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Haploids in Genetic and Cytogenetical Research

S. Georgiev

To cite this article: S. Georgiev (2008) Haploids in Genetic and Cytogenetical

Research, Biotechnology & Biotechnological Equipment, 22:2, 644-651, DOI:
10.1080/13102818.2008.10817528

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 REVIEW
HAPLOIDS IN GENETIC AND CYTOGENETICAL RESEARCH
S. Georgiev
Sofia University, Faculty of Biology, Department of Genetics, Sofia, Bulgaria
Correspondence to: Sevdalin Georgiev
E-mail: georgievs@biofac.uni-sofia.bg
ABSTRACT
The aim of this review is to summarize our current understanding of the meiotic behaviour of chromosomes in haploids and
polyhaploids and to try to elucidate some aspects of chromosome pairing in haplo- and polyhaploids during meiotic division.
Keywords: euhaploids, aneuhaploids, haploidy, monoploids,
Zea mays, Hordeum vulgare, nulisomic haploids, disomic
haploids
Introduction
Since the first report of haploidy in Datura stramonium
(2), this phenomenon has been found among numerous
other plant species. A large number of haploids have been
produced in cereal crops including wheat (22, 23, 24, 25,
28, 47, 48,). Spontaneously and induced haploidy were
reported in several animal species, including Drosophila,
salamander, frog, mouse and chicken. Usually haploidy in
animals produces physiologically abnormal individuals that
die during embryogenesis. In certain insects (Hymenoptera:
honeybees, wasps, etc.) haploid is part of the sex-determination
system. Haploids with gametic chromosome complements,
offer excellent opportunities for the investigation of many
fundamental problems concerned with genetics, cytogenetics
and genomics. The study of haploids has helped considerably
in our understanding of the cytogenetic structure and evolution
of several plant species (25).
Doubled haploids (DH, plants or animals with chromosome
doubling of haploids) also are particularly useful in the genetic
improvement of trees which have very long generation cycles
and hence production of homozygous lines by continuous
inbreeding in them. The DH-derived homozygous populations
could be usefully in genetic studies, particularly in elucidating
the genetic control of traits by recessive alleles and analysis
Fig. 1. A-C. Haploid cells of Zea mays L. (56)
644
A&EB
of quantitative trait loci (7, 17). The DH are completely
homozygous at every locus, since haploids have only one allele
per locus, they are useful for linkage mapping and studies of
the genetical control of chromosome pairing.
Despite the great importance of haploids, no book is
currently available that gives an adequate coverage to basic
aspects and application of haploidy in various facets of
biological research.
The purpose of this article is to review mainly the recent
literature on haploids and to discuss their possible use in
genetic and cytogenetical research.
Satisfactory classification of haploids for the first time
has been suggested by Kimber and Riley (31). According
this classification haploids are divided into two broad groups,
euhaploids and aneuhaploids. A euhaploid may possess either
the basic chromosome number itself or an exact multiple
of it, depending upon the ploidy level of the species from
which it is derived. If a plant with the gametic chromosome
number is produced from a diploid species it may be known
either as a monoploid or simply as a haploid (Fig. 1A-C).
On the other hand if it is derived from a polyploid species
the term polyhaploid is used. The cytogenetic constitution
of a polyhaploid depends upon the nature of the polyploid
species from which it originates. Therefore it is important to
distinguish between polyhaploids derived from autopolyploids
species (autopolyhaploids) and those from allopolyploid
species (allopolyhaploids). Unlike euhaploids, aneuhaploids
possess chromosome numbers that differ from the haploid
BIOTECHNOL. & BIOTECHNOL. EQ. 22/2008/2
number by one or more chromosomes. They possessed extra
chromosomes or may be deficient for chromosomes, relative
to the euhaploid condition (Fig. 2).
Furthermore the extra chromosomes may be members of
the gametic set of the species (disomic haploids) or they may be
alien (from a different species or genus) – additional haploids.
Haploids deficient for one chromosome of the normal gametic
complement of the species are known as nullisomic haploids
and those having one or more alien chromosomes substituted
for one or more chromosomes of the haploid complement are
known as substitution haploids. Misdivision haploids possess
telocentric or misdivision derivatives (isochromosomes) of
some chromosomes.
