more
facts.
it is
Our
human
of the first
must
heredity
importance to
actual knowledge
is
con-
much of
foundations. The
upon questionable
is to learn more facts
to sift the
and to
truth from error in what is already known
which
rests
reduce
all
Much progress
is
being
made
in this direction,
by the
owing
revival of Mendel's
in its infancy.
GENETICS
246
human
beings and
a word,
it
is
and practice
science
of
eugenics."
The
Record
publica-
Office has
twenty-five
is
Graham
and has now systematically arranged particulars concerning the history of over 20,000 individuals.
In
Association
States.
HUMAN CONSERVATION
^2^7
3.
practice.
any program
of
an academic
affair for
is
bound
to remain largely
rize over.
It
is
analysis
is, it
must be followed by a
or
woman
overwhelming demands
In the midst
of the present,
the
plasm he
it
would be
when
it is
it
on to posterity
and
it is
in the
only
when
rank and
file
this
of society as well as
among
the
GENETICS
248
leaders,
ment
of
better-
4.
plasm
negative
world
way
Davenport and
"dry up the streams that feed the torrent of defective and degenerate protoplasm."
This may
be partially accomplished, at least in America, by
employing the following agencies control of immigration
more discriminating marriage laws
a
is
and
finally, drastic
zation or sterilization
a.
when
sexual segregation of
measures of asexuali-
necessary.
Control of Immigration
much
HUMAN
Eugenically, the
CONSER\\\TION
weak point
210
way.
in the
knowledge
of the
germplasm
GENETICS
250
shall
country.
The United
already has
field
desirable animals
amount increased.
Department of Agriculture
sees this
States
this
human
blood
Is the inspection
less
important
and super-
make customs
marriage.
*' Marriage,"
says
Davenport, "can be looked at from many points of
view.
In novels as the climax of human courtship
in law largely as two lines of property descent
in
society, as fixing a certain status
but in eugenics,
which considers its biological aspect, marriage is an
experiment in breeding."
Certain of the United States have laws for-
HUMAN CONSERVATION
251
bidding the marriage of epileptics, the insane, liahitual drunkards, paupers, idiots, feeble-minded, and
those afflicted with venereal diseases.
It would be
such laws were not only more uniform and
widespread, but also more rigidly enforced.
well
if
It
is
for
illegitimacy
influence
is
in
regulating
marriage
and consequent
reproduction.
An
Educated Sentiment
GENETICS
252
with insanity, and also in America with respect to miscegenation, but a cautious and intelligent
afflicted
make up
tracting parties
The
less usual.
scientific attitude is
to the romantic
is
is
way
simply "organized
of looking at things.
common
sense,"
Science
and romance,
it
may
Segregation of Defectives
and certain
criminals,
of
plasm
may
HUMAN
CONSER^'ATIOx\
258
society.
disease
denied
'
'
(Davenport)
"The
There
administration
sexual segregation
is
same
institutions, unless
may
GENETICS
254
"In the
for centuries
In the competition of life a prehas thus been placed on imbecility and disease.
est of struggles.
mium
The
cretin
goitre,
the union.
and
The
result
cretin,
religion
is
that idiocy
multiplied
is
The cretin of Aosta has been dea new species of man. In fair weather
intensified.
veloped as
the roads about the city are lined with these awful
human beings with less intelligence than
paupers
a goose, with less decency than the pig."
Why mper, writing in 1880, further observes: "It
is
as
known
is
it
is
well
are sanctioned
is
and
all
in
gone.
e.
fifth
plasm
in
method
Drastic Measures
of restricting undesirable
germ-
HUMAN CONSERVATION
255
imbeciles,
fined to the
quent parenthood.
In the female the corresponding operation, which
consists in removing a portion of each Fallopian
tube, is much more severe, but not impracticable or
dangerous.
Eight states
our land.
5.
Not only
able germplasm, but also positively by the conservation of desirable germplasm, may the eugenic
ideal be approached.
