Spontaneous haploids in durum wheat: their cytogenetic characterization

Summary Durum or macaroni wheat (Triticum turgidum L., 2n = 4x = 28; AABB genomes) arose as a natural hybrid between two wild species, Aegilops speltoides Tausch (2n = 2x = 14; BB genome) and Triticum urartu Tumanian (2n = 2x = 14; AA genome). The two progenitors hybridized in nature about half a million years ago and gave rise to tetraploid wheat, presumably in one step as a result of functioning of unreduced gametes in their hybrid BA (amphihaploid). It is easily possible to go back on the evolutionary scale and obtain durum haploids BA, and then regenerate tetraploid durum plants from them. Interestingly, such a reversion to haploidy does occur in nature as well, although at a very low frequency. This article reports on the occurrence of two spontaneous durum haploids and describes their chromosomal characteristics. The haploids (euhaploids, to be precise) had 14 somatic chromosomes, which, on fluorescent genomic in situ hybridization (fl-GISH), could be distinguished as 7 A-genome and 7 B-genome chromosomes. At meiosis, only 2.3 and 2.7% of the chromosomes paired in the two haploids, because of the presence of the homoeologous pairing-suppressor gene, Ph1. The Ph1-induced lack of pairing is a prerequisite for chromosome doubling through the formation of unreduced gametes that give rise to tetraploid durum wheats.Mention of trade names or commercial products in this publication is solely to provide specific information and does not imply recommendation or endorsement by the U.S. Department of Agriculture.     

Discovery of 2n gametes in tetraploid oat Avena vaviloviana

Summary Sexual polyploidization via the action of 2n gametes (gametes with the sporophytic chromosome number) has been identified as the most important evolutionary mode of polyploidization among plant genera. This study was conducted to determine whether 2n gametes are present in the tetraploid level of the genus Avena (2n=4×=28) Twenty tetraploid Avena lines, representing four species and one interspecific hybrid, were screened for pollen grain size in order to differentiate between n and 2n pollen. Avena vaviloviana (Malz.) Mordv. line PI 412767 was observed to contain large pollen grains at a 1.0% frequency. Cytogenetic analyses of pollen mother cells of PI 412767 revealed cells with double the normal chromosome number (i.e., 56 chromosomes at metaphase I and anaphase I). The mode of chromosome doubling was found to be failure of cell wall formation during the last mitotic division that preceded meiosis. The resulting binucleate cells underwent normal meiotic divisions and formed pollen grains with 28 chromosomes. Based on the formation and function of 2n gametes, three models involving diploid and tetraploid oat lines are proposed to describe possible evolutionary pathways for hexaploid oats. If stable synthetic hexaploid oat lines could be developed by utilizing 2n gametes from diploid and tetraploid oat species through bilateral sexual polyploidization, the resulting hexaploids could be used in breeding programs for transferring genes from diploids and tetraploids to cultivated hexaploids.     

Bilateral sexual polyploidization in potatoes

Summary Bilateral sexual tetraploidization was achieved by intercrossing diploid Phureja × haploid Tuberosum hybrids. Both parents contribute gametes with the somatic number of chromosomes (diplogynoids and diplandroids). More than 100 tetraploid progeny appeared in nine 2x–2x families. The tetraploids are generally more vigorous and significantly outyield their diploid full-sibs. The superiority of the tetraploids is interpreted on the basis of the mode of diplandroid (2n pollen) formation by first meiotic division restitution (FDR). Heterozygosis, epistasis and genetic diversity, if present in the parental diploid genotype, are largely maintained in the 2n gametes formed by FDR. and syngamy brings about more opportunities for heterotic responses in conjunction with polyploidization. Differences in tuber yield among tetraploid clones within families were found to be statistically significant (P <0.05). Thus, unlike somatic chromosome doubling, which can merely increase chromosome numbers, or somatic hybridization, which could in addition bring about heterosis, sexual polyploidization is also associated with genetic variability. The degree of inbreeding and the genotypic structure in the population of newly arisen tetraploids is discussed. The relationship among asexual reproduction, preservation of favorable nonadditive genetic effects by FDR. restoration of sexuality, and polyploidization is explored. The possible evolutionary significance of these factors is emphasized.     