Haploids and genetic analyses
Haploids (haploid from the Greek word for “single”) and their
homozygous diploid derivatives represent perfect materials
for the study of many genetic problems, but the successful
utilization of haploids in genetic research depends not only
the efficient production of large numbers of haploids but also
on the availability of reliable techniques for the production of
chromosomally doubled and genetically stable derivatives. The
monoploid plant sporophyte provides a unique opportunity to
study the expression of a single allele and its dosage effect
on plant physiology and morphogenesis. Lindstrom and Koos
(34) showed that the recessive gene (p) for pubescent fruit
Fig. 2. Classification of Haploids (51)
BIOTECHNOL. & BIOTECHNOL. EQ. 22/2008/2
character was incapable to produce any phenotypic effect in a
single dose. In contrast Subrahmanyan and Kasha (53) found
that in barley the eight recessive marker genes were expressed
in monoploid progeny.
For the study of gene dosage effects with incomplete
dominance autopolyhaploids and their somatically induced
polyploids also offer good possibilities. For example assuming
no locus alteration during evolution of a parental autotetraploid,
autopolyhaploid progeny with genotypes AA, Aa and aa and
their somatically induced tetraploids (AAAA, AAaa, aaaa)
would allow for critical analysis of the dosage effect.
Genetic analysis in autotetraploids is more complex than in
diploids since they exhibit tetrasomic inheritance (6, 19). Their
autopolyhaploid derivatives may thus be expected to show
disomic inheritance (25). Thus the use of autopolyhaploids
may simplify the complexities involved in the genetic analysis
of autotetraploid species. The inherent genetic variability
of autotetraploid species after inbreeding comes to express
much slower rate that of diploid species (44). Thus their
autopolyhaploid derivatives which are at the diploid level,
may offer a faster means of testing and utilizing the genetic
variability in autotetraploid species. Such information would
be of special interest for plant breeding programms.
Haploidy provides an opportunity to study the effect of
different levels of ploidy on vigour and productivity. Such
studies will be useful in determining the optimum level of
645
ploidy for the maximum expression of desired characteristics.
The fact that triploid sugar beets yield more sugar per acre than
diploids or tetraploids is an example of an application of such
knowledge in plant breeding research (5).
In monopliods, since each chromosome is represented only
once, the occurrence of mutations can be readily detected.
Thus the study of monoploids and homozygous diploids
derived from them may be useful in determining the rates
of spontaneous and induced mutations. Studies on haploids
and their diploid derivatives have led to the detection of
genetic factors controlling phenomena such as apomixis,
meiotic behaviour, pollen production and fertility (35). In a
vegetatively propagated monoploid cultivar of Pelargonium
(n=9), Daker (13) observed a low frequency of bivalents and a
preponderance of neocentric univalents. However, in contrast
with expectations, microsporogenesis in the colchicine doubled
haploids was very erratic and pairing was never complete.
Dyads were the major products which were observed in
meiosis.
Monoploids have been found suitable for studding the
influence of alien cytoplasm on meiotic behaviour and the
expression of both qualitative and quantitative characters.
Sadasivaiah and Kasha (50), comparing Hordeum vulgare
monoploids carrying vulgare and H. bulbosum cytoplasm,
respectively, observed no difference in meiotic behaviour,
but found detectable effects of bulbosum cytoplasm on the
expression of morphological traits. Plants with bulbosum
cytoplasm tended to be more prostrate, shorter, later heading and
maturing, lower yielding and more sensitive to environmental
stress than those with vulgare cytoplasm.
The homozygous diploid derivatives of haploids can be
used as genetic constants in many types of laboratory and field
experiments to measure the variation due to environmental
effects (35).
The great numbers of discovery that haploids could be
produced at will, in many crop plants (22, 23, 24, 25, 26, 27, 28,
42, 47, 48, 54,), has determined the interest in the utilization of
haploids in various genetic studies. However, despite the rapid
progress achieved in the production of haploids, their actual
use in genetic research has not generally proceeded beyond
the determination of simple genetic ratios (4, 9, 40). However,
Collins, Legg and Kasperbauer (10), from an extensive study
of alkaloid content of several allopolyhaploids and tetraploid
forms in tobacco (Nicotiana tabacum), have demonstrated the
suitability of allopolyhaploids for the study of genetic control
of plant chemistry.