1
GENETICS
256
It is possible that
some
if
of the philanthropic
endeavor now directed toward alleviating the condition of the unfit should be directed to enlarging the
opportunity of the fit, greater good would result in
the end. In breeding animals and plants the most
notable advances have been made by isolating and
developing the best, rather than by attempting to
raise the standard of mediocrity through the elimination of the worst.
exceptional pupil
who
is
it is
up to a passing standard.
The campaign
for
human
The
following
By
fit
is
what
is
meant by subsidizing
Dec.
"Berlin.
11,
1911.
The Empe-
newspaper clipping
unconfirmed
concerned.
ror
improvement
the
this
German type
of
humanity,
type which
HUMAN CONSERVATION
will
if
257
in its characteristics,
out.
The
Only
'typical' couples
The man
the
fixed type
is
is
to be not
to be produced as follows
more than
woman
Neither the
Its tint
or
'beefiness.'
to
be
woman must
bear a
be
of
German
the language of
It
should
eugenic ideal
ical
is
traits alone.
GENETICS
258
b.
By
Much
is
in
better
homes,
better
all
euthenic endeavor,
living conditions,
in
short,
is
present in
man.
c.
Much
By
good protoplasm
fails
to find expression in
1.
Preventable Death
unqualified stay at
When
eration.
the hospital,
off,
but
it is
strain of
home
it is
man who
is
cut
germplasm.
The Thirty
Years'
War
in
HUMAN CONSERVATION
Germany
259
Napoleon in his
campaigns drained the best blood of France.
David Starr Jordan has presented this matter
very clearly.
He
"man
with a
not the
oppression, as he has been
pictured, but rather the dull progeny resulting from
result of
centuries
is
of
who were
left
behind when
fit
2.
Social Hindrances
life
is
tell
their
own
GENETICS
260
Who
shall
sit in
Judgment
unfit
many
difficulties, for
who
fit
is
quali-
from the
The
list
who were
commanding appearance,
HUMAN CONSERVATION
^2G1
ralities.
of her sisters
murdered
his
owed
his
condemn
her.
"The
very conflicting.
The
subject
is
many and
difficult one,
and
I for
judgment may
err as well
GENETICS
262
what
capable
of,
fearful blunders
but
it is
human judgment
applying to
man
is
that in
eugenic inquisition.
It
is
path
of
human
achievement.
the future
BIBLIOGRAPHY
A
listed
below.
Bateson, W.
1909.
Baur, E.
Castle,
1911.
Gebriider Borntraeger.
lehre.
W.
Cambridge
Cambridge.
University Press.
E.
Berlin.
1911.
New
Animal Breeding.
Tower.
York.
1912.
Chicago.
CoRRENS, C.
1912.
Borntraeger.
Darbyshire, a. D.
East, E.
M.
ciples
1907.
to
The Mac-
York.
Gebriider
Berlin.
1912.
Cassell
coverv.
1912.
New
millan Co.
The Relation
Plant Breeding.
Exp. Sta.
263
London.
158.
Conn. Agric.
BIBLIOGRAPHY
264
GoDLEWSKi, E.
1909.
Engelmann.
Entwicklungsmechanik.
GoLDSCHMiDT, R.
Einfuhrung
1911.
Engelmann.
schaft.
JoHANSSEN, W.
1909.
Fischer.
Jena.
in die
Leipzig.
Vererbungswissen-
Leipzig.
Vieweg
Haecker, V. 1912. Allgemeine Vererbungslehre.
und Sohn. Braunschweig.
1911.
The Social Direction of Human
Kellicott, W. E.
Evolution.
D. Appleton and Co. New York.
Lock, R. H.
Recent Progress
1909.
LoTSY,
1906-1908.
P.
J.
Fischer.
theorien.
Morgan, T. H.
Vorlesungen
Study
of Variation,
London.
iiber
Descendenz-
Jena.
Experimental Zoology.
1907.
New
millan Co.
in the
Murray.
The Mac-
York.
Montgomery, T. H.
in
Animals.
PuNNETT, R. C.
New
York.
Reid, Archdall,
man and
Thomson, J. A.
Watson, J. A.
Co.
Heredity.
1912.
Murray.
