Transferring alien genes from related species and genera for wheat improvement

Wide hybrids in the Triticeae tribe have been attempted and studied for over a 100 years. The first such
hybrid was between wheat and rye (Wilson, 1876). Rimpau (1891) described 12 plants recovered from
seed of a wheat-rye hybrid that represented the first triticale. Farrer (1904) reported studies on wheat-
barley hybridization; however, Shepherd and Islam (1981) considered it improbable that these were true
intergeneric hybrids. Several perennial grasses were hybridized with wheat in the early 1930s with the
objectives of transferring disease resistance and perenniality into annual crops. Many hybrids
involving Triticum and several Aegilopsspecies were also made during the 1920s and 1930s from which
the genomic relationships of the two genera were derived. The large-scale practical use of the hybrids,
however, was delayed until the advent of colchicine treatment in the late 1930s. The ability to double
the chromosome number of hybrids using colchicine had both practical and theoretical consequences.
The production of fertile amphiploids provided the way to develop X Triticosecale Wittmack as a new
cereal crop. Also advanced were evolutionary studies when McFadden and Sears (1946) resynthesized T.
aestivum, discovering that Ae. tauschii (syn. T. tauschii) was the D-genome donor to bread wheat.

With the advancement of hybridization techniques (Kruse, 1973) and embryo culture (Murashige, 1974),
wide hybridization became a more common practice involving more perennial species. In reviews of the
progress of wide hybridization, intense interest was expressed among breeding programmes in utilizing
the genetic resources available in the perennial Triticeae for cereal improvement (Dewey, 1984; Sharma
and Gill, 1983a; Mujeeb-Kazi and Kimber, 1985; Sharma, 1995; Wang, 1989).

Of the approximately 325 species in the tribe Triticeae, about 250 are perennials and 75 are annuals
(Dewey, 1984). Relatively few perennials have been hybridized with wheat essentially because of the
complexity of doing so and due to embryo rescue/regeneration constraints. The perennials, which
include many important forage grasses, have the potential to serve as a vital genetic reservoir for the
improvement of annual grasses. These include the major cereals: bread wheat, durum wheat, triticale,
barley and rye. Perennials successfully utilized for improving wheat are predominantly in
the Thinopyrum group.

This chapter will focus on achieving agricultural production targets with emphasis on bread and durum
wheat. Such production targets are to be achieved by enforcing crop improvement protocols based
upon the utilization of genetic diversity, crucial for durability of stress resistances and tolerances and for
ensuring sustainability. Major emphasis is devoted to consideration of the exploitation of ‘alien’ genetic
diversity, encompassing interspecific and intergeneric hybridization categories.

DISTRIBUTION OF GENETIC DIVERSITY: GENE POOLS

Though genetic diversity can be induced, for more controlled, well-directed incorporation, diversity
naturally present in the annual and perennial Triticeae species has priority. This natural diversity resides
in the conventional wheat germplasm and in closely or distantly related alien species sources. The
species resources are distributed within gene pools, and genetic transfers can be realized for wheat
improvement from these pools over short- or long-term time frames. The gene pools are structured
upon the genomic constitution of the species and are comprised of three groups: primary, secondary
and tertiary.

The primary gene pool species include the hexaploid landraces, cultivated tetraploids, wild T.
dicoccoides and diploid donors of the A and D genomes to durum and bread wheats. Genetic transfers
from these two genomes occur as a consequence of direct hybridization and homologous recombination
with breeding protocols contributing different back-crossing and selection strategies. Some cross
combinations require embryo rescue, but no cytogenetic manipulation procedures are necessary. The
secondary gene pool is formed of the polyploid Triticum plus Aegilops species, which share one genome
with the three genomes of wheat. The diploid species of the Sitopsis section are included in this pool,
and hybrid products within this gene pool demonstrate reduced chromosome pairing. Gene transfers
occur as a consequence of direct crosses, breeding protocols, homologous exchange between the
related genome or through use of special manipulation strategies among the non-homologous genome.
Embryo rescue is a complementary aid for obtaining hybrids. Diploid and polyploid species are members
of the tertiary gene pool. Their genomes are non-homologous. Hence, genetic transfers require special
techniques that assist homoeologous exchanges, facilitated by irradiation or callus culture mediated
translocation induction.

