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Subgenome integrity in bread wheat (Triticum aestivum; BBAADD) makes possible the extraction of its BBAA component to restitute a novel plant type. The availability of such a ploidy-reversed wheat (extracted tetraploid wheat [ETW]) provides a unique opportunity to address whether and to what extent the BBAA component of bread wheat has been modified in phenotype, karyotype, and gene expression during its evolutionary history at the allohexaploid level. We report here that ETW was anomalous in multiple phenotypic traits but maintained a stable karyotype. Microarray-based transcriptome profiling identified a large number of differentially expressed genes between ETW and natural tetraploid wheat (Triticum turgidum), and the ETW-downregulated genes were enriched for distinct Gene Ontology categories. Quantitative RT-PCR analysis showed that gene expression differences between ETW and a set of diverse durum wheat (T. turgidum subsp durum) cultivars were distinct from those characterizing tetraploid cultivars per se. Pyrosequencing revealed that the expression alterations may occur to either only one or both of the B and A homoeolog transcripts in ETW. A majority of the genes showed additive expression in a resynthesized allohexaploid wheat. Analysis of a synthetic allohexaploid wheat and diverse bread wheat cultivars revealed the rapid occurrence of expression changes to the BBAA subgenomes subsequent to allohexaploidization and their evolutionary persistence.

The Ph1 gene has long been considered the main factor responsible for the diploid-like meiotic behavior of polyploid wheat. This dominant gene, located on the long arm of chromosome 5B (5BL), suppresses pairing of homoeologous chromosomes in polyploid wheat and in their hybrids with related species. Here we report on the discovery of genotypic variation among tetraploid wheats in the control of homoeologous pairing. Compared with the level of homoeologous pairing in hybrids between Aegilops peregrina and the bread wheat cultivar Chinese Spring (CS), significantly higher levels of homoeologous pairing were obtained in hybrids between Ae. peregrina and CS substitution lines in which chromosome 5B of CS was replaced by either 5B of Triticum turgidum ssp. dicoccoides line 09 (TTD09) or 5G of Triticum timopheevii ssp. timopheevii line 01 (TIM01). Similarly, a higher level of homoeologous pairing was found in the hybrid between Ae. peregrina and a substitution line of CS in which chromosome arm 5BL of line TTD140 substituted for 5BL of CS. It appears that the observed effect on the level of pairing is exerted by chromosome arm 5BL of T. turgidum ssp. dicoccoides, most probably by an allele of Ph1. Searching for variation in the control of homoeologous pairing among lines of wild tetraploid wheat, either T. turgidum ssp. dicoccoides or T. timopheevii ssp. armeniacum, showed that hybrids between Ae. peregrina and lines of these two wild wheats exhibited three different levels of homoeologous pairing: low, low intermediate, and high intermediate. The low-intermediate and high-intermediate genotypes may possess weak alleles of Ph1. The three different T. turgidum ssp. dicoccoides pairing genotypes were collected from different geographical regions in Israel, indicating that this trait may have an adaptive value. The availability of allelic variation at the Ph1 locus may facilitate the mapping, tagging, and eventually the isolation of this important gene.Key words: diploid-like meiotic behavior, genetic control of pairing, Ph1 gene, Triticum turgidum ssp. dicoccoides, wild emmer.

Interspecific or intergeneric hybridization, followed by chromosome doubling, can lead to the formation of new allopolyploid species. Recent studies indicate that allopolyploid formation is associated with genetic and epigenetic changes, although little is known about the type of changes that occur, how rapidly they occur, and the type of sequences involved. To address these matters, we have surveyed F1 hybrids between diploid species from the wheat (Aegilops and Triticum) group and their derived allotetraploids by screening a large number of loci using amplified fragment length polymorphism and DNA gel blot analysis and by assaying the extent of cytosine methylation. We found that sequence elimination is one of the major and immediate responses of the wheat genome to wide hybridization or allopolyploidy, that it affects a large fraction of the genome, and that it is reproducible. In one cross between AE: sharonensis x AE: umbellulata, 14% of the loci from AE: sharonensis were eliminated compared with only 0.5% from AE: umbellulata, with most changes occurring in the F1 hybrid. In contrast, crosses between AE: longissima x T. urartu showed that sequence elimination was more frequent after chromosome doubling. Alterations in cytosine methylation occurred in approximately 13% of the loci, either in the F1 hybrid or in the allopolyploid. For eight of nine bands that were isolated, the sequences that underwent elimination corresponded to low-copy DNA, whereas alterations in methylation patterns affected both repetitive DNA sequences, such as retrotransposons, and low-copy DNA in approximately equal proportions.

Wild emmer wheat, Triticum turgidum ssp. dicoccoides, the progenitor of most domesticated wheats, was discovered by Aaron Aaronsohn in 1906 in the Levant. The contribution of this discovery to the study of the cytogenetic structure and evolution of the wheats, to an understanding of the processes and site of their domestication and evolution under cultivation, and to the improvement of domesticated wheat is discussed.

An Israeli accession (TTD140) of wild emmer, Triticum turgidum var. dicoccoides, was found resistant to several races of powdery mildew. Inoculation of the chromosome-arm substitution lines (CASLs) of TTD140, in the background of the Israeli common wheat cultivar ‘Bethlehem’ (BL), with five isolates of powdery mildew revealed that only the line carrying the short arm of chromosome 2B of wild emmer (CASL 2BS) exhibited complete resistance to four of the five isolates. To map and tag the powdery mildew resistance gene, 41 recombinant substitution lines, derived from a cross between BL and CASL 2BS, were used to construct a linkage map at the gene region. The map, which encompasses 69.5 cM of the distal region of chromosome arm 2BS, contains six RFLP markers, a morphological marker (glaucousness inhibitor, W1 I), and the powdery mildew resistance gene. Segregation ratios for resistance in F2 of BL × CASL 2BS and in the recombinant lines, combined with the susceptability of F1 progeny to all tested isolates, indicate that resistance is controlled by a single recessive allele. This alleleco-segregated with a polymorphic locus detected by the DNA marker Xwg516, 49.4 cM from the terminal marker Xcdo456. The new powdery mildew resistance gene was designated Pm26.