Chromosome pairing in durum wheat haploids with and without <Emphasis Type="Italic">ph1b</Emphasis> of bread wheat

Durum or macaroni wheat (Triticum turgidum L., 2n = 4x = 28; AABB) is an allotetraploid with two related genomes, AA and BB, each with seven pairs of homologous chromosomes. Although the corresponding chromosomes of the two genomes are potentially capable of pairing with one another, the Ph1 (Pairing homoeologous) gene in the long arm of chromosome 5B permits pairing only between homologous partners. As a result of this Ph1-exercised disciplinary control, durum wheat and its successor, bread wheat (Triticum aestivum L., 2n = 6x = 42; AABBDD) show diploid-like chromosome pairing and hence disomic inheritance. The Ph mutants in the form of deletions are available in bread wheat (ph1b) and durum wheat (ph1c). Thus, ph1b-haploids of bread wheat and ph1c-haploids of durum wheat show extensive homoeologous pairing that has been shown by us and several others. Here we study the effect of ph1b allele of bread wheat on chromosome pairing in durum haploids, whereas we studied earlier the effect of ph1c allele in durum haploids that we synthesized. In durum wheat, the ph1b-haploids show much higher (49.4% of complement) pairing than the ph1c-haploids (38.6% of complement). Mention of a trademark or proprietary product does not constitute a guarantee or warranty of the product by the USDA or imply approval to the exclusion of other products that also may be suitable.     

2n gametes in potato species and their function in sexual polyploidization

Summary The extent and pattern of polyploidy in the tuber-bearing Solanums varies among the many taxanomic series that have been identified in this subsection of Solanum. While several series appear to be entirely diploid, others exhibit a range of ploidy levels from 2x to 6x, and some contain only polyploid species.In many diploid, triploid and tetraploid species 2n gametes (gametes or gametophytes with the sporophytic chromosome number) have been detected. Both 2n eggs and 2n pollen occur. 2n gametes provide the opportunity for unilateral and bilateral sexual polyploidization. The genetic determination and consequences of sexual polyploidization strongly suggest that 2n gametes have been the major instrument in the polyploid evolution of the tuber-bearing Solanums. Somatic doubling of species and interspecific hybrids appears to be of very limited importance.New evidence for the occurrence of 2n eggs and 2n pollen in many species is reported, and data from the literature are added to illustrate the widespread distribution of 2n gametes throughout the subsection. A very high correlation is found between polyploidy and 2n gametes, and its significance is discussed. Proof is presented for the occurrence of alleles governing 2n pollen production in the cultivated tetraploids, providing additional evidence for the hypothesis that 2n gametes have been involved in their origin.Multiple unilateral and bilateral sexual polyploidizations are proposed for the origin of the cultivated tetraploids: this accounts for the large variability encountered in this group, which closely resembles that of the related diploids. Similar evolutionary pathways are hypothesized for the other polyploid complexes in the subsection.A scheme is proposed in which participation of both 2n and n gametes link together all ploidy levels in the tuber-bearing Solanums, thus overcoming the ploidy barriers and providing for gene flow throughout the sympatric species of the subsection.Paper No. 2093 from the Laboratory of Genetics. Research supported by the College of Agricultural and Life Sciences, a gift from Frito-Lay, Inc., and grants from NIH (GM 15422) and The International Potato Center. Plant materials, unpublished chromosome counts and technical help of IR-1 are gratefully acknowledged.     

Hybrids between durum wheat and Thinopyrum junceiforme: Prospects for breeding for scab resistance