With improving the techniques of isolation and induced
fusion of plant protoplasts have opened new avenues for
genetic analysis and the study of genetic regulation. Fusion
of plant protoplasts as a means of circumventing the sexual
cycle to produce interspecific and intergeneric hybrids, will be
of great interest in the study of phylogenetic relationships and
gene transfer between widely divergent groups of plant species.
Isolated haploid protoplasts and their regeneration into entire
646
plants offer possibilities for improved and highly efficient
methods for the induction and selection of mutations and thus
for the analysis of genetic fine structure in eucaryotes.
The phenomenon of transgenosis (gene transfer) provides
new possibilities for investigating various problems concerned
with prokaryote-eukaryote gene interaction and control
mechanisms involved in their expression (28). More studies
in this area may provide a powerful tool to plant breeders for
transferring defined genetic material into crop plants.
Polyhaploids also offers an excellent example to investigate
the evolution in many plants, thus they can be used as a tool to
elucidate the mechanism of evolution in cereals (25).
From the above discussion it is clear that the potentialities
of haplods as tools in genetic research are numerous.
Haploids in Cytogenetical Studies
Mitotic Studies
Chromosome numbers have been studied in haploids and
polyhaploids of a number of species (13, 20, 23, 29, 33,
37, 49, 56). These studies are important for two reasons:
i) in the first place, since each homologue is represented
only once in haploids, the task of matching homologues in
karyotype analysis is eliminated. This is especially true of
polyhaploids, where the lower chromosome number provides
an added advantage in karyotype studies and ii) secondly, the
identification of chromosomes, especially in aneuhaploids,
will be extremely useful in determining the genetic influence
exerted by chromosomes on the meiotic behaviour of the
individuals concerned.
Apart from the study of somatic chromosomes, the study of
the mitotic cycle in haploids are of considerable interest with
regard to possible genomic effects – Genome size and genome
structure. It is well known that in higher plants interphase
nuclear volume and minimum mitotic cycle time are linearly
correlated (58).
Meiotic Studies
The study of chromosome pairing in monoploids is important
since in this case each chromosome usually lacks its normal
homologous partner and pairing affinities.
Monoploids of a number of species have been reported in
the literature (27, 31, 32, 35). However, in most of the earlier
monoploid studies, observation on meiotic chromosome
behaviour are largely restricted to diakinesis and later stages
of meiosis, and very little attention has been given to the study
of prophase pairing. In this regard, a generalized description
of monoploid meiosis based on results obtained from recent
studies may be helpful towards a clearer interpretation of the
various chromosome configurations observed in monoploid
meiosis.
The earliest meiotic stage at which the pairing properties
of non-homologous chromosomes may be critically studied in
monoploids is pachytene.
BIOTECHNOL. & BIOTECHNOL. EQ. 22/2008/2
 Pachytene (from the Greek for “thick”) begins at the
completion of synapsis when homologous chromosomes are
united along their length. The pairing properties between non-
homologous chromosomes at this stage are not the same in
monoploids derived from different plant species. In rice Chu
(8) observed an occasional pairing between parts of some non-
homologous chromosomes. Pairing within single univalent
(foldback pairing) has also been observed in monoploids of
maize a (15, 57, 59). In barley monioploids derived from
interspecific crosses between H. bulbosum (2n=14) and H.
vulgare (2n=14), Symko (55), observed a completed lack of
pairing at pachytene. In contrast Sadasivaiah and Kasha, (50),
observed a very high degree of non-homologous pairing in two
types of barley monoploids, obtained from reciprocal crosses
between H. vulgare and H. bulbosum (Figs. 3-5).
Figs. 3-5. Meiosis in barley haploids. Mid-pachytene stages showing non-
homologous pairing. (Reproduced from 50).
From the above discussion it is clear that the obtained
results are rather controversial, probably due to a failure to
achieve a good spread of pachytene chromosomes required
for an accurate analysis. It is important to note that in many
haploid plants (intergeneric hybrids of grasses) there is a very
little chromosome pairing (61). It is possible that lack of pairing
may be a prerequisite for the occurrence of meiotic restitution
and hence chromosome doubling (24).
The introduction of electron microscopy and new
fluorescent techniques to biological research has opened a new
approach to studies of pairing. The synaptonemal complex, a
structure found when homologous chromosomes are involved
in meiotic pairing, has been reported in monoploids of tomato
(38), maize (57), wheat (24). These synaptonemal complexes
BIOTECHNOL. & BIOTECHNOL. EQ. 22/2008/2
were similar in nature to those formed in their corresponding
diploid species.