RFLP diversity in the nuclear genome was estimated within and among Israeli populations of wild emmer wheat (Triticum turgidumvar.dicoccoides) from a long-term study site at Ammiad (NE Israel), and from several other geographical locations. Using 55 enzyme-probe combinations, high levels of genetic diversity were revealed in wild emmer in general and within the Ammiad site. In spite of high diversity, observed heterozygosity was low and populations consisted of a patchwork of alternate multilocus homozygotes, consistent with the reproductive biology of a predominant self-fertilizing species. Retention of genetic diversity in wild emmer may be promoted by large population sizes, microhabitat diversity, and occasional gene flow through both pollen and seed. Population genetic structure in wild emmer appears to have been influenced by historical founder events as well as selective factors. Multivariate analyses indicated that individuals tend to cluster together according to their population of origin, and that there is little geographical differentiation among populations. Sampling of 12 domesticated land-races and both primitive and modern cultivars ofT. turgidumrevealed high levels of diversity and a large number of alleles that were not detected in the wild emmer populations. This may reflect a long-term domestication process in which wild, semi-domesticated, and domesticated types grew sympatrically, continuing introgression from wild populations, and perhaps also gene flow from trans-specific sources.

The Rbc S multigene family in hexaploid wheat, Triticum aestivum , is composed of at least 22 genes that were classified into four subfamilies (SFs). In this study, the relative expression of three SFs was analysed, (1) in various plant organs at different developmental stages of one hexaploid line and two tetraploid lines—a wild and a cultivated one—and (2) in fully expanded flag leaves of 16 diploid, 54 tetraploid, and 40 hexaploid lines. The expression of Rbc S genes was found to be tissue-specific (highest expression in leaves and stems) and negatively correlated with organs’ age. In all cases—at all ploidy levels, in all plant organs and developmental stages, and in most light treatments—SF-1 genes accounted for at least 80% of the Rbc S mRNA. Similar to other plant species, wheat Rbc S genes were found to be light regulated, with the expression of SF-1 genes being less affected by light than the other two SFs. A negative correlation between the expression level and RFLP was found. Accordingly, the SF with the highest level of expression (SF-1) was the most conserved, suggesting a higher selection pressure for the more active Rbc S genes.

The analysis of the pattern of isochromosome pairing allows one to distinguish factors affecting presynaptic alignment of homologous chromosomes from those affecting synapsis and crossing-over. Because the two homologous arms in an isochromosome are invariably associated by a common centromere, the suppression of pairing between these arms (intrachromosome pairing) would indicate that synaptic or postsynaptic events were impaired. In contrast, the suppression of pairing between an isochromosome and its homologous chromosome (interchromosome pairing), without affecting intrachromosome pairing, would suggest that homologous presynaptic alignment was impaired. We used such an isochromosome system to determine which of the processes associated with chromosome pairing was affected by the Ph1 gene of common wheat—the main gene that restricts pairing to homologues. Ph1 reduced the frequency of interchromosome pairing without affecting intrachromosome pairing. In contrast, intrachromosome pairing was strongly reduced in the absence of the synaptic gene Syn-B1. Premeiotic colchicine treatment, which drastically decreased pairing of conventional chromosomes, reduced interchromosome but not intrachromosome pairing. The results support the hypothesis that premeiotic alignment is a necessary stage for the regularity of meiotic pairing and that Ph1 relaxes this alignment. We suggest that Ph1 acts on premeiotic alignment of homologues and homeologues as a means of ensuring diploid-like meiotic behavior in polyploid wheat.

The cytologically diploid-like meiotic behavior of hexaploid wheat (i.e., exclusive bivalent pairing of homologues) is largely controlled by the pairing homoeologous gene Ph1. This gene suppresses pairing between homoeologous (partially homologous) chromosomes of the three closely related genomes that compose the hexaploid wheat complement. It has been previously proposed that Ph1 regulates meiotic pairing by determining the pattern of premeiotic arrangement of homologous and homoeologous chromosomes. We therefore assume that Ph1 action may be targeted at the interaction of centromeres with spindle microtubules—an interaction that is critical for movement of chromosomes to their specific interphase positions. Using monosomic lines of common wheat, we studied the effect of this gene on types and rates of centromere division of univalents at meiosis. In the presence of the normal two doses of Ph1, the frequency of transverse breakage (misdivision) of the centromere of univalent chromosomes was high in both first and second meiotic divisions; whereas with zero dose of the gene, this frequency was drastically reduced. The results suggest that Ph1 is a trans-acting gene affecting centromere-microtubules interaction. The findings are discussed in the context of the effect of Ph1 on interphase chromosome arrangement.

Recently, we reported on the characterization of the calmodulin (CaM) gene family in wheat [44]. We classified wheat CaM genes into four subfamilies (SFs) designated SF-1 to SF-4, each representing a series of homoeoallelic loci on the homoeologous chromosomes of the three genomes of common wheat. Here we studied the expression of these wheat CaM genes in the course of wheat development. Northern blot analysis using SF-specific probes revealed differences in SF expression levels in different organs and stages of development. Subsequently, cell-specific expression of CaM SFs was investigated by in situ RNA hybridization. In developing seeds, all CaM SFs showed highest expression in the embryo and less in the aleurone and in the starchy endosperm. In primary roots, all four CaM SFs were expressed in the root cap, meristematic regions and in differentiating cells. During development of the roots, expression gradually decreased. The wheat glutenin gene, which was used as a control throughout our experiments, was found to be expressed in the starchy endosperm but not in the aleurone, embryos or vegetative tissues. In stems, at advanced stages of growth, differences in cell-specific expression of CaM SFs were found. For example, SF-2 was highly expressed in differentiating phloem fibers. Thus, CaM genes in common wheat exhibit a developmentally regulated organ-, tissue-, cell- and SF-specific expression patterns.

International audience; Wild emmer wheat, Triticum turgidum subsp. dicoccoides, (2n = 49x = 28; genome BBAA), the progenitor of domesticated wheat, is genetically closely related to durum and common wheat. This wild taxon has characteristics that would be valuable if transferred to domesticated wheat. A series of chromosome-arm substitution lines (CASLs) of wild emmer wheat were produced in the background of an Israeli common wheat cultivar. These CASLs were evaluated in a pot experiment and in field trials in Israel and California for their grain yield (GY) and its components and for grain protein percentage. In addition, the extent of genetic interactions (epistatic effects) between "wild" and "domesticated" alleles, within and between homoeologous groups 1 and 7 as expressed in grain and protein yields and other quantitative traits, were determined. The research has shown that wild emmer wheat harbors genetic variability for quantitative traits and that the "wild" genes interact among themselves in a non-additive way in the common genetic background. Several chromosome arms improve GY and protein percentage in common wheat, but their effects will be presumably enhanced when combination of genes from several "wild" arms are integrated into a single "domesticated" genotype. Hence, the interaction between these genes and those in the recipient common wheat must be accounted for when higher yield or protein content is desired. The results of this study indicate on the potential of this material for breeding and genetic analysis, and support the idea of pyramiding genes from a wild species.