Tetraploid wheatgrass, Thinopyrum junceiforme(2n = 4x = 28; J₁J₁J₂J₂), a wild relative of wheat, is an excellent source of resistance to Fusarium head blight. Intergeneric F₁ hybrids (2n = 4x = 28; ABJ₁J₂) between durum wheat (Triticum turgidum; 2n = 4x = 28; AABB) cultivars Lloyd or Langdon and Th. junceiforme were synthesized. Most of the pairing in F₁ hybrids was between the J₁- and J₂-genome chromosomes. Some pairing occurred between wheat chromosomes and alien chromosomes, resulting in segmental exchange that was confirmed by fluorescent in situ hybridization (FISH). The F₁hybrids were largely male-sterile and were backcrossed, as the female parent, to the respective durum cultivar. Backcrosses from Lloyd × Th. junceiforme hybrids yielded fertile partial amphiploids (2n = 6x = 42; AABBJ₁J₂) as a result of functioning of unreduced female gametes of the hybrid. Lloyd proved to be a more useful durum parent than Langdon in crosses with Th. junceiforme designed to transfer scab resistance genes. Pairing in the amphiploids was characterized by preferential pairing,which resulted in bivalent formation. However, some intergeneric pairing also occurred. Several fertile hybrid derivatives were produced by further backcrossing and selfing. The introduction of alien chromatin into the durum complement was confirmed by FISH. Hybrid derivative lines had significantly lower mean infection scores (p = 0.01), the best showing 10.93% infection, whereas the parental durum cultivars had 70.34% to 89.46% infection. Hybridization with wild relatives may offer an excellent means of introducing scab resistance into durum wheat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Occurrence, inheritance and use of reproductive mutants in alfalfa improvement

The widespread occurrence of 2ngametes (i.e. gametes with the somatic chromosome number) in the Medicago sativa-coerulea-falcata complex supports the concept that gene flow from diploid to tetraploid species occurs continuously in nature and plays a key role in alfalfa evolution. Breeders realized early that gene transfer between ploidy levels via 2n pollen and 2n eggs would have had potential use in cultivated alfalfa improvement. Cytological investigations provided insights into the types of meiotic abnormalities responsible for the production of 2n gametes. Alterations were defined as genetically equivalent to first (FDR) or second division restitution(SDR) mechanisms. For breeding purposes,data have proven that 2n gametes of the FDR type are more advantageous than those obtained by SDR for transferring parental heterozygosity and retaining epistatic interactions. The use of diploid meiotic mutants that produce 2ngametes is now recognized as one of the most effective methods available for exploiting heterosis and introgressing wild germplasm traits into cultivated tetraploid alfalfa via unilateral (USP) and bilateral sexual polyploidization (BSP) schemes. Both2n egg and 2n pollen producers could be used for direct gene transfer from wild diploid relatives into cultivated alfalfa by means of 2x-4x and4x-2x crosses. Although data have shown that forage yield improvement can be achieved when plants are sexually tetraploidized, problems related to reduced plant fertility and seed production remain largely unexplained. Apomixis has the potential of cloning plants through seed and thus provides a unique opportunity for developing superior tetraploid cultivars with permanently fixed heterosis and epistatic effects. A main goal in alfalfa breeding could be the introduction of functional apomixis (i.e. Apomeiosis and parthenogenesis) in cultivated alfalfa stocks. In the future, the efficiency of alfalfa breeding programs based on the use of reproductive mutants could be improved by direct selection at the genotype level using RFLPs and PCR-based markers. Suitable DNA markers and detailed linkage maps of alfalfa mutants should help to discover apomictic mutants and address basic genetic issues such as the extent of genomicre combination in polyploid hybrids and the effect of sexual polyploidization on heterosis. Molecular markers have recently been used in alfalfa for studying the inheritance of 2n gamete formation and identifying polymorphisms associated to genes involved in meiotic abnormalities. Molecular tagging of 2n egg and 2n pollen formation not only should explain the genetic control and regulation of these traits, but may also be an essential step towards marker-assisted selection of 2n gamete producers and implementation of USP and BSP breeding schemes. Future perspectives include strategies for the map-based cloning of genomic DNA markers,and screening of EST mini-libraries related to flowers at different developmental stages from meiotic mutants and wild-type scan lead to the identification of mRNAs and thus of candidate genes that control 2n gamete formation in alfalfa. This revised version was published online in July 2006 with corrections to the Cover Date.     