Diplotene (from the Greek for “twofold” or “double”) is
signalled by the gradual dissolution of the synaptonemal zipper
complex and a slight separation of regions of the homologous
chromosomes. This is ideal stage for the study of chiasmata, if
these exist between non-homologously paired chromosomes.
In majority of haploids reported, practically no attention has
been given to this stage. However, in barley monoploids,
Sadasivaiah and Kasha (49) observed up to four chiasma-
like configurations in some bivalent-like structures found at
diplotene (Figs. 6 and 7). The occasional association of more
than two chromosomes was also observed at this stage.
Figs. 6-11. Meiosis in barley haploids. Diplotene and Diakinesis. Fig. 10.
Diakinesis with two bivalent-like and one trivalent. Fig. 11. Diakinesis with
one ring univalent-like, one bivalente-like and one quadrivalent-like structures.
(Reproduced from 50).
Diakinesis (from Greek for “double movement”) is
accompanied by further condensation of the chromatids.
At this stage chromosomes occur mostly as univalents (56).
However, associations of two or more, resulting in bivalent-
like and multivalent-like configurations have been observed in
many monoploids (51). Sadasivaiah and Kasha (50) observed
univalents and bivalent-trivalent- and quadrivalent-like
configurations in barley (Figs. 8-11).
647
 Metaphase I. The most striking phenomena observed at this
stage are the lack of metakinesis and the disorganized spindle.
Chromosomes occur mostly as univalents and lie scattered in
the cell. As a result, only a very low frequency of bivalents
(mostly rod types) and some very rare higher associations
have been recorded at this stage (Figs. 12, 13). Some times the
observed bivalents were pseudobivalents formed as a result of
random pairing between chromosomes (36).
Figs. 12-17. Meiosis inbarlez haploids. Fig. 12. MI with five univalents and
one bivalent-like structure. Fig. 13. MI with one trivalent configuration. Fig.
14. MI showing secondary associations of univalents. (side-to-side) Fig. 15.
AI. (the division of univalents). Figs. 16, 17. AI showing univalent division
and chromatidbridge. (Reproduced from 50)
Metaphase I has been studied extensively in the majority of
monoploids (25, 31, 35, 37). In monoploids of barley, Symko
(1969) observed a complete absence of pairing. However,
Sadasivaiah and Kasha (50) found only a single bivalent-
like structure in barley haploids (Fig. 12). A similar low
frequency of bivalent-like structures has also been observed
in monoploids of N. otophora (11), in tomato (14), Linum
648
africanum and Ricinus communis (62). In addition to bivalents,
one trivalent like structure was also observed rarely in some
cells of monoploid barley (Fig. 13).
Besides taking part in bivalent and trivalent like
configurations, univalents at metaphase I may participate in
so called secondary associations like side-by-side (s-s), end-
to-end (e-e), end-to-side (e-s), multiple s-s, multiple e-e,
multiple associations involving combinations of s-s, e-e and
e-s associations. These associations can be clearly identified
where univalents are oval or rod shaped, as in wheat, barley,
oats (Fig. 14). In species where the univalents are more or less
round, classification of associations into s-s, e-e, or e-s may
not be possible.
Anaphase I. In diploids homologous chromosomes move
to opposite poles of the spindle.
In many monoploids the absence of metaphase I plates
and the distribution of univalents to the poles make it difficult
to distinguish from the anaphase I (31, 11). Hence the term
“meta-anaphase” has been used in such cases to denote this
period of meiotic division. On the other hand, in monoploids
of certain species such as H. vulgare, it is possible to make a
clear distinction between metaphase I and anaphase I stages
(Fig. 15). The segregation of chromosomes at anaphase
I is usually random (36, 50). FISH analyses (24, 56) also
showed the random movement of the A- and B-genome
chromosomes of T. durum and maize, or in extreme cases,
all chromosomes moved to one pole, which resulted in non
reduction of chromosome number at the first meiotic division.