Summary Isolation, physical mapping and polymorphism of chromosome-specific DNA sequences in wheat are reported. Following the microdissection of the long arm of chromosome 5B (5BL) of common wheat, its DNA was amplified by degenerate oligonucleotide-primed PCR and directly cloned into plasmid vectors. Characterization of the chromosome arm library showed that ~55% of the inserts are of low-copy nature. Southern analysis using aneuploid lines of common wheat revealed that five of 11 low-copy inserts analyzed map to chromosome arm 5BL; four of these are 5BL-specific. By deletion mapping, the 5BLspecific sequences were located to sub- chromosome arm regions. Based on the hybridization patterns of three 5BLspecific sequences to DNA from a diverse collection of goat-grass (Aegilops) and wheat (Triticum) species, it was concluded that these sequences emerged at different times in the course of evolution of this group of plant species.

The wheat group has evolved through allopolyploidization, namely, through hybridization among species from the plant genera Aegilops and Triticum followed by genome doubling. This speciation process has been associated with ecogeographical expansion and with domestication. In the past few decades, we have searched for explanations for this impressive success. Our studies attempted to probe the bases for the wide genetic variation characterizing these species, which accounts for their great adaptability and colonizing ability. Central to our work was the investigation of how allopolyploidization alters genome structure and expression. We found in wheat that allopolyploidy accelerated genome evolution in two ways: (1) it triggered rapid genome alterations through the instantaneous generation of a variety of cardinal genetic and epigenetic changes (which we termed “revolutionary” changes), and (2) it facilitated sporadic genomic changes throughout the species’ evolution (i.e., evolutionary changes), which are not attainable at the diploid level. Our major findings in natural and synthetic allopolyploid wheat indicate that these alterations have led to the cytological and genetic diploidization of the allopolyploids. These genetic and epigenetic changes reflect the dynamic structural and functional plasticity of the allopolyploid wheat genome. The significance of this plasticity for the successful establishment of wheat allopolyploids, in nature and under domestication, is discussed.

The importance of hybridization and polyploidization in wheat speciation has been recognized for close to a century (Sakamura 1918; Kihara 1919, 1924, 1954; Percival 1921; Sax 1927). Following these pioneering works, it quickly became apparent that polyploid wheats are not the sum of their constituent genomes. This is not unexpected because the nascent hybrids/polyploids are equipped with a complex set of regulatory elements and of copy number variation that originate from two or more divergent genomes and that generate novel types of interactions and dosage effects. Moreover, they have to adjust at the cytological level, at the level of gene expression, and at the protein level. They also have to maintain genome stability through the regulation of meiotic pairing and recombination, the orchestration of cell division, and the silencing of transposons. The recent studies described here provide an impressive account with regard to the extent and the rapid time course at which a new genetic variant was established upon hybridization and polyploidization. We describe here the current knowledge on the changes that occurred in the wheat genome upon allopolyploidization, starting from the early evolutionary and cytological studies to the recent genomic analyses.

We report here on the successful painting of a specific plant chromosome within its own genome. Isochromosomes for the long arm of chromosome 5 of the wheat B genome (5BL) were microdissected from first meiotic metaphase spreads of a monoisosomic 5BL line of the common wheat Triticum aestivum cv. Chinese Spring. The dissected isochromosomes were amplified by degenerate oligonucleotide-primed PCR in a single tube reaction. The amplified DNA was used as a complex probe mixture for fluorescent in situ hybridization on first meiotic metaphase spreads of lines carrying 5BL as a distinctive marker. Hybridization signals were observed, specifically, along the entire 5BL. In some of the cells, labeling was also detected in two bivalents, presumably those of the 5B "homoeologues" (partial homologues) found in common wheat (5A and 5D). The probe also revealed discrete domains in tapetal nuclei at interphase, further supporting the probe's high specificity. These data suggest that chromosome and homoeologous group-specific sequences are more abundant in 5BL than genome-specific sequences. Chromosome-painting probes, such as the one described here for 5BL, can facilitate the study of chromosome evolution in polyploid wheat.

The Ph1 (pairing homoeologous) gene of common wheat, the main factor responsible for the cytological diploid-like behavior of this polyploid species, has been a subject of intensive cytogenetic studies in the last three decades. This paper reviews the effects of Ph1 on chromosomal pairing and spindle microtubule dynamics. It presents data supporting the hypothesis that this gene exerts its effect at presynaptic stages, affecting the premeiotic alignment of homologous and homoeologous chromosomes, and thereby controls the regularity and pattern of pairing at meiosis

The inheritance of grain protein percentage and of grain weight were studied by crossing common and durum wheat cultivars with hexaploid and tetraploid breeding lines that excel in grain protein percentage. All high protein lines were descendants of the tetraploid wild emmer Triticum turgidum var. dicoccoides. One hexaploid cultivar was also crossed with a high-protein var. dicoccoides genotype. All crosses were made between low- and high-protein genotypes and were carried out reciprocally for any combination of genotypes; some of them between genotypes of the same ploidy level and some between hexaploid and tetraploid lines. Weight and protein percentage were determined in selfed and crossed grains that developed on the same spike. Mean weight and protein percentage were also determined in F2 grains of all crosses of the same ploidy level, either tetraploid or hexaploid. At any ploidy level, F1 grains resembled the selfed grains of the mother plant both in grain weight and in grain protein percentage, indicating a major maternal effect on both traits. F2 grains had similar grain weight to the heavy-grained parent, and their protein percentage was close to the midparents value. However, a slight indication of cytoplasmic inheritance of grain protein percentage was found in the comparison between most pairs of F2 reciprocals. The interspecific crosses (hexaploid with tetraploid combinations) yielded shrivelled seeds with highly reduced weight but relatively unchanged protein percentage. Weight reduction in the shrivelled hybrid grains (compared with the selfed ones) was more severe when the mother plant was hexaploid rather than tetraploid. The significance of the different tissues in determining grain weight and protein percentage is discussed.Key words: grain weight, grain protein percentage, maternal effect, paternal effect, reciprocal crosses, wheat, Triticum aestivum, Triticum turgidum var. dicoccoides.