Progenies of allotriploids of Oriental × Asiatic lilies (Lilium) examined by GISH analysis

With the aim of utilizing allotriploid (2n = 3x = 36) lily hybrids (Lilium) in introgression breeding, different types of crosses were made. First, using diploid Asiatic lilies (2n = 2x = 24), reciprocal crosses (3x − 2x and 2x − 3x) were made with allotriploid hybrids (AOA) obtained by backcrosses of F₁ Oriental × Asiatic hybrids (OA) to Asiatic cultivars (A). Secondly, the AOA allotriploids were crossed with allotetraploid (OAOA, 2n = 4x = 48), in 3x − 4x combination. Finally, the AOA allotriploids where crossed to 2n gamete producer F₁ OA hybrids (3x − 2x (2n)). Two types of triploids were used as parents in the different types of crosses, derived from: (a) mitotic polyploidization and (b) sexual polyploidization. Ploidy level of the progeny was determined by estimating the DNA values through flowcytometry as well as chromosome counting. The aneuploid progeny plants from 3x − 2x and reciprocal crosses had approximate diploid levels and in 3x − 4x crosses and 3x − 2x (2n) the progeny had approximate tetraploid levels. Balanced euploid gametes (x, 2x and 3x) were formed in the AOA genotypes. Recombinant chromosomes were found in the progenies of all crosses, except in the case of 2x − 3x crosses through genomic in situ hybridization (GISH) analyses. Recombinant chromosomes occurred in the F₁ OA hybrid when the triploid AOA hybrid was derived through sexual polyploidization, but not through mitotic polyploidization with two exceptions. Those recombinant chromosomes were transmitted to the progenies in variable frequencies.     

Fluorescent in situ hybridization as an aid to introducing alien genetic variation into wheat

Summary Fluorescent in situ hybridization (FISH) has been used to assess the occurrence and frequency of wheat-alien chromosome pairing in a wheat/Thinopyrum bessarabicum hybrid and in wheat/rye hybrids with different levels of chromosome pairing by examining pollen mother cells at metaphase I of meiosis. The use of FISH to identify the presence and size of alien chromatin in a wheat background is also demonstrated.The value of FISH as an aid to the introgression of alien genetic variation into wheat is discussed.Abbreviations FISH fluorescent in situ hybridization - GISH genomic in situ hybridization - PRINS primer-induced in situ hybridization     

Cytological analyses of hybrids and derivatives of hybrids between durum wheat and Thinopyrum bessarabicum,using multicolour fluorescent GISH*

The wheat progenitors and other wild relatives continue to be important sources of genes for agronomically desirable traits, which can be transferred into durum wheat (Triticum turgidum; 2n = 4x = 28; AABB genomes) cultivars via hybridization. Chromosome pairing in durum × alien species hybrids provides an understanding of genomic relationships, which is useful in planning alien gene introgression strategies. Two durum cultivars, ‘Lloyd’ and ‘Langdon’, were crossed with diploid wheatgrass, Thinopyrum bessarabicum (2n = 2x = 14; JJ), to synthesize F₁ hybrids (2n = 3x = 21; ABJ) with Ph1. ‘Langdon’ disomic substitution 5D(5B) was used as a female parent to produce F₁ hybrids without Ph1, which resulted in elevation of pairing between durum and grass chromosomes – an important feature from the breeding standpoint. The F₁ hybrids were backcrossed to respective parental cultivars and BC₁ progenies were raised. ‘Langdon’ 5D(5B) substitution × Th. bessarabicum F₁ hybrids were crossed with normal ‘Langdon’ to obtain BC₁ progeny. Chromosome pairing relationships were studied in F₁ hybrids and BC₁ progenies using both conventional staining and fluorescent genomic in situ hybridization (fl‐GISH) techniques. Multicolour fl‐GISH was standardized for characterizing the nature and specificity of chromosome pairing: A–B, A–J and B–J pairing. The A–J and B–J pairing will facilitate gene introgression in durum wheat. Multicolour fl‐GISH will help in characterizing alien chromosome segments captured in the durum complement and in their location in the A and/or B genome, thereby accelerating chromosome engineering research.     

