 Hexaploid triticale, including its primary and secondary forms, is an important forage crop and a

promising energy plant. Primary forms are usually developed by crossing Triticum turgidum L.
with rye, with secondary forms obtained by crossing primary hexaploid triticale and/or
hexaploid wheat with octoploid triticale
 Multivalent formation at meiosis is seen in case of autopolyploids and no multivalent formation
in case of allopolyploids
 Segmental allopolyploids may undergo univalent, bivalent and/or multivalent pairing
 The competitive advantage of polyploids over their diploid progenitors is mostly related to
transgressive segregation, i.e., formation of extreme phenotypes, and increased vigor.
 Some changes promoted by polyploidy allowed the polyploid individuals to survive the
environmental changes of the KT boundary (EXTINCTION) and outcompete many of their diploid
progenitors. •These changes included increase in phenotypic variability, heterosis, mutational
robustness, subfunctionalization, and alterations in the reproduction modes

 Genome ‘‘buffering’’ and heterozygosity widely applied in polyploid breeding programs •
Genome redundancy (EXTRA) promotes a ‘‘buffering’’ effect in which the deleterious alleles are
masked by the extra copies of wild-type alleles. Another benefit of genome redundancy is the
possibility of functional diversification of redundant gene copies, in which one member of a
duplicated gene pair mutates and acquires a novel function, without compromising essential
functions • The increment in heterozygosity is another feature that accompanies polyploidy.
 autopolyploids are also expected to have higher levels of heterozygosity compared to their
related diploids due to polysomic inheritance and the possibility of outcrossing occurrence
 Nullisomichas been produced in wheat and otas only
 Monosomicsdo not survive in diploid species •Nullisomiesdo not survive even in some polyploid
species, eg, in tobacco
 •In diploid species, only trisomicssurvive; full series of trisomicsare available in maize, barley,
peas, tomato, bajra, spinach and several other species
 Trisomics vary considerably in vigour; some trisomics may be comparable to the normal diploid,
e.g., Trisomics vary considerably in vigour; some trisomics may be comparable to the normal
diploid, e.g., trisome 5 (that is, trisomic for chromosome 5) in maize, while others are greatly
reduced in vigour
 Logical Features of Aneuploids • During meiosis, nullisomics generally show regular bivalent
formation and chromosome distribution producing gametes with n-1 chromosomes •
Tetrasomics are relatively less regular; they usually form one quadrivalent (involving four
chromosomes) at metaphase I. (which aften separates 2: 2 at anaphase 1. But sometimes the
separation of the four chromosomes of the quadrivalent is not as regular; therefore, all the
gametes produced by a tetrasomic are not n+ 1 ) • In monosomics, one chromosome does not
have a pair and remains as a univalent at metaphase I. At anaphase I, the univalent may move
 one of the two poles, may lag and be lost, or may divide (as in mitosis) into two chromatids,
which move to the opposite poles. Due to the irregular behaviour of the univalent, monosomics
produce an excess of n-1 gametes, e.8., wheat produces -75% n-1 gametes. • In trisomics, the
extra chromosome often forms a trivalent with the other two homologous chromosomes, or
may remain as a univalent. The univalent benaves ifneguiarly: nence the proportion of n +1
gametes is usually less than 50%
  The transmission of aneuploid condition is very poor through pollen grains, primarily due to a
slower tube growth of aneuploid pollen. As a result, normal pollen outcompetes the aneuploid
one; this is called certation.

APOMIXIS

The first discovery of this phenomenon was credited to Leuwenhock as early as 1719 in Citrus seeds.

Most frequently in Compositae, Rosaceae & Graminae.

Apomictic species are usually perennials derived from strongly out crossing ancestors, and apomixis
is usually associated with interspecific hybridization and polyploidy.

Haploid parthenogenesis occurs accidently and has been reported in Nicotiana, Crepis and maize.
Diploid parthenogenesis occurs in many grasses, e.g.. Taraxacum.

In case of Pennisetum, Panicum and Citrus, apomixis is controlled by single gene or locus, apomixis
being dominant in Panicum and Citrus. qHowever, recent studies with Tripsacum indicate that a
single gene may, in fact, comprise several extremely tightly linked genes. qWhile in Pennisetum
ciliare obligate apomixis shows inhibitory gene interaction.

Apomictic lines can be developed by the following three different approaches

(1) Gene transfer from wild species - from Tripsacum dactyloides into maize.

(2) Induced mutations -In sorghum, two mutant lines showing facultative apospory have been
isolated and are being used in hybridization programmes in an effort to develop obligate apomictic
lines .

(3) Selection of apomictic genotypes in the segregating generations of interspecific hybrids.-

apomictic seed formation has been reported in the intergeneric hybrids between T. aestivum and
Avena sativa, H. vulgare and T. aestivum or T. turgidum, etc

Application of APOMIXIS

1) fixation of heterosis

Apomixis (apospory and diplospery) leads of fixation of heterosis since apomictic progeny of hybrids
have the same genotype as the hybrid. Obligate apomixis results in complete fixation of heterosis.