Three different 3' noncoding sequences of wheat rubisco small subunit (SSU) genes (RbcS) were used as probes to identify the gene members of different RbcS subfamilies in the common wheat cultivar Chinese Spring (CS). All genes of the wheat RbcS multigene family were previously assigned to the long arm of homoeologous group 5 and to the short arm of homoeologous group 2 chromosomes of cv CS. Extracted DNA from various aneuploids of these homoeologous groups was digested with four restriction enzymes and hybridized with three different 3' noncoding sequences of wheat SSU clones. All RbcS genes located on the long arm of homoeologous group 5 chromosomes were found to comprise a single subfamily, while those located on the short arm of group 2 comprised three subfamilies. Each of the ancestral diploid genomes A, B, and D has at least one representative gene in each subfamily, suggesting that the divergence into subfamilies preceded the differentiation into species. This divergence of the RbcS genes, which is presumably accompanied by a similar divergence in the 5' region, may lead to differential expression of various subfamilies in different tissues and in different developmental stages, in response to different environmental conditions. Moreover, members of one subfamily that belong to different genomes may have diverged also in the coding sequence and, consequently, code for distinguishable SSU. It is assumed that such utilization of the RbcS multigene family increases the adaptability and phenotypic plasticity of common wheat over its diploid progenitors.

Recent studies in the genera Aegilops and Triticum showed that allopolyploid formation triggers rapid genetic and epigenetic changes that lead to cytological and genetic diploidization. To better understand the consequences of cytological diploidization, chromosome pairing and seed fertility were studied in S1, S2, and S3generations of 18 newly formed allopolyploids at different ploidy levels. Results showed that bivalent pairing at first meiotic metaphase was enhanced and seed fertility was improved during each successive generation. A positive linear relationship was found between increased bivalent pairing, improved fertility, and elimination of low-copy noncoding DNA sequences. These findings support the conclusion that rapid elimination of low-copy noncoding DNA sequences from one genome of a newly formed allopolyploid, different sequences from different genomes, is an efficient way to quickly augment the divergence between homoeologous chromosomes and thus bring about cytological diploidization. This facilitates the rapid establishment of the raw allopolyploids as successful, competitive species in nature.

The genes coding for the Rubisco small subunit (SSU) and for the α-subunit of the Rubisco-binding protein were located to chromosome arms of common wheat. HindIII-digested total DNA from the hexaploid cultivar Chinese Spring and from ditelosomic and nullisomic-tetrasomic lines was probed with these two genes, whose chromosomal location was deduced from the disappearance of or from changes in the relative intensity of the relevant band(s). The Rubisco SSU pattern consisted of 14 bands, containing at least 21 different types of DNA fragments, which were allocated to two homoeologous groups: 15 to the short arm of group 2 chromosomes (4 to 2AS, 7 to 2BS, and 4 to 2DS) and 6 to the long arm of group 5 chromosomes (2 on each of arms 5AL, 5BL, and 5DL). The pattern of the Rubisco-binding protein consisted of three bands, each containing one type of fragment. These fragments were located to be on the short arm of group 2 chromosomes. The restriction fragment length polymorphism (RFLP) patterns of several hexaploid and tetraploid lines were highly conserved, whereas the patterns of several of their diploid progenitors were more variable. The variations found in the polyploid species were mainly confined to the B genome. The patterns of the diploids T. monococcum var. urartu and Ae. squarrosa were similar to those of the A and D genome, respectively, in polyploid wheats. The pattern of T. monococcum var. boeoticum was different from the patterns of the A genome, and the patterns of the diploids Ae. speltoides, Ae. longissima, and Ae. Searsii differed from that of the B genome.

One of the intriguing issues concerning the dynamics of plant genomes is the occurrence of intraspecific variation in nuclear DNA amount. The aim of this work was to assess the ranges of intraspecific, interspecific, and intergeneric variation in nuclear DNA content of diploid species of the tribe Triticeae (Poaceae) and to examine the relation between life form or habitat and genome size. Altogether, 438 plants representing 272 lines that belong to 22 species were analyzed. Nuclear DNA content was estimated by flow cytometry. Very small intraspecific variation in DNA amount was found between lines of Triticeae diploid species collected from different habitats or between different morphs. In contrast to the constancy in nuclear DNA amount at the intraspecific level, there are significant differences in genome size between the various diploid species. Within the genus Aegilops , the 1C DNA amount ranged from 4.84 pg in A. caudata to 7.52 pg in A. sharonensis; among genera, the 1C DNA amount ranged from 4.18 pg in Heteranthelium piliferum to 9.45 pg in Secale montanum . No evidence was found for a smaller genome size in annual, self-pollinating species relative to perennial, cross-pollinating ones. Diploids that grow in the southern part of the group’s distribution have larger genomes than those growing in other parts of the distribution. The contrast between the low variation at the intraspecific level and the high variation at the interspecific one suggests that changes in genome size originated in close temporal proximity to the speciation event, i.e., before, during, or immediately after it. The possible effects of sudden changes in genome size on speciation processes are discussed.

The aim of the present study was to assess the genetic variation in several Israeli and Turkish populations of wild emmer wheat, Triticum turgidum ssp. dicoccoides, the progenitor of most domesticated wheat. Single spikes were collected in 2002 from 60 plants that grew in six different habitats in Ammiad, northeastern Israel (8-12 plants from each habitat), and in 1998 from 56 plants that grew in seven different habitats in Diyarbakir, southeastern Turkey (8 plants from each habitat). Seeds were planted in a nursery and DNA was extracted from every plant and analyzed by the fluorescent-based amplified fragment length polymorphism (AFLP) method. Seven primer combinations produced 788 discernible loci of which 48.6% were polymorphic in Israel and 40.5% in Turkey. The genetic diversity estimates P (frequency of polymorphic loci) and He (gene diversity) were higher in Ammiad than in Diyarbakir (means of P = 0.34 and He = 0.13 in Ammiad vs. P = 0.20 and He = 0.08 in Diyarbakir). Ammiad populations contained more unique alleles than Diyarbakir populations. The relative genetic diversity estimates (theta) values were 0.188 in Ammiad and 0.407 in Diyarbakir, suggesting better differentiation of the populations in Turkey. Genetic distance was larger between Israeli and Turkish populations than between populations of each country. The data indicate that the Israeli and Turkish populations are considerably diverged and that the Israeli populations are more polymorphic than the Turkish ones, having a larger within-populations genetic variation than among-populations one. The significance of the results in relation to the differentiation pattern of wild emmer in the Near East is discussed.