Relevance of sexual polyploidization for crop improvement – A review

Colchicine induced polyploids have not directly contributed for crop improvement in the past. On the other hand, the so-called natural polyploids, derived from the functioning of numerically unreduced(2n) gametes have been shown to be more relevant for crop improvement in many cases. Different types of cytological abnormalities during meiosis can give rise to 2n gametes and the genetic composition of these gametes is variable. Depending on the type meiotic abnormalities, various types of 2ngametes, such as first division restitution(FDR), second division restitution (SDR),indeterminate meiotic restitution (IMR) and post meiotic restitution (PMR) gametes,among others, have been described in recent years. For the improvement of autopolyploids such as potato, alfalfa,Vaccinium spp., and some of the fodder grasses, FDR gametes have been proved to be highly useful. However, the use of 2n gametes for the improvement of allopolyploid crops has received much less attention so far. Some of the investigations on allopolyploids, derived from Festuca-Lolium, Alstroemeria and Lilium species hybrids, have revealed that 2ngametes can be most useful for the introgression of alien genes and chromosomes into cultivars. An important feature of using sexual polyploidization in the case of allopolyploids is that introgression can be achieved through recombination due to genetic crossing-over between alien chromosomes as well as addition of alien chromosomes, which is extremely difficult or impossible to achieve in the case of colchicine induced allopolyploids. Because of the recent developments in the field of plant molecular biology, methods have become available for the analysis of 2ngametes and sexual polyploid progenies more accurately and to develop systematic breeding approaches. The methods include DNA in situ hybridization (GISH and FISH)and molecular mapping (AFLP, RFLP, RAPDs).In addition to providing basic information on the genetic and genome composition of the polyploid progenies, these methods can be potentially useful for a more efficient creation of desirable breeding material and cultivars. This revised version was published online in July 2006 with corrections to the Cover Date.     

Screening <Emphasis Type="Italic">A. ventricosa</Emphasis> populations for 2n gametes

Sexual polyploidization via the formation of 2n gametes has been acknowledged as the most significant evolutionary mode of polyploidization among plant species. The present study was conducted in order to determine whether 2n gametes are present in the C-genome diploid Avena ventricosa Bal. ex Coss., a species that contributed to the evolution of the cultivated hexaploid species (Avena sativa L). Individual plants belonging to four different Cypriot populations, were screened for pollen grain size variation with the aim to distinguish 2n gametes. Avena ventricosa ARI00-845 was identified to produce large pollen grains at a low percentage (1.21%). Subsequent analysis using flow cytometry confirmed the presence of 2n gametes in the pollen. Cytogenetic analyses of pollen mother cells revealed cells with twice the typical chromosome number at metaphase I (i.e., 28 chromosomes). We postulate that irregularities in cell wall formation preceding meiosis could have contributed to the mode of chromosome doubling.     

Relevance of unilateral and bilateral sexual polyploidization in relation to intergenomic recombination and introgression in <Emphasis Type="Italic">Lilium</Emphasis> species hybrids

Sexual polyploids were induced in diploid (2n = 2x = 24) interspecific F1 hybrids of Longiflorum × Asiatic (LA) and Oriental × Asiatic (OA) Lilium hybrids by backcrossing to Asiatic (AA) parents as well as by sib-mating of the F1 LA hybrids. A majority of the BC1 progenies were triploid and the progenies from sib-mating were tetraploid or near tetraploids. Genomic in situ hybridization (GISH) technique was applied to assess the intergenomic recombination in the BC1 populations of LA and OA hybrids obtained after unilateral sexual polyploidization. A total of 63 LA (LA × AA and AA × LA) and 53 OA hybrids were analysed. LA hybrids were originated through the functioning of 2n gametes either as 2n eggs or 2n pollen while those of OA hybrids originated through functional 2n pollen of diploid OA genotype. In both type of crosses, a majority of the progenies had originated through First Division Restitution (FDR) mechanism of functional 2n gamete either with or without a cross over. However, there were nine LA- and four OA-genotypes where Indeterminate Meiotic Restitution (IMR) was the mechanism of 2n gamete formation. Based on GISH, total amount of introgression of Longiflorum and Oriental genome into Asiatic genome was determined. Most of the BC progenies exhibited recombination and the amount of recombination was higher in LA hybrids as compared to OA hybrids. Intergenomic recombination was also determined cytologically in the 16 plants of sib-mated LA hybrids where both parents had contributed 2n gametes. Based on these results the nature of interspecific lily hybrids obtained from uni- and bilateral sexual polyploidization leading to allotriploid and allotetraploid formation is discussed in the context of introgression and intergenomic recombination.     