Heredity.
London.
York.
1904.
ScHALLMAYER, W.
DE Vries, H.
Chap-
Principles of Heredity.
London.
1908.
S.
New
Weismann, a.
The
1905.
Hall.
1905.
Mutation.
Fischer.
Jena.
by
Chicago.
INDEX
Abnormal
Bateson,
fertilization, 30.
Abraxas, 216.
Ageniapsis, 202.
Agouti, 163, 171.
Albinism, 59, 151, 232.
Alcoholism, 93.
Alternative inheritance, 121.
Ambly stoma, 90, 129.
American Breeders' Assoc, 245.
Amitosis, 20.
Ammonites, 71.
Amphimixis, 55, 81.
Anaphase, 20.
Ancon
sheep, 68.
Angora
hair, 129.
of,
Beecher,
129.
171.
70.
Bees, 210.
Begonia, 7.
Belemnites, 71.
Belgian hare, 183, 193.
Bell, 246.
Beneden, van, 21, 23.
Bertillon system of identification. 38.
BiFFEN, 129, 224.
Bimodal polygons, 48.
Antirrhinum, 173.
Ants, 210.
Aphids, 204.
Biometry. 42.
Birth-marks, 87.
Blending inheritance, 121, 174, 182.
Asexual spores, 9.
Atavism, 146.
Autosomes, 209.
Average deviation, 45.
Axolotl, 90.
Azaleas, double, 63.
Blumenbach,
Booted poultry,
Born, 200.
BURBANK,
156.
Banana
Banded
Capsella, 89.
Drosophila).
Bryonia, 222.
Bryozoa, 8.
Calvin, 260.
Canaries, 129.
shell, 129.
182.
Balls, 129.
Baltzer, 209, 220.
fly (see
197.
265
INDEX
266
Castle, 4,
17.
Chapin, 261.
Characters, congenital, 80.
moral and mental, 230.
240, 251.
Datura, 180.
unit, 61.
Davenp6rt,
90.
Chromatin,
222.
Cotton, 129.
Cousin marriage, 234, 238, 242, 243.
Crabs, 212.
Crested head, 69, 129.
Cretinism, 253.
tailless, 68.
Centrosome,
16.
Chromosomes,
16.
Death,
extra, 207.
"x," 207.
"y," 209.
Chromosome
7.
preventable, 258.
Defects, hereditary, 232.
Chrysanthemum, 50.
Circumcision, 87.
Dermal
Cirripede, 212.
Clover, four-leaved, 39.
Cobb, 173, 183.
Cochins, 182.
Cocoon, yellow, 129.
theory, 29.
Colorado potato-beetle
(see
Lep-
tinotarsa)
Color-blindness, 214.
characters, 69.
Complementary
factor, 159.
Compound
determiners, 159.
Congenital characters, 80.
Docked
factor, 170.
Comb
spines, 151.
Dogs,
tails, 88.
hairless, 68.
tailless, 68.
INDEX
267
hypothesis, 150.
inhibitory, 160.
intensifying, 165, 171.
pattern, 163, 171.
pigment, 101.
DUGDALE, 227.
Duplex dose, 133.
spotting, 171.
Durham,
165, 171.
East, 192.
Echidna, 150.
Echinoderms, 31.
Farrabee,
Echinus, 32.
Education, 3.
Edwards, Jonathan, 228, 260.
Richard, 261.
Egg, abortive, 23.
cell,
21.
fertilized, 24.
branched, 69.
recurved, 69.
twisted, 69.
Feeble-mindedness, 149.
Feet of Chinese women, 87.
Feral animals, 150, 152.
Fertilization, 24.
human,
abnormal, 30.
28.
mature, 23.
selective, 202.
ElMER, 40.
Elderton, 232.
Elementary species,
Environment, 2, 88.
Enzyme
129.
Fay, 233.
Firefly, 207.
Fission, 7.
62, 72.
theory, 33.
Flavism, 151.
Fluctuation, 57.
Four-leaved clover, 39.
Erinaceus, 150.
Erithizon, 150.