Corrigendum: Transcriptional activation of retrotransposons alters the expression of adjacent genes in wheat

Genetic structure of natural populations of wild crop relatives has been the subject of many studies. Yet, most of them focused on the assessment of spatial genetic diversity, while information on long-term variation, affected by yearly changes, has been considered only in few cases. The present study aimed therefore, to estimate the spatio-temporal genetic variation in populations of wild emmer wheat, the progenitor of domesticated wheat, and to assess the contribution of spatial versus temporal factors to the maintenance of genetic variation in a population. Single spikes were collected in the years 1988 and 2002 from plants that grew in the same sampling points, from six different habitats in the Ammiad conservation site, Eastern Galilee, Israel. Seeds were planted in a nursery and DNA was extracted from each plant and analyzed by the AFLP method. Fourteen primer combinations yielded 1,545 bands of which 50.0 and 48.8% were polymorphic in the years 1988 and 2002, respectively. Genetic diversity was much larger within populations than between populations and the temporal genetic diversity was considerably smaller than the spatial one. Nevertheless, population genetic structure may vary to some degree in different years, mainly due to fluctuations in population size because of yearly rainfall variations. This may lead to predominance of different genotypes in different years. Clustering the plants by their genetic distances grouped them according to their habitats, indicating the existence of genotype-environment affinities. The significance of the results in relation to factors affecting the maintenance of polymorphism in natural populations is discussed.

One major strategy to increase the level of zinc (Zn) and iron (Fe) in cereal crops, is to exploit the natural genetic variation in seed concentration of these micronutrients. Genotypic variation for Zn and Fe concentration in seeds among cultivated wheat cultivars is relatively narrow and limits the options to breed wheat genotypes with high concentration and bioavailability of Zn and Fe in seed. Alternatively, wild wheat might be an important genetic resource for enhancing micronutrient concentrations in seeds of cultivated wheat. Wild wheat is widespread in diverse environments in Tarkey and other parts of the Fertile Crescent (e.g., Iran, Iraq, Lebanon, Syria, Israel, and Jordan). A large number of accessions of wild wheat and of its wild relatives were collected from the Fertile Crescent and screened for Fe and Zn concentrations as well as other mineral nutrients. Among wild wheat, the collections of wild emmer wheat, Triticum turgidum ssp. dicoccoides (825 accessions) showed impressive variation and the highest concentrations of micronutrients, significantly exceeding those of cultivated wheat. The concentrations of Zn and Fe among the dicoccoides accessions varied from 14 to 190 mg kg -1 DW for Zn and from 15 to 109 mg kg -1 DW for Fe. Also for total amount of Zn and Fe per seed, dicoccoides accessions contained very high amount of Zn (up to 7 µg per seed) and Fe (up to 3.7 µg per seed). Such high genotypic variation could not be found for phosphorus, magnesium, and sulfur. In the case of modern cultivated wheat, seed concentrations of Zn and Fe were lower and less variable when compared to wild wheat accessions. There was a highly significant positive correlation between seed concentrations of Fe and Zn. Screening different series of dicoccoides substitution lines revealed that the chromosome 6A, 611, and 5B of dicoccoides resulted in greater increase in Zn and Fe concentration when compared to their recipient parent and to other chromosome substitution lines. The results indicate that Triticum turgidum L. var. dicoccoides (wild emmer) is an important genetic resource for increasing concentration and content of Zn and Fe in modern cultivated wheat. © 2004 Taylor & Francis Group, LLC.

To better understand genetic events that accompany allopolyploid formation, we studied the rate and time of elimination of eight DNA sequences in F1 hybrids and newly formed allopolyploids of Aegilops and Triticum. In total, 35 interspecific and intergeneric F1 hybrids and 22 derived allopolyploids were analyzed and compared with their direct parental plants. The studied sequences exist in all the diploid species of the Triticeae but occur in only one genome, either in one homologous pair (chromosome-specific sequences [CSSs]) or in several pairs of the same genome (genome-specific sequences [GSSs]), in the polyploid wheats. It was found that rapid elimination of CSSs and GSSs is a general phenomenon in newly synthesized allopolyploids. Elimination of GSSs was already initiated in F1 plants and was completed in the second or third allopolyploid generation, whereas elimination of CSSs started in the first allopolyploid generation and was completed in the second or third generation. Sequence elimination started earlier in allopolyploids whose genome constitution was analogous to natural polyploids compared with allopolyploids that do not occur in nature. Elimination is a nonrandom and reproducible event whose direction was determined by the genomic combination of the hybrid or the allopolyploid. It was not affected by the genotype of the parental plants, by their cytoplasm, or by the ploidy level, and it did not result from intergenomic recombination. Allopolyploidy-induced sequence elimination occurred in a sizable fraction of the genome and in sequences that were apparently noncoding. This finding suggests a role in augmenting the differentiation of homoeologous chromosomes at the polyploid level, thereby providing the physical basis for the diploid-like meiotic behavior of newly formed allopolyploids. In our view, this rapid genome adjustment may have contributed to the successful establishment of newly formed allopolyploids as new species.

The general goal was to identify, map, and tag, with DNA markers, segments of chromosomes of a wild species (wild emmer wheat, the progenitor of cultivated wheat) determining the number, chromosomal locations, interactions, and effects of genes that control quantitative traits when transferred to a cultivated plant (bread wheat). Slight modifications were introduced and not all objectives could be completed within the human and financial resources available, as noted with the specific objectives listed below: 1. To identify the genetic contribution of each of the available wild emmer chromosome-arm substitution lines (CASLs) in the bread wheat cultivar Bethlehem for quantitative traits, including grain yield and its components and grain protein concentration and yield, and the effect of major loci affecting the quality of end-use products. [The quality of end-use products was not analyzed.] 2. To determine the extent and nature of genetic interactions (epistatic effects) between and within homoeologous groups 1 and 7 for the chromosome arms carrying "wild" and "cultivated" alleles as expressed in grain and protein yields and other quantitative traits. [Two experiments were successful, grain protein concentration could not be measured; data are partially analyzed.] 3. To derive recombinant substitution lines (RSLs) for the chromosome arms of homoeologous groups 1 and 7 that were found previously to promote grain and protein yields of cultivated wheat. [The selection of groups 1 and 7 tons based on grain yield in pot experiments. After project began, it was decided also to derive RSLs for the available arms of homoeologous group 4 (4AS and 4BL), based on the apparent importance of chromosome group 4, based on early field trials of the CASLs.] 4. To characterize the RSLs for quantitative traits as in objective 1 and map and tag chromosome segments producing significant effects (quantitative trait loci, QTLs by RFLP markers. [Producing a large population of RSLs for each chromosome arm and mapping them proved more difficult than anticipated, low numbers of RSLs were obtained for two of the chromosome arms.] 5. To construct recombination genetic maps of chromosomes of homoeologous groups 1 and 7 and to compare them to existing maps of wheat and other cereals [Genetic maps are not complete for homoeologous groups 4 and 7.] The rationale for this project is that wild species have characteristics that would be valuable if transferred to a crop plant. We demonstrated the sequence of chromosome manipulations and genetic tests needed to confirm this potential value and enhance transfer. This research has shown that a wild tetraploid species harbors genetic variability for quantitative traits that is interactive and not simply additive when introduced into a common genetic background. Chromosomal segments from several chromosome arms improve yield and protein in wheat but their effect is presumably enhanced when combination of genes from several segments are integrated into a single genotype in order to achieve the benefits of genes from the wild species. The interaction between these genes and those in the recipient species must be accounted for. The results of this study provide a scientific basis for some of the disappointing results that have historically obtained when using wild species as donors for crop improvement and provide a strategy for further successes.