Diploid and polyploid species with genomes that are non-homologous to wheat reside in the tertiary
gene pool. Homologous exchanges cannot affect genetic transfers, but genomic homoeology of these
species does permit the transfer of genes by somewhat complex protocols.

UTILIZATION OF GENE POOL DIVERSITY

For practical end products to be obtained, some transfer prerequisites that encompass all three gene
pool species are hybrid production, embryo rescue, plant regeneration, cytological diagnostics, breeding
methodology and stress screening, culminating in the stability of the advanced derivatives contributed
by homozygosity. Based upon these prerequisites and genetic transfer ease, primary gene pool diversity
is most important for wheat improvement. The species of the diploid A and D genomes contribute novel
genes and allow direct recombinational exchanges with their respective genome partners to facilitate
both durum and bread wheat improvement over a relatively short-term time frame than what is
provided by the secondary or tertiary gene pool species. Setting the above as a basis for genetic
introgression, the strategy and outcome of exploiting the diversity of the three gene pools and their
accessions is illustrated.

PRIMARY GENE POOL

The A genome (Triticum boeoticum, T. monococcum, T. urartu; 2n=2x=14, AA)

One avenue of using the A-genome accessional diversity is via bridge-crossing of the AABBAA
amphiploids (Figure 11.1a). In general, the durum parents x A-genome accession crosses are simplistic
and of high frequency (Table 11.1). Meiosis of F1 hybrids (2n=3x=21, ABA) with up to six bivalents for
metaphase I chromosome associations per meiocyte is indicative of genomic exchange among the A
genomes (Table 11.2). The meiotic stability of the AABBAA, 2n=6x=42 amphiploids suggests an ease of
maintenance of these genetic stocks (Table 11.3). The durum cultivars in these amphiploids are
susceptible for the stresses being addressed in the authors’ studies. Hence, upon stress screening, a
resistant amphiploid implies that the particular A-genome accession contributes the expressed
resistance. So far, some diversity has been identified in the AAB-BAA amphiploids for Cochliobolus
sativus, Fusarium graminearum (scab) and leaf rust resistance (Delgado et al., 2001). Diversity is more
extensively observed for Septoria tritici resistance.

FIGURE 11.1

Schematics showing the production of A - and D - genome 2n = 6x = 42 chromosome stocks as a

consequence of hybridizing durum cultivars with A-genome diploid (a) and D - genome diploid (b)
accessions

The D genome (Aegilops tauschii = goat grass; 2n=2x=14, DD)

The A-genome strategy cover the D genome that demonstrates an genetic diversity for stresses as
observed the AABBDD synthetic (MujeebKazi, 2001a). Crossing the resistant synthetic hexaploids with
elite but susceptible bread wheat (BW) cultivars has yield resistant BW/SH derivatives (mujeeb-Kazi et
al., 2001a, 2001b). Currently, 620 SH wheats (Plate 9, Table 11.4) have been produced, and a wide array
of these wheats are being globally utilized for wheat improvement either at the SH or at the BW/SH
advanced derivative level. The most advanced in the authors’ wheat breeding programmes are the D-
genome resistances for Karnal bunt (Tilletia indica), S. tritici and C. sativus. Promise also exists for
resistances and tolerances in this SH germplasm for leaf rust, stripe rust, mineral toxicities, drought,
salinity (Pritchard et al., 2001), heat, cold, sprouting, waterlogging (Villareal et al., 2001), high molecular
weight (HMW)/low molecular weight (LMW) quality subunits, powdery mildew, loose smut, cereal cyst
nematode (CCN), yield and its components. The least accessional diversity observed so far in the D
genome is for scab or F. graminearum (less than 1.0 percent), but under evaluation tests conducted at
one location in Mexico, the observed scab resistance is promising and superior than that of the leading
bread wheat cultivars Frontana and Sumai-3 with their assemblage of four genes (van Ginkel et al.,
1996). No accessional diversity has been observed for aluminiu…

Taxonomic Literature of Plants: An Overview

Taxonomy is fundamentally a descriptive and highly documented science. Its literature is voluminous
and constitutes so vital a part of its structure that, irrespective of whether the problem is one of
identification of an unknown plant, solution of a nomenclatural puzzle, or a monographic or floristic
study, acquaintanceship must be made with the more important publications of the subject.