2) production of homozygous lines- Some types of apomixis involve parthenogenetic development of

reduced egg cells, followed by reduplication or fusion, or development of embryos from secondary
diploid cells of the imbryo sac derived through fusions. Such types to apomixis would give rise directly to
homozygous diploids. Such apomixis is claimed to occur in Rubus and has been shown to occur in
sorghum (S. bicolor); in sorghum true-breeding, sexually reproducing progenies were identified in F3 of
the cross 8473 (apomictic line) x sexual lines (3 different lines) and the fusion of haploid nuclei of the
embryo sac has been cytologically verified.
 (3) development of phenotypically stable populations called vybrids.- vybrid is the progeny obtained
from a cross between two facultative apomicts; the vybrid itself reproduces through facultative
apomixis and is maintained by harvesting the seeds of apomictic plants in every generation

Many studies suggest that hybridization, either on diploid or polyploid cytotypes, is a major trigger for
the formation of unreduced female gametophytes, which represents the first step toward apomixis, and
must be combined to parthenogenesis, the development of an unfertilized egg cell. Nevertheless,
fertilization of endosperm is still needed for most apomictic plants. Coupling of these three steps
appears to be a major constraint for shifts to natural apomixis..

CIMMYTmountedaconcertedefforttounderstandandtransferapomictic traitstomaize-
anadvancethatcouldforeverchangethedevelopmentanduseof
improvedvarietiesinsomeoftheworld’sneediestregions.

TheCIMMYT ApomixisProjectwaslaunchedin1989withthegoaloftransferringthenaturallyoccurring,
untappedtraitofapomixistomaize,anachievementthatcoulddramaticallyanddirectlyimprovefarmer
productivityinthedevelopingworld.

In addition, mobile DNA sequences known as transposons, which induce mutation when inserted into
genes, have been introduced into apomictic materials. The aim is to identify and isolate the individual
genes associated with apomictic control.

T.dactyloide and T.zopilotense , this two species showed high cross abilities with maize

This long-term strategy has provided very encouraging results. It was found that all Tripsacum
chromosomes can be transferred into addition lines and that about 10% of these plants also show
maize-Tripsacum translocations, suggesting that 20- chromosome recovered maize plants with small
Tripsacum DNA segments can be produced from the backcross series.

Moleculargeneticsstudieshaveledtotheidentificationofaverypromising”candidate”gene
involvedinthecontrolofapomixis.Thisgene(calledelongateinmaize)isnowbeingclonedand
experimentswillsoonbeunderwaytoverifyitsfunction.
Asecondstrategy,addedin1993,isbasedonidentifyingandisolatingtheapomixisgenesin
Tripsacumandtransferringthemdirectlytomaize.Thiscomplexapproachreliesheavilyon
biotechnologyandrequiresextensivegeneticmapping,theisolationof”candidategenes”from
maizeandtheircounterpartsfromTripsacum,andthesuccessfulsplicingofthegenesintomaize.
Ifsuccessful,thiseffortmayyieldresultsmorequicklythanthebackcrossingapproach,with
technologicalimplicationsextendingfarpastmaize.( Last paragraph is important)

1st unit

1.On the extreme, round worm shows only two chromosomes, while the other extreme is represented
by Protozoa having 300 or more chromosomes.

The giant chromosomes found in diptera and they may be as long as 300 µ and up to 10 µ in diameter.

The size of the chromosomes in mitotic phase of animal and plants sp generally varies between 0.5 µ
and 32 µ in length, and between 0.2 µ and 3.0 µ in diameter. The longest metaphase chromosomes
found in Trillium32 µ. The giant chromosomes found in diptera and they may be as long as 300 µ and up
to 10 µ in diameter. In general, plants have longer chromosomes than animal and species having lower
chromosome numbers have long chromosomes than those having higher chromosome numbers Among
plants, dicots in general, have a higher number of chromosome than monocots. Chromosomes are
longer in monocot than dicots.

• All house mouse chromosomes are telocentric, while human chromosomes include both metacentric
and acrocentric, but no telocentric.

For example, darker bands are generally found near the centromeres or on the ends (telomeres) of the
chromosome, while other regions do not stain as strongly. • The position of the dark-staining are
heterochromatic region or heterochromatin. • Light staining are euchromatic region or euchromatin.

• In E.coli, about 3000 to 4000 genes are organized into its one circular chromosome. • The
chromosome exists as a highly folded and coiled structure dispersed throughout the cell. • The folded
nature of chromosome is due to the incorporation of RNA with DNA. • There are about 50 loops in the
chromosome of E.coli. • These loops are highly twisted or supercoiled structure with about four million
nucleotide pairs. • Its molecular weight is about 2.8 X109 • During replication of DNA, the coiling must
be relaxed. • DNA gyrase is necessary for the unwinding the coils.

Negative supercoiling or left hand side coiling of the DNA is the most common

There are many supercoiled loops (~100 in E. coli) attached to a central core. Each loop can be
independently relaxed or condensed. Topoisomerase enzyme– (Type I and II) that introduce or remove
supercoiling.

• The complexes between eukaryotic DNA and proteins are called Chromatin, which typically contains
about twice as muchprotein as DNA. • The major proteins of chromatin are the histones– small proteins
containing a high proportion of basic aminoacids (arginine and lysine) that facilitate binding negatively
charged DNA molecule . • There are 5 major types of histones: H1, H2A, H2B, H3, and H4– which are
very similar among different sp of eukaryotes.

Thus it is generally accepted that structural integrity and individuality of chromosomes is maintained
due to telomeres.