Spontaneous wheat/rye translocations from female meiotic products of hybrids between octoploid triticale and wheat

Summary Heptaploid hybrids between octoploid triticale and wheat were backcrossed as female parents with wheat to examine the rye chromosome distribution in the resultant progenies using genomic in situ hybridization (GISH). One hundred and one backcross (BC) seeds were examined and whole rye chromosome additions and substitutions, wheat/rye centric and noncentric translocations and rye telocentric chromosomes were detected. Dicentric wheat/rye translocated chromosomes were also observed. Comparisons were made with previous results on the rye chromosome distribution from male gametes of the same cross and differences were found, where in the female derived population a deficit of plants with more than two rye chromosomes was apparent relative to the anther derived population.     

Occurrence of numerically unreduced (2n) gametes in Alstroemeria interspecific hybrids and their significance for sexual polyploidisation

The F1 hybrids of seven diploid Alstroemeria species (2n=2x=16) were investigated for the production of numerically unreduced (2n) gametes and their mode of origin. Based on a survey of 17 interspecific hybrid combinations,consisting of 119 genotypes, it was found that the F1 hybrids of Chilean-Brazilian species mostly produced first division restitution (FDR) 2n gametes. These F1 hybrids were self-pollinated in order to obtain F2 seeds, which was an indication that the F1 plants also produced 2neggs simultaneously. All the F2 progeny plants were typical allotetraploids, most of which formed 16 bivalents and a small proportion formed multivalents during metaphase I stages of meiosis. Through genomic in situ hybridisation (GISH) it was proved that multivalent formation in F2plants, derived from A. inodora ×A. pelegrina hybrid, was due to homoeologous recombination but not from reciprocal translocations. In order to test the segregation pattern of the recombinant chromosomes, an F3 population from one genotype, P6C49-6, was investigated. The recombinant chromosomes assorted independently from each other supporting the hypothesis that the segregation of chromosomes in ring quadrivalents did not behave like those in translocation heterozygotes. It was concluded that in allopolyploids of Alstroemeria,bilateral sexual polyploidisation could accomplish genetic recombination by both homoeologous crossing-over as well as through the assortment of chromosomes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Polyploidy and sexual polyploidization in the genus Vaccinium

Genus Vaccinium,consisting of blueberries, cranberries, lingonberries and many related wild species, includes diploid, tetraploid, and hexaploid species. Most evidence indicates that the tetraploid species are autotetraploids, with non-preferential bivalent chromosome pairing. Although homoploid interspecific crosses usually produce numerous fertile hybrids if the parents are from the same section of the genus, inter sectional crosses at the diploid level normally produce no seedlings, weak seedlings, or seedlings that are very low in fertility. There is a strong but not complete triploid block within Vaccinium. Even with insections, tetraploid × diploid (and the reciprocal) crosses normally give only a few tetraploid and a few triploid hybrids. Hexaploid × diploid crosses within sections are very hard to make, and the few hybrids that have been obtained are pentaploid. The frequency of 2n gametes varies,both among genotypes within species and among species. Vaccinium pollen is normally shed in tetrads, and the frequency of large pollen grains shed in dyads can be used to estimate 2n gamete frequency. Cultivated blueberries occur at both the tetraploid and the hexaploid levels, and there are important genetic resources in the diploids. Unreduced gamete production has permitted transfer of genetic material from the diploid to the tetraploid level and from the diploids and tetraploids to the hexaploid level via triploid hybrids. Intersectional crosses can occasionally produce tetraploid Vaccinium hybrids that appear to behave as amphidiploids and have medium to high fertility. CommercialVaccinium cultivars are normally propagated by cuttings. Intersectional hybridization, chromosome doubling, and asexual propagation could permit the production of novel hybrid combinations with value as ornamentals or in fruit production. This revised version was published online in July 2006 with corrections to the Cover Date.     