Every developed and civilized nation has had taxonomists, and this literature may be found in many
languages. There is at present a tendency to write in one of the leading languages such as English,
German, Russian and of course some literature, especially original descriptions, is still written in Latin.
Even so, taxonomy is truly international in its character; hence its studies involve the use of many
different languages.

According to Porter (1967) “Taxonomic literature runs the gamut from ponderous volumes to obscure
notes in periodicals and even letters of correspondence between workers”.

Taxonomic literatures have always occupied a prominent position in botanical literature, the
classification of which is difficult task. An attempt will be made here to supply representative examples
of the subject.

Classics:

The works which have been profoundly influenced the development of plant taxonomy and regarded as
landmarks in the history of Botany are called Classics. They include the works of Theophrastus, Pliny,
Dioscorides, Albertus Magnus, Brunfels, Cesalpino, the Bauhins, Ray, Tournefort and Linnaeus.Although
they exerted a powerful influence in their day, these books are principally of value from the historical
standpoint.

Taxonomic Indexes:

The taxonomic indexes are indexes of plant names and not to literature concerning the plants. Indexes
serve as an aid to locating quickly the source of original publication of a name, to learn if a particular
name has been applied to a plant or to what order, family, subfamily or tribe, a plant of a given name
may belong.

These indexes are the nucleus of any significant taxonomic library, and it is incumbent on the student of
taxonomy to know of their availability and importance e.g. Index Kewensis plantarum phanerogam
arum.

Floras:

A flora is a systematic arrangement of the species of a given area or a particular region, usually
restricted to a major segment of the plant kingdom (flowering plants etc.), with keys and descriptions
and often illustrations, by the use of which a student may determine the names and characteristics of
the wild plants of the area, A flora covers a country, a section of a country, a state, a valley, a desert, or
a vicinity of a city.
The student of taxonomy should be familiar with the floras covering his own region and some rare
important floras covering other regions. In the regional floras, the student finds the available details of
the plant life of his immediate environment.

In any flora the plants are arranged according to one or another of the available systems (Engler, Bessey,
Hutchinson, etc.), giving for each plant the complete scientific name, author citation, reference to
source of original publication, synonymy, and geographic distribution within the area in question.

There is no single world flora that accounts for every species, on the earth. The herbarium material on
which to base, such, a flora does not exist. The so-called world floras (e.g., Bentham and Hooker’s
Genera plantarum, Engler and Prantl’s Die naturalichen Pflanzenfamilien), for the most part, do not
treat units below the category of genus, and some of them do not account for units below the category
of family.

Any one engaged in taxonomic work, is called or frequently to identify plants, and when these are
unknown indigenous plants, they are identified usually by the aid of a manual or flora of the area.
Knowledge of the more important floras is invaluable to the traveler interested in the plants of an area.

Monographs and Revisions:

A monograph is a treatise including all significant information of a morphologic or taxonomic nature

covering the group such as family or genus.

A taxonomic monograph is a comprehensive treatise representing an analysis and synthesis of existing

taxonomic knowledge of that taxon, plus the results of original research of that in systematics. In other
words, it is “as complete an account as can be made at a given time of any one family, tribe, or genus,
‘nothing being neglected which is necessary for a perfect knowledge of it.”

The usual subject of a taxonomic monograph is the genus or the family. All elements of the treatise are
accounted for by dichotomous keys, full synonymies, complete descriptions, precise designations of
types, together with notes as to where the types are deposited, citations of specimens examined,
distributional ranges (supplemented by maps of the same), notes on habitats, and discussions of
taxonomic and nomenclatorial considerations as may be appropriate.

A taxonomic revision differs from a monograph primarily in degree of scope and completeness. Often it
accounts for only a section of a genus or for the elements as restricted to a continent or smaller
geographical area. Many revisions make no attempt to review all previous work on the taxon or to take
cognizance of the interrelated sciences of cytotaxonomy, genetics, ecology, etc.