Freaks, 58.
rules, 238.
sentiment, 251.
Eugenics, 244.
Record
Office, 245.
Euthenics, 244.
Evening primrose (see (Enothera)
Freckles, 81.
Frogs, 200.
G.A.GE, 83.
Galen,
198.
G.A.LTON, 42, 77, 98, 117. 120-122.
151, 164, 244.
Galton Laboratory
for
Gamete, 23.
Garden peas, 124-128,
Geddes, 197.
Gemmules, 8.
132. 134.
complementary, 159.
Generation, spontaneous,
Genetics, 2.
compound,
Eugenics,
246.
Germ-cells, 21.
6.
INDEX
268
GOLDSCHMIDT, 44.
Graduated variants, 108.
Greyhounds, 33, 201.
Greening apple, 8.
Green peas, 134.
Growth, 7.
Gruber,
16.
Haecker,
Hair
17, 129.
Hairlessness, 68.
Hallet, 152.
Hare, Belgian, 183, 193.
Harelip, 149.
Hauser, 256.
Hedgehog, European, 150.
Heine, 260.
Helix, 178.
Henking, 207.
Hereditary bridge, 27.
Heredity, definition of, 4.
Hereford
cattle, 68.
Heritage,
2.
Hodge, 92.
Homozygote,
HOOKE, 15.
Hymenoptera, 210.
Illegitimacy, 251.
Imbecility, 234, 238.
Immigration, 248.
Immunity to rust, 129.
Imperfect dominance, 175.
Inachus, 212.
Inbreeding, 238.
Independent unit characters, 144.
Indian corn, 241.
Induction, parallel, 83, 93.
somatic, 83.
Influence of proximity, 241.
Inheritance, alternative, 121.
biological, 75.
blending, 121.
particulate, 121, 164.
sex-limited, 213.
Inhibitory factor, 181.
Insanity, 234.
Insects and flowers, 240.
Instinct, 92.
Japanese
art, 37.
Jamestown weed,
180.
131.
Horns
Hyalodaphnia, 54.
Hybridization, 120.
Jungle-fowl, 148.
in sheep, 179.
in cattle, 179.
Horses, birth-rate
of,
201.
hairless, 68.
pacing, 129.
trotting, 129.
Human
betterment, 244.
birth-rate, 201.
Kammerer,
90.
egg, 28.
Human
stature, 98.
traits, 231.
Hurst,
133.
Lamarck,
53, 76.
INDEX
Laughlin, 246.
Lederbaur, 89.
Leghorn poultry,
1^09
MOHL, VON,
177, 182.
15.
Montgomery,
Live-for-ever, 54.
Mud
LoEB, 27.
Lop-eared rabbit, 183.
Mullenix,
MacDougall,
67, 83.
Mantle
fibers, 19.
puppy, 91.
173, 183.
Mutilations, 87.
Maturation, 22.
Nat. Assoc, of
British
and
McClung,
Mean,
Necturus, 91.
207.
arithmetical, 45.
Melanism, 151.
Mendel,
Metaphase,
19.
Neo-Darwinians, 77.
Neo-Lamarckians, 77.
Neo-Mendelian theory of
Nicodemus,
85.
Night-blindness, 238.
Nilsson-Ehle, 185.
NiTOBE, 37.
Nuclear membrane,
(Enothera, 64.
Oesterleben, 201.
Oil-gland, 69.
Oocyte, 23, 211.
Oogonia, 23.
Mode,
Ovid,
45.
15.
Nucleus, 15.
Nulliplex dose, 133.
Nutrition theory of sex, 198.
6.
sex, 205.
Nettles, 129.
Newman, 202.
Newton, 260.
Ophiotrocha, 212.
Origin of life, 5.
of species, 1.
Milton,
Irish
Millers, 224.
6.
INDEX
270
Paramecium,
Pug
Ovists, 22.
48, 110.
Pasteur,
6.
jaws, 69.
PUNNETT,
Pure line,
127, 216.
103.
Puritan stock, 239.
Purple beech, 63.
Patterson, 202.
Pearson, 42, 43,
49.