Gene transfer from non-tuberous to tuberous Solanum species: Evidence from meiosis in hybrids

Summary The distant hybrids between non-tuberous Solanum species and tuberous S. pinnatisectum display little or no pairing in F1 and predominantly bivalent formation (preferential pairing) after chromosome doubling. In such a situation the question about the potential and extent of gene transfer from the non-tuberous parent to the tuberous one is relevant to potato breeding. This question was investigated by studying meiosis in triploid and hexaploid hybrids from crosses between diploid TV⁵ x tetraploid (S. etuberosum x S. pinnatisectum). TV⁵ is similar to S. verrucosum with cytoplasm of S. tuberosum. The following evidence was found for the desirable transfer of S. etuberosum genes to the tuberous species.The triploid F1 hybrids did not display the configurations 12 II+12 I expected if no gene exchange would take place between S. etuberosum and the tuberous species; however, a considerable number of multivalents per cell was observed in all plants studied.In the hexaploid F1 hybrids, obtained from the triploids through somatic doubling in vitro, 36 bivalents could reasonably be expected. Although bivalents were predominant (an overall average of 24.2 per cell) quite a few chromosomes were associated as multivalents in all plants investigated.It is concluded that in the hybrids studied a considerable amount of pairing and chiasma formation occurs between chromosomes of non-tuberous and those of tuberous Solanum species. This pairing affinity is larger than that found in 2x and 4x hybrids from S. etuberosum x S. pinnatisectum. Some hypotheses are put forward to explain this increased pairing affinity.     

Transfer of resistance to race 2 of Plasmodiophora brassicae from Brassica napus to cabbage (B. oleracea ssp. capitata). III. First backcross and F2 progenies from interspecific hybrids between B. napus and B. oleracea ssp. capitata

Summary The first backcross and F₂ progenies from triploid F₁ and tetraploid F₁ hybrids between B. napus and 2x and 4x B. oleracea ssp. capitata (cabbage) were studied for their general morphology, resistance to race 2 of the clubroot pathogen, chromosome number and meiotic chromosome behavior. No linkage was apparent between resistance and the major morphological characters. Unreduced gametes played a large part in the successful formation of seed of the B₁ and F₂ progeny. B₁ plants with low chromosome numbers were selected for use in recurrent backcrosses. The potential use of anther culture to extract gametic progenies from resistant B₁ and F₂ plants with higher chromosome numbers was suggested. The presence of homoeologous pairing observed in all the plants is considered advantageous for selecting suitable progeny in later generations.     

Fluorescent in situ hybridization — a useful aid to the introduction of alien genetic variation into wheat

Summary Fluorescent in situ hybridization (FISH) of DNA to plant chromosomes has proved to be a powerful cytogenetic tool. The value of fluorescent in situ hybridization of total genomic DNA (GISH) of related species is demonstrated in the determination of wheat/alien chromosome pairing in hybrids. Its use for assessing the relative merits of the various genes that affect chromosome pairing is also shown.The ability of GISH to identify the presence in wheat of whole alien chromosomes or alien chromosome segments is illustrated. The potential of FISH for detecting repeated DNA sequences, low copy sequences and single copy genes is discussed.Abbreviations FISH fluorescent in situ hybridization - GISH genomic in situ hybridization - PRINS primer-induced in situ hybridization     

Formation of unreduced gametes is impeded by homologous chromosome pairing in tetraploid <Emphasis Type="Italic">Triticum turgidum</Emphasis> × <Emphasis Type="Italic">Aegilops tauschii</Emphasis> hybrids

It is believed that unreduced gametes with somatic chromosome numbers play a predominant role in natural polyploidization. Allohexaploid bread wheat originated from spontaneous hybridization of Triticum turgidum L. with Aegilops tauschii Coss. Unreduced gametes originating via meiotic restitution, including first-division restitution (FDR) and single-division meiosis (SDM), are well documented in triploid F₁ hybrids of T. turgidum with diploid Ae. tauschii (genomic constitution ABD, usually with 21 univalents in meiotic metaphase I). In this study, two T. turgidum lines known to carry genes for meiotic restitution were crossed to tetraploid Ae. tauschii. The resulting F₁ hybrids (genomes ABDD), had seven pairs of homologous chromosomes and regularly formed 14 univalents and seven bivalents at metaphase I. Neither FDR nor SDM were observed. The distribution of chromosome numbers among progeny obtained by self pollination and a backcross to T. turgidum showed the absence of unreduced gametes. These results suggest that high homologous pairing interfered with meiotic restitution and the formation of unreduced gametes. This may be related to asynchronous movement during meiosis between paired and unpaired chromosomes or to uneven distribution of chromosomes in anaphases, resulting in nonviable gametes.