The RbcS multigene family of hexaploid (bread) wheat, Triticum aestivum (genome BBAADD), which encodes the small subunit of Rubisco, comprises at least 22 genes. Based on their 3′ non-coding sequences, these genes have been classified into four subfamilies (SFs), of which three (SF-2, SF-3 and SF-4) are located on chromosomes of homoeologous group 2 and one (SF-1) on homoeologous group 5. In the present study we hybridized three RbcS subfamily-specific probes (for SF-1, SF-2 and SF-3) to total DNA digested with four restriction enzymes and analyzed the RFLP patterns of these subfamilies in eight diploid species of Aegilops and Triticum, and in two tetraploid and one hexaploid species of wheat (the diploid species are the putative progenitors of the polyploid wheats). The three subfamilies varied in their level of polymorphism, with SF-2 being the most polymorphic in all species. In the diploids, the order of polymorphism was SF-2 > SF-3 > SF-1, and in the polyploids SF-2 > SF-1 > SF-3. The RbcS genes of the conserved SF-1 were previously reported to have the highest expression levels in all the wheat tissues studied, indicating a negative correlation between polymorphism and gene expression. Among the diploids, the species with the D and the S genomes were the most polymorphic and the A-genome species were the least polymorphic. The polyploids were less polymorphic than the diploids. Within the polyploids, the A genome was somewhat more polymorphic than the B genome, while the D genome was the most conserved. Among the diploid species with the A genome, the RFLP pattern of T. urartu was closer to that of the A genome of the common wheat cultivar Chinese Spring (CS) than to that of T. monococcum. The pattern in Ae. tauschii was similar to that of the D genome of CS. Only partial resemblance was found between the RFLP patterns of the species with the S genome and the B genome of CS.

Chromosome painting is a technique that allows the specific visualization of an entire chromosome in metaphase spreads and in interphase nuclei by in situ hybridization with a mixture of sequences generated from that particular chromosome (Pinkel et al., 1988; Carter, 1994). In the first chromosome painting experiments the source of composite probes were recombinant libraries from specific human chromosomes isolated by flow cytometry (Pinkel et al., 1988; Lichter et al., 1988). Painting probes are now generated more efficiently from microdissected chromosomes by a general PCR amplification of the chromosomal DNA with a degenerated primer (Telenius et al., 1992a, b; Guan et al., 1994).

Among the major crops, cereals constitute about 50% of the annual world protein production used for animal and human consumption (Hanson et al. 1982). Among cereals, wheat production exceeds all other crops, accounting for about 40% of the total protein production. Wheat, therefore, has the major contribution of any single crop to the world protein production (Harlan and Starks 1980). Naturally, the least expensive and most effective means of increasing protein production and upgrading its quality is through the improvement of wheat protein. This chapter deals primarily with the quantitative aspects of high protein wheat.

The mode of inheritance, linkage groups, and chromosomal location of 23 morphological and 4 biochemical traits were characterized in the wild tetraploid emmer wheat,Triticum turgidum var.dicoccoides. These traits were described and their mode of inheritance was determined by their segregation in four F2 populations derived from crosses between four var.dicoccoides accessions and a tetraploiddurum cultivar. Linkage groups among the genes encoding for these traits were determined or postulated, and their chromosomal location was deduced by linkage to previously located genes. The genetic control of the following traits was characterized and is first reported here: black keel; hairy leaf sheath; hairy auricles; hairy rachilla; hairy kernel brush; obtuse flag leaf; and curved neck/peduncle. The linkage data indicated that developmentally-related genes tended to occur in clusters.

The recently described Aegilops searsii Feld. &Kis. is morphologically closely related to Ae. longissima Schweinf. et Muschl. The cytogenetic affinity of these two species was estimated by studying karyotypes, chromosomal pairing, pollen fertility and seed set in parental lines and in intra- and inter-specific F1 hybrids. Pairing in Ae. longissima × Ae. searsii hybrids was incomplete and indicative of several structural rearrangements of parental chromosomes. The hybrids were highly sterile, indicating that the two species are well isolated from one another by internal chromosomal and genetic barriers. These isolating mechanisms are superimposed upon strong ecological specialization of each species. The data further support the validity of the specific rank of Ae. searsii and indicate that it contains a distinct genome which was designated Ss.

Polymorphism of the high molecular weight glutenin subunits was studied in 456 accessions of the wild wheat Triticum turgidum var. dicoccoides (2n=4x=28; genomes AABB), originating from 21 populations in Israel. A total of 50 different SDS PAGE migration patterns were observed, resulting from the combinations of 15 subunit patterns of the A genome and 24 subunit patterns of the B genome. Most migration patterns consisted of five subunits, varying between three and six. The migration patterns of the A genome had zero to three subunits—two being most common. The apparent molecular weights (MWs) of the slowest migrating subunit (114,000 to 103,500) and of the next in rate of migration (106,000 to 96,000) were highly correlated (r = 0·97). Also, both subunits were either present (in most accessions) or absent. In 82.3 per cent of the accessions, the third subunit (MW 76,000 to 71,500) was absent, while in 16.9 per cent of the accessions all three subunits of the A genome were absent. The migration patterns of the B genome had one to three subunits—three being most common. The slowest migrating subunit (99,500 to 93,000) was present in almost all cases (99·3 per cent). The MWs of the next two subunits (90,500 to 82,000 and 86,000 to 78,000, respectively) were highly correlated (r = 0·95). Also, either both subunits were present, as in most cases (94·4 per cent), or absent (5·6 per cent). A nomenclature for the genes encoding for the HMW glutenins is proposed based on the following model: The three subunit groups controlled by each genome are encoded by two genes. In genome A, one gene (Glu-A1–1), with 12 alleles, encodes for the two correlated subunit groups 1Ax and 1Ax'; the other gene (Glu-A1–2), with 3 alleles, encodes for the fast-migrating subunit group (1Ay). In genome B, one gene (Glu-B1-1), with 8 alleles, encodes for the slow-migrating subunit group (1Bx), and the other gene (Glu-B1–2), with 10 alleles, encodes for the two correlated subunit groups, 1By and 1By'. The polymorphism of the HMW glutenin genes found in var. dicoccoides is much higher than that of cultivated wheats as well as of genes coding for enzymes in var. dicoccoides.