A revision may be based only on herbarium studies, where as monograph should cover the morphology,
anatomy, cytology, genetics and ecology.

Catalogues:
Catalogues account for the books of special libraries rich in botanical titles, and are of especial value in
taxonomic studies. It is often necessary to know the full name of a particular author, to know the
unabridged and exact title of a work, to know when it was published, or when a particular edition was
issued. These data are usually available from such catalogues.

Review Serials:

Review serials are periodicals, usually issued at regular intervals that provide either:

(1) A bibliography of current literature of a particular subject,

(2) An abstract of papers or books in special fields,

(3) Reviews of titles of current literature, or

(4) Any combination of these functions.

In evaluating their treatments it is well to remember that an abstract is a brief factual summary of a
paper, frequently prepared by its author, whereas a review is an often critical appraisal and evaluation
of the paper, and is by a person other than the original author.

Periodical:

A periodical is a publication appearing usually at regular intervals. Each issue is called a number, or
sometimes is termed a fascicle. Collectively these numbers or fascicles comprise a volume. In the case of
periodicals appearing at regular intervals – biweekly, monthly, or quarterly- a volume usually comprises
the issues of a calendar year.

Scientific periodicals usually are sponsored either by a scientific organization, such as a learned society,
or an educational or non-profit research institution, such as a university or museum. Serial may appear
at regular or irregular intervals.

A society may publish a monthly serial to provide a source of publication for a variety of relatively short
papers contributed by its members, together with records of its own proceedings. Such a periodical is
usually entitled Journal, Anal, Bulletin, or Proceeding.

The same society also may publish at much less frequent intervals another serial to account for longer
and more monumental works, often by a single author, such as monographs of floras. These are often
entitled Memoirs or Transactions. The titles of some periodicals are long, and inciting them it is
customary to abbreviate or condense them. Unfortunately, there is no uniformity of practice.

The number of botanical serials is so great that probably no library contains complete sets of them all.

When a local library does not have a desired serial, or a particular volume of a serial, it is necessary:

(1) To borrow that volume,

(2) To visit a library known to possess it, or

(3) To obtain a photographic copy such as microfilm or Photostat.

Dictionaries and Glossaries:

Dictionaries and glossaries form the nucleus of taxonomic literature. Dictionaries and glossaries are
invaluable in a subject matter with so large a vocabulary as that of Botany. A botanical dictionary may
list and describe all known genera of certain plant groups e.g. A Dictionary of Flowering, Plants and
Ferns by J. C. Willis. A glossary is an alphabetical list of difficult terms with their interpretations.

Almost all modern manuals, and many floras, include a glossary of the botanical terms employed.
Several comprehensive and nearly all-inclusive glossaries have been published as separate works of
definitive character.

Most of the data available in botanical dictionaries are to be found nowhere else, least of all in the usual
unabridged dictionaries of common use. Most botanical dictionaries are of plant names and are sources
for the etymology of Latin or vernacular names, for biographical data of persons for whom plants have
been named, and for vernacular names in various languages.

Legislation:

Since 1867 botanists have met somewhat regularly at the international level and agreed upon legislation
in plant names. These rules and regulations are contained in the International Code of Botanical
Nomenclature (I.C.B.N.). They are subject to revision at each Botanical Congress and a new edition is
then prepared.

Maps and Cartography:

This subject may seem remote or apart from that of the rest of the text; however, the taxonomic
botanist knows the need of:

(1) An understanding of maps, the various projections, and the uses, for which they were designed,

(2) The kinds of maps available as sources of supplementary information, and

(3) Of the availability and sources of maps that treat any area of investigation.

Soil maps and geological maps are equally important in any detailed floristic or monographic research.
These maps show type of soil profile.

Aerial photographs are not strictly classifiable as maps but they may be used in taxonomic research as
valued adjuncts to maps of various types.

Icons:
Icone is the perfect, natural diagram or picture of plant. A picture can convey the mind’s eye a quicker
perfection than words. It gives a clean idea at a instance to non-students of botany who are eagerly
devoted in plant uses e.g. Ayurveda students, general public interested in medical uses of plants, nature
lovers etc. without botanical background.