177.
Pigment
factor, 161.
Pomace
fly (see
Poppy, Shirley,
Redfield,
79.
flowers, 129.
Regeneration, 210.
Regression, 98, 151.
Red
Drosophila).
63.
Population, 115.
Porcupine, 150.
Potency, defined, 179.
Reinfection, 94.
partial, 180.
total, 179.
Poultry, birth-rate
booted, 182.
of,
201.
173.
Riddle, 83.
RiMPAU, 153.
Ritzema-Bos, 241.
Roan
Rock-pigeon, 148.
Roses, double, 63.
INDEX
271
Rotifers, 204.
Spermists, 22.
Spha;r echinus, 32.
Spines, dermal, 151.
Sacculina, 212.
Sadler, 198.
Salamandra, 90.
Saunders, 129.
Sponges,
14.
Sedum,
G.
Spores, 9.
Sports, 39, 56.
Schwann,
7, 8.
Spontaneous generation,
54.
Sprengel, 63.
Spurs, extra in poultry, 69.
Standard deviation, 46.
Standfuss, 70.
Starchy kernel, 129.
Starfish rays, 44.
Statoblasts, 8.
Stentor, 16.
Sterilization laws, 255.
Stevens, 207.
Stevenson, R. L., 260.
Stockard,
39.
Silk-worm, 129.
Simplex dose, 133.
SiTOWSKI, 83.
Skew polygons, 50.
Skin color in man, 196, 231.
Smith, 212.
Supplementary
Smooth-margined nettle
Survival of the
fittest,
leaves, 129.
Smooth
Sumner,
89.
Sunburn, 88.
Sunflower, 129.
226.
Susceptibility to disease, 93.
to rust, 129.
Sweet pea, 13, 160.
Taillessness, 68.
Tails, docked, 88.
Tatusia, 202.
cycle, 71.
Telophase, 20.
Tennent,
Theory,
31.
cell, 14.
chromosome, 29.
enzyme, 33.
Spencer,
origin of,
1.
92, 260.
nutrition, 198.
Theromorphs, 71.
Thigh bones absent,
69.
INDEX
272
Thomson,
Thumb
non-hereditary, 41.
normal, 40.
orthogenetic, 39.
146, 197.
prints, 38.
Thury,
198.
Tineola, 83.
Toes, crippled, 87.
extra, 69, 178.
webbed,
physiological, 38.
psychological, 38.
quantitative, 41.
qualitative, 41.
single, 39.
somatic, 40.
useful, 39.
Varieties, 62, 72.
Vasectomy, 255.
69.
Tomato,
108.
Vermiform appendix,
2.
human, 231.
Treasury of
Human
Inheritance,
246.
150.
Vestigial structures, 149.
ViLMORIN, 155.
Viola, 37.
Vitalism, 82.
Volta Bureau, 246.
Voltaire, 260.
Trihybrids, 140.
TrUobites, 71.
Trimmed ears of dogs, 88.
Walter,
Tuberculosis, 38.
Tuttle, Elizabeth, 260.
Twins, 201.
W^ar, 258.
Two-celled
Tyndall,
fruit, 129.
6.
Wart-hog, 91.
Webbed
Uniformity factor, 164, 171.
Unit character, 61.
Urosalpinx, 47.
toes, 69.
Weismann,
81,
10,
7,
86-88,
84,
55,
94,
77,
95,
241.
129, 152, 186, 224.
Whymper, 254.
Wheat,
Variability, relative, 47.
Variation, abnormal, 40.
continuous, 40.
convergent, 150.
definite, 39.
discontinuous, 41, 69.
fluctuating, 43.
fortuitous, 39.
germinal, 40.
graduated, 52.
harmful, 39.
hereditary, 41.
indefinite, 39.
indifferent, 39.
integral, 52.
morphological, 38.
multiple, 39.
fHOfERTT
Wiedersheim, 87.
Wilson, 34, 35, 207, 208.
Wing
absent, 69.
WiNSHiP, 228.
Woltereck,
53.
X chromosome,
Y
207.
chromosome, 209.