Several aneuploid lines and one intervarietal substitution line of the hexaploid wheat Triticum aestivum (2 n = 6 x = 42; genomes AABBDD) cv. Chinese Spring were used to study the effects of different doses of chromosomes 1B, 1D, or 1A on the amount of the high molecular weight (“HMW”) glutenins and gliadins of endosperm. These homeologous chromosomes carry HMW glutenin and gliadin gene clusters on their long and short arms, respectively. Increasing the dosage of chromosome 1B of Chinese Spring in plants having in their 3 n endosperm zero or the normal three doses of the homeologue 1D, as well as in plants carrying in their endosperm one dose of 1B of the cultivar Timstein, had a dual effect: on one hand, a nonlinear increase in the amount of each subunit encoded by the chromosome whose dosage was elevated and, on the other hand, a compensating nonspecific decrease in the amount of other HMW glutenin and gliadin subunits encoded either by the homeoalleles on 1D or by the homoalleles on 1B of Timstein, respectively. Deletion of chromosome arm 1BL, which carries only a few HMW glutenin genes, had no significant effect on the amount of HMW glutenins encoded by 1DL and HMW gliadins encoded by 1DS and 1BS. However, deletion of 1BS or 1DS, each carrying many gliadin genes, caused a significant but nonspecific increase in the HMW glutenins and gliadins encoded by the remaining arms of 1B and 1D. The possible mechanism and evolutionary implications of gene-dosage compensation in polyploid wheat are discussed.

Distances between marked pairs of bivalents were determined at first meiotic metaphase of common wheat (2n=6x=42; genome AABBDD) by tallying the number of bivalents intervening between the marked bivalents. Bivalents of the same genome tended to lie relatively close to one another, whereas those of different genomes were further apart on the metaphase plate. Within each genome the association of telocentric bivalents representing the two different arms of one chromosome was much more intimate than that of genetically nonrelated bivalents. Among chromosomes of different genomes, homoeologous bivalents were relatively closer to one another than nonrelated ones. These results were more pronounced in cells with circularly arranged bivalents on the metaphase plate than in those with a linear arrangement. The data, consistent with previous findings in cold-treated somatic cells, indicate that a similar pattern of chromosomal arrangement exists in somatic and meiotic cells and that chromosomal arrangement observed in arrested metaphases is also maintained on the untreated, functional spindle system.

The frequency of interlocking bivalents at first meiotic metaphase of common wheat Triticum aestivum L., which is normally very low, is significantly increased by raising the dosage (from two to three, four and six) of the Ph1 gene, located on the long arm of chromosome 5B (5BL). In several cells more than three bivalents were interlocked in one chain configuration indicating involvement of non-homoeologous bivalents. Plants with reduced dose (one or zero) of Ph1 also exhibited an increased frequency of interlocking but to a lesser extent than those with high gene dosage. However, chains of more than three interlocked bivalents were never observed in these plants, suggesting that with one or zero doses of Ph1 interlocking is restricted to homoeologous bivalents only. Chromosomal arm 5BS affected interlocking in an opposite manner to 5BL; namely, two and four doses of 5BS markedly reduced interlocking frequency. The modification in the frequency of interlocking bivalents by these genetic manipulations represents the first successful attempt to affect interlocking by genetic means. The results are explained on the basis of the hypothesis that this gene system controls somatic and premeiotic association of both homologous and homoeologous chromosomes.

The inheritance of the high molecular weight (HMW) glutenins and of several gliadins controlled, respectively, by the long and short arms of chromosome 1B of common wheat was studied. Analysis was carried out on the progeny of two inter-varietal crosses in which the parental lines possessed differentially migrating subunits as revealed by sodium dodecyl sulphate polyacrylamide gel electrophoresis. No recombination event was detected either within the fraction of the HMW glutenins or among most of the gliadin subunits studied indicating that they are controlled by tightly linked gene clusters. One gliadin subunit (B30) showed 25.5% recombination frequency with the rest of the gliadin subunits and 23.5% recombination frequency with the fraction of the HMW glutenin subunits. It has been concluded that this subunit is controlled by a separate locus (Gld-B6), proximal to the major gliadin gene cluster on the short arm of chromosome 1B. Consequently, the recombination percentage between the glutenin loci and most of the gliadin loci was calculated as 49.0 and the distance in centi-Morgans (cM) as 53.6. The estimated distance in cM is very close to the observed recombination percentage. A genetic map of these storage protein genes is presented.

Grain protein percentage (GPP) was studied in 910 accessions of the wild tetraploid wheat, Triticum turgidum var. dicoccoides, collected from 22 populations representing different ecogeographical conditions in Israel. High values of GPP were found, ranging from 19.7% to 28.0% for population means, and from 14.1% to 35.1% for single accessions. Marginal populations had usually lower GPP and smaller intra-population variation than central ones. Repeated sampling of some central populations for four consecutive years revealed relatively large intra-population fluctuations. A high and significant genetic component of variation was found within and between populations by a nested analysis of variance in two nurseries. However, the regression coefficients of parents vs. offsprings were relatively low, indicating a smaller genetic component of variation which may be accounted for by a significant genotype × environment interaction. No correlation was found between GPP and ecological factors, except for soil type: accessions growing on terra-rossa had higher GPP than those growing on basaltic soil. Accessions with black glumes, or with glabrous auricles, or with large grains exhibited high GPP values. A strategy for collecting accessions with high GPP is presented, and the potential use of high GPP genotypes in breeding programs is discussed.

Total endosperm protein subunits, extracted from the common wheat cultivar Chinese Spring and from some of its aneuploid lines, were fractionated according to their molecular weight (MW) in an improved high resolution one-dimensional sodium dodecyl sulphate (SDS) polyacrylamide gel electrophoresis (PAGE). The resolution obtained by this method and, in particular, that of the high molecular weight (HMW) glutenin and gliadin subunits approached that of a previous report in which two-dimensional fractionation system based on charge and MW was used. In the cultivar Chinese Spring, 21 discrete protein bands were resolved and the chromosomes controlling many of them were either reconfirmed, or, in some cases, established. The advantages of this high resolution SDS PAGE technique are discussed.