Icons are full elucidation, namely of the distinctive characters of the natural plants order, as explained in
the descriptive protein of the work. In this the properties and uses of plants, are mentioned as meriting
attention on account of properties, they are known to possess, but of whose forms, the name
communicates no definite ideas, and one can just form a conception of their fitness for the purposes
indicated.

In these sufficient acquaintance with the plants named, that is to be able to recognize these even when
laid before. The non-botanical readers have no other means of acquiring knowledge then through the
oral communication of natives, whose acquaintance with the plants indicated, being entirely traditional
without any guide to direct them always to the same plant is often as likely to be wrong or right.

This is no imaginary statement it is one, the truth of which have seen and verified in a thousand
instances, another, and not the least important lineation, the plants meant by the author, and at the
same time, to establish the Native names, of at least so many of an indigenous plants, on the firm basis,
by combining them with the representation of the objects named.

Such a work still retains an important deaideratum to all classes of community.

In Icones there is the great advantage of pictures in conveying to the mind’s eye a quicker perception
than words can do, of the distinctive peculiarities of an unknown plant. At a single instance the
analytical details of plants apart from general habit are accomplished. The Icones cenvey precise,
definite idea and almost accurate picture while a botanist describe these with a laxity of terms.

In description of plants by professional Botanist the number objects of study are so great, they are, even
in the present advanced state of science, quite indispensible, especially to the young botanist.

In Icones an intimate acquaintance with a few species only of an order, will even a young Botanist
rapidly to acquire a competent knowledge of the rest.

The Icones are indispensable to the student of botany to study the system (systematic) botany, since by
seeing several species of an order arranged together and put in contrast with those of some other order,
student may acquire such an idea of the appearance of a group although he may not be able to explain
it to others, as will made a strong impression on himself and prove eminently useful in advancing his
own researches and in preparing his mind for entering on the more abstract and sub-time parts of the
study.

Icones familiarises student with the appearance of groups of indigenous plants by furnishing correct
figures of numerous species of the order.

Icone acts as a ready reckoner of plant species of that area or region or part or state or country.
Classical Literature:

Linnaeus, C. 1753. Species Planturum, 2 volts. First volume was published in May 1753 and second
volume in August 1753. The system of binomial nomenclature was first adopted in this work and it is
therefore accepted as strating point for application of the ‘Rule of Priority’ in nomenclature of flowering
plants and pteridophytes.

De Candolle, A.P., A and C. 1824-73. Prodromus Systematis Naturalist Regni Vegetabilis. 17 volumes,
Paris, France only dicotyledonous plants.

De Candolle, A& C. de candolle. 1879-91. Monographiae Phanerogamarum. 7 Volumes, Paris, France.

Bentham, G. & J.D. Hookder. 1862-1883. Genera Plantarum, 3 volumes, London, U.K.

The families and genera are arranged according to Bentham and Hooker’s system of classification, in
many important world herbaria, including Kew Herbarium and almost all India herbaria.

Genera Siphanogamarum, 1900-1907, Berlin, Germany.

This book was edited by C.G. Dalla Tore and H. Harms and includes names of families and genera of
spermatophytes. The genera are arranged according to Engler’s system of classification and bear
numbers 1 (Cycas) to 9629 (Thamnoseris). In some herbaria these numbers are used for arranging the
specimens.

Engler A. & K. Prantl 1887-1960. Die Naturlichen Pflanzenfamilien. 23 volumes, Leipzig, Germany.

Englier, A. & L. Diels, 1936. Syllabus der Pflanzenfamilien, 11th ed. berlin, Germany. Some herbaria in
the world, particularly in Europe, are arranged according to this system.

Pflanzeneich:

Engler, A 1900-1937. Das Pflanzenreich. regni Vegetablis Conspectus. Lepzig, Germany. This work has
monographs on many families and genera.

Wallichian Catalogue:

It is a list of 9148 plants collected during Nathaniel Willich’s superintendence of Royal Botanic Gardens,
Calcutta. 1828 to 1829 at East India Company’s Museum in London. It lists not only Wallich’s own
collections, but also plants collected by B.’ Heyne, P. Rusell, W. Roxburgh, R. Wight, etc.