Yellow mice, 203.
peas, 134.
200.
Yung,
Zygote, 24.
UMAIF
78,
123,
HTHE
following
pages
contain
advertisements
on
kindred
of
subjects
The
Human
Science of
Behavior
By
MAURICE PARMELEE
Professor of Sociology, University uf Missouri
This
recent
is
the
work
first
book
neurology
in biology, zoology,
work
the determination of
feehng,
and the
The book
most
in particular, in genetic
in anthropology,
showing the
sig-
fundamental factors in
instinct, inteUigence,
of work which
behavior and
work
lege
it
will
is
of so
human
much
nature.
To
those engaged
will
in
in certain courses in
psychology and
sociology.
human
educational
New York
An
Introduction to Zoology
By
ROBERT W. HEGNER,
Ph.D.
Illustrated,
in a
all
way
that
the phyla
sized, but
is
is
is
as
viewed
inten-
The
animals are not treated as inert objects for dissection, but as living organ-
No arguments are
necessary to justify the " type course," developed with the problems of or-
ganic evolution in mind, and dealing with dynamic as well as static phe-
nomena.
" I have read your chapter (The Crayfish and Arthropods in Genand can express my satisfaction with reference to the general
eral)
"
The plan
is
New York
COLLEGE ZOOLOGY
By
ROBERT W. HEGNER,
Ph.D.
Illustrated, Cloth,
This book
12mo, xxiv
+ 733 pp.,
$2.60 net
intended to serve as a text for beginning students in uniwho have already taken a course in
general biology and wish to gain a more comprehensive view of the animal
kingdom. It differs from many of the college text-books of zcxilogy now
on the market in several important respects: (i) the animals and their
organs are not only described, hut their functions are pointed out; (2) the
animals described are in most cases native species; and (3) the relations
of the animals to man are emphasized.
Besides serving as a text-book, it
is believed that this book will he of interest to the general reader, since it
gives a bird's-eye view of the entire animal kingdom as we know it at the
present time.
Within the past decade there has been a tendency for teachers of zoology
to pay less attention to morphology and more to physiology.
As a prominent morphologist recently said, " Morphology ... is no longer in favor
and among a section of the zoological world has almost fallen into
disgrace" (Bourne). The study of the form and structure of animals is,
however, of fundamental importance, and is absolutely necessary before
physiological processes can be fully understood; but a course which is
built up on the " old-fashioned morphological lines" is no longer adequate
versities
and
is
The present volume has not been made by merely adding a descripti'>n
of the vertebrates to the author's " Introduction to Zoology" (for a brief
account of which see the last pages of this circular). On the contrar)-. it
is a new work throughout, although the same general method of treatment,
book, has been employed in this
book the author has submitted
the manuscript of each chapter to a scholar and teacher of un(|uestioned
in the earlier
field.
The
criticisms
se-
cured have greatly increased both the accuracy and the practicability of
the text.
New York
The
in
Cefl
EDMUND
B.
WILSON,
Ph.D.
first
it
changed, most notably in case of those that are focussed in the centrosome
cell-division
and
This has
fertilization.
page.
It
illustrations
made
The endeavor
has, however,
fifty
still
new
been
limits, even at some cost of comand the present edition aims no more than did the first
to cover the whole vast field of cellular biology.
Its limitations are, as
before, especially apparent in the field of botanical cytology.
Here progress has been so rapid that, apart from the difficulty experienced by a
zoologist in the attempt to maintain a due sense of proportion in reviewing the subject, an adequate treatment would have required a separate
to
prehensiveness
volume.
more
fully
considered
renders increasingly
difficult
been followed.
New York
An
the
Theory
Outline of
of Organic Evolution
By
Phenomena 'which
MAYNARD
M.
it
explains
METCALF,
Woman's
Ph.D.
College of Baltimore
The
lectures out of
which
this
Woman's
for the
this
than a burden.
In
many
form an
in
This method of
treatment has been chosen both for the sake of the greater vividness thus
Many
it
later to
become
familiar.
New York
II
i i
ill
III!
HI
11
lii
liipi