Acetylene reduction activity (ARA) was measured in cores containing roots of various Israeli wild and cultivated wheat lines colonized by Azospirillum. The inoculated plants were grown under greenhouse or field conditions. Although, no measurable ARA was detected during earlier stages of wheat development, 50–600 nmol C2H4 g−1 dry root h−1 was measured during heading and flowering stages. By using N yield balance and 15N dilution techniques, it was found that Triticum aestivum cv. Miriam inoculated with Azospirillum accumulated 20% more N (14N and 15N) at the booting stage than did the uninoculated control. This difference in N content became less apparent in grains. No significant 15N dilution could be found and the contribution of atmospheric N2 to the N content of grains of inoculated plants was negligible. It was concluded that the potential contribution of biological N2 fixation to spring wheat cultivation in Israel is very low.

Two lines of common wheat cv. Chinese Spring carrying mutant alleles for the Ph2 homoeologous pairing-suppressor gene on chromosome arm 3DS promoting homoeologous pairing in wheat interspecific and intergeneric hybrids have been tested for their mitotic sensitivity to colchicine. Both the ph2a mutation, which corresponds to a fairly long deletion of 3DS, and ph2b, which is either an intragenic change or a very small deletion, as well as the ph2a/ph2b heteroallelic combination and the deficiency for the entire 3DS arm (ditelo 3DL), conditioned a significantly lower sensitivity to colchicine than that determined by the wild-type allele Ph2 (euploid and ditelo 3DS). Observation of both metaphase and anaphse root-tip cell populations, treated with various colchicine concentrations, provided similar results. The degree of spindle disruption in 2 × 10−4 M colchicine, as measured by the percentage of fully affected metaphases (C type), was significantly reduced in ph2 and Ph2− genotypes and, consequently, a larger proportion of cells could proceed toward anaphase and also had a regular segregation pattern at this stage. The differential sensitivity of ph2 genotypes to colchicine is in the opposite direction to that previously found for ph1 genotypes, lacking the homoeologous pairing suppressor on chromosome arm 5BL. The ph2 mutation, while promoting homoeologous pairing as ph1 does, decreases spindle sensitivity to colchicine with respect to the wild-type (ph2) allele rather than increasing it, as ph1 does. The observed alteration of spindle sensitivity to colchicine that mutants for structurally unrelated but functionally related genes (Ph1 and Ph2) condition is interpreted as a highly probable coincidence in their cellular target. It is assumed therefore that these loci affect the equilibrium between tubulin and microtubules. Through this effect they presumably determine chromosome positioning in somatic and premeiotic stages, leading to different pairing patterns at meiosis. Key words: Triticum aestivum, tubulin, pairing suppressors, somatic association.

Five field experiments were conducted during 1982 and 1983 to evaluate the yield response of two lines of common wheat and one line of durum wheat to several rhizospheric bacteria. The two lines tested in the 1982 experiment (a common wheat and a durum wheat) showed a considerable yield response to incoulation withAzospirillum brasilense, with an average significant yield incresae of 23%. In the 1983 experiments, variable yield responses, some of which were significant, were obtained by inoculating three wheat lines (two common wheat and one durum wheat) with Azospirillum or with other rhizospheric bacteria, isolated from various Israeli wild relatives of wheat. A certain specificity between the tested wheat lines, the applied bacteria and the particular local conditions was evident. Inverse relationships were found between the maximum percentage yield increase due to inoculation with the most efficient bacteria and the yield level of the non-inoculated plots.

Several quantitative traits were studied in Aegilops longissima addition lines and in tetrasomic lines of the common wheat cultivar Chinese Spring. The effects of gene dosage on these traits, which were highly significant and specific for each of the seven homoeologous groups, were more numerous and caused greater differences than the alien genetic effects. It is therefore suggested that in using alien addition lines for evaluating the effect of alien chromosomes on quantitative traits, the homoeologous tetrasomic lines should be used as a control. In comparison to the suggested approach, the use of substitution lines, which are much more difficult to obtain than addition lines, proved, in some cases, less accurate because of the inability of the alien chromosome to fully compensate for the missing homoeologous chromosome.Key words: addition lines, substitution lines, tetrasomes, Triticum aestivum, Aegilops longissima.

Lines of common wheat cv. 'Chinese Spring' carrying different doses of the Ph1 gene, located on the long arm of chromosome 5B were treated with the antimitotic agents griseofulvin and isopropyl-N-phenyl-carbamate (IPC). Treatments with low griseofulvin concentrations and IPC resulted mainly in the production of a high percentage of cells exhibiting spindle disorganization at metaphase and multipolar cells at anaphase–telophase. These treatments did not differentially affect the tested genotypes. On the other hand, higher griseofulvin concentrations induced the appearance of frequent C-metaphases and C-anaphases because of complete disruption of the spindle microtubules; in such cases, more pronounced sensitivity was observed in a line lacking the Ph1 gene than in lines disomic and tetrasomic for chromosome 5B. From this evidence it can be concluded that subcellular structures regulating microtubule organization and orientation, which are apparently the target of IPC and low griseofulvin concentrations, are not related to the action of the Ph1 gene. Rather, tubulin–microtubules equilibrium, which is affected by colchicine and high griseofulvin concentrations, is influenced by the action of this gene. Thus, the Ph1 gene product may correspond to a tubulin or a microtubule-associated protein which in turn stabilizes microtubule structure. In either case, the equilibrium tubulin–microtubules would be shifted towards microtubules in plants containing two doses of Ph1 and even more in plants containing an extra dose of this gene.Key words: spindle, Triticum, microtubules, antitubulins, griseofulvin.

While an extra dose of chromosome 2A of common wheat, previously reported to carry a pairing promoter on its short arm, did not increase pairing between homoeologous chromosomes in F1 hybrids between common wheat cv. Chinese Spring (CS) and Aegilops variabilis, two doses of chromosome 2D or 2B caused a significant increase in homoeologous pairing. Evidently, chromosomes 2D and 2B carry a pairing promoter(s). Studies of F1 hybrids between aneuploids of CS, either deficient for chromosome 2D or having it in an extra dose, and Ae. variabilis, Ae. longissima, and Secale cereale supported the finding that this chromosome carries a pairing promoter. Using ditelosomic lines, the promoter was found to be located on the short arm of 2D (2DS). It was deduced that the promoter of 2B is also located on the homoeologous short arm, i.e., on 2BS. Evidence was obtained that the long arm of 2D may carry a suppressor(s) of pairing. Thus, the short arm of 2A, 2D, and 2B carries a pairing promoter(s), while the long arm of 2D and possibly of 2A and 2B carry a minor suppressor(s). The promoters are more potent than the suppressors and the overall effect of group-2 chromosomes is pairing promotion.Key words: wheat, homoeologous pairing, pairing promoter, pairing suppressor, intergeneric hybrid, meiosis.