In spontaneous haploids of pearl millet, Jauhar (21) and Powell
et al. (45) observed that all seven univalents moved to one
pole. This phenomenon has essentially same consequence as
the FDR (first division restitution). When all univalents move
to one pole, they, in essence, bypass the reductional division of
meiosis and than, on equational division, produce unreduced
gametes. The phenomena described above could have breeding
implications. In this case the diploid chromosome complement
is first restored and than the diploid complement undergoes
mitotic (equational second meiotic division) division resulting
in unreduced gametes. During the equational division, sister
chromatids of each chromosome (univalents of haploids) move
to opposite poles, and therefore the two resultant nuclei are
essentially similar to each other or doubled haploids (DH) arisen
(7, 17). The instant homozygosity derived through chromosome
doubling of haploids can accelerate breeding programs (1,
30). However, the mechanism involved in the movement of
univalents is not well understood. Lagging of varying numbers
of univalents (Fig. 16) and division of univalents have been
commonly observed in monoploids of a number of species (11,
15, 24, 25, 37). In addition, a small percentage of chromatid
bridges with or without acentric fragments (Fig. 17) have been
observed in haploids of barley and maize (59). Chromatid
bridges and acentric fragments could be formed as a result of:
i) recombination within duplicated segments present in reverse
order with respect to the centromere, ii) U-type exchange and
BIOTECHNOL. & BIOTECHNOL. EQ. 22/2008/2
iii) recombination in univalent foldbacks, if the centromere
divides at anaphase I.
Telophase I. takes place when nuclear membranes begin to
form around the chromosomes that have moved to the poles. In
many haploids, varying numbers and sizes of micronuclei are
observed at telophase I (11, 25).
Second Meiotic Divisions. The second metaphase division
are usually normal, with a normal spindle, although some
irregularities have been reported in some haploids. Irregular
distributions of chromosomes, laggards (Fig. 18) and chromatid
bridges (usually without fragments; Fig. 19) are also observed
at anaphase II. Bridges at anaphase II may be formed as a result
of failure of chromatids to separate at anaphase I. At the tetrad
stage, varying frequencies of dyads, triads, and monads, with
micronuclei which again vary in size and frequency have been
observed in many haploids (24, 25, 37, 50).
Figs. 18 and 19. Meiosis in barley haploids. Late AII with laggards and
chromatid bridge. (Reproduced from 50)
Monoploids are usually sterile and reports of seedset on
haploids are extremely rare. However, some seed set has been
observed after open pollination or cross pollination using the
diploid parental species as male parent (24, 59).
Some aspects of Non-homologous Chromosome
Association in Haploids
The pairing of homologous chromosomes is essential not
only for meiotic chromosomal recombination, but also for the
faithful segregation of homologous chromosomes (12, 18,
41).
Pachyten Pairing. Haploids derived from different plant
species vary with regard to the degree of non-homologous
pairing at pachytene. A high degree of intimate pairing
between non-homologues chromosomes has been observed in
haploids of H. vulgare (50), while such pairing is almost absent
in haploids of N. otophora (11). However, types intermediate
between the two extremes are also known in the literature (8,
11). Although the cause for this differential behaviour is by no
means clear, some possible explanations may be considered.
One such possibility is that segmental duplications provide a
basis for non-homologous association, and that the extent of
duplication determines the amount of pairing. It should be
point out that such an interpretation may seems valid in cases
where extensive genetic studies have provided evidence of
the existence of gene duplications. However, according to this
explanation, a very extensive duplication, both between and
BIOTECHNOL. & BIOTECHNOL. EQ. 22/2008/2
within the chromosomes of the genome, would be necessary to
explain the high degree of intimate pairing observed in haploid
barley (50). Similarly, a high degree of interchromosomal
duplication is also required to account for pairing observed
between non-homologues in haploids. If duplicated segments
exist within a genome, then preferential pairing should lead to
a non-random association of non-homologous chromosomes.
However the presence of foldback pairing and occasional
heteromorphic bivalents and multivalents in barely haploids
may suggest that pairing is random.
A second possibility is that postulated by Rieger (46), who
has suggested that all chromosomes in general have a tendency
to pair in meiotic prophase. In case where some homologous
segments exist in a set of non-homologous chromosomes,
preferential pairing may occur between such identical
segments, but does not suppress non-homologous pairing.
However it appears that the Rieger’s hypothesis does not
satisfactorily explain the partial to almost complete absence of
pairing between non-homologues in some haploids (8, 11).
A third possibility is distributive pairing, a hypothesis
advanced by Grell, (16) to account for the non-random
disjunction of non-homologously paired univalents in
Drosophila melanogaster females. Distributive pairing is
thought to occur subsequent to exchange pairing and between
chromosomes that have failed to become involved in pairing
that results in exchange between homologues. According
to Grell (16) the similarity of chromosomal size rather than
homology between chromosomes is the determining factor for
distributive pairing. Now on the basis of the occurrence of ring
univalents and heteromorphic bivalents as well as multivalent
associations at diplotene and diakinesis, the distributive
hypothesis is not so important.
A fourth possibility is the involvement of heterochromatin
as a causative factor in non-homologous pairing (49). Recent
biochemical evidence in situ and molecular hybridization
shows that the DNA of eukaryotic organisms possessed
various classes of repetitive nucleotide sequences (3) and some
of these repetitive DNAs are localized in the heterochromatic
regions of chromosomes (43). In view of this, the possibility
of heterochromatic attraction, being involved in the non-
homologous associations observed in haploids cannot be
excluded.
A further possibility, that differential non-homologous
pairing observed in haploids may be under the control of genetic
factors. According Martinez et al. (37) there were differences in
the meiotic behaviour of the haploids from different cultivars.
Variation in the effectiveness among cultivars of wheat is
proposed to have a genetic origin and the role of the Ph1 locus
in different haploids is discussed (24).
According Jauhar et al. (23, 25). Haploids without Ph1
substitution line from Langdon 5D/5B, showed extensive
pairing, unlike of those with Ph1. The Ph1-haploids showed
very little pairing which was limited mostly to intergenomic
649
rod bivalent formation between the chromosomes of the A and
B genomes.
At present, alternative ideas are developing. Attractive is
the concept that double strand DNA repair mechanisms that
find and use the undamaged homologue for repair have evolved
into a meiotic mechanism for the recognition and pairing of
homologous sequences (39).
Now more attractive and acceptable is the hypothesis
which concerns the structure of eucharistic chromosomes. The
ends of the chromosomes, telomeres become tightly clustered
together on the inner surface of the nuclear envelope, resulting
in a structure resembling a bouquet of flowers. The close
relationship between the telomere clustering and chromosome
pairing has led to the suggestion that the bouquet may help to
facilitate the pairing. Consistent with this potential role is the
general observation that synapsis is typically initiated near the
telomeres. According Wojciech et al. (60) mutants defective in
bouquet formation show delay in the progression of meiosis
and severe defects in pairing and synapsis. Evidence that this
telomere clustering is important for homologous chromosome
clustering is important for homologous chromosome pairing in
S. pombe has been obtained from several recent studies (52).
In the microsporocytes of a haploid of T. monococcum (x=7),
foldback and other nonhomologous pairing was observed
at pachytene. At the diplotene equivalent stage of meiosis,
nonhomologous chromosomes were connected by their
telomeres in associations involving two to seven chromosomes.
According Jackson et al. (20) these connections may have
resulted from earlier base-pairing of repeated sequences
of guanine-rich telomere overhangs of non-homologous
chromosomes. Recent molecular studies of several widely
divergent organisms have shown that all telomeres of non-
homologous chromosomes in a genome are identical and
telomere structure is conserved among widely divergent
eukaryotes.
For allopolyploids to produce viable gametes and be
fertile, they must behave as diploids during meiosis, so that
only identical chromosomes (homologues) pair. A solution
to this problem is an enhanced ability to resolve incorrect
pairing, which in turn promotes correct pairing. This gives
non-homologous chromosomes an almost “Teflon”-like status,
so that only the correct pairs “stick”.
Conclusions
Haploids, as a consequence of their unique genomic constitution,
offer improved means of investigating many fundamental
problems in genetics, cytogenetics and genomic. Mutations,
gene-cytoplasmic and gene-environmental interactions can be
more readily detected and studied, without interference from
factors such as heterozygosity. In addition, critical information
regarding chromosomal homologies, chromosomal and genome
evolution, chromosomal and gene dosage effects, the basis of
meiotic pairing and other genetically controlled phenomena
may be obtained from the study of haploids. Haploidy has
special suitability for the detection of genetic regulatory systems
650
of an operon-like nature that may exist in higher organisms,
and in studies of genetic fine structure, genetic transformation
(transgenosis) and nucleus – cytoplasmic organelle interactions
in higher plants.
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