Durum wheat as a candidate for the unknown female progenitor of bread wheat: an empirical study with a highly fertile F<Subscript>1</Subscript> hybrid with <Emphasis Type="Italic">Aegilops tauschii</Emphasis> Coss.

Hexaploid bread wheat was derived from a hybrid cross between a cultivated form of tetraploid Triticum wheat (female progenitor) and a wild diploid species, Aegilops tauschii Coss. (male progenitor). This cross produced a fertile triploid F₁ hybrid that set hexaploid seeds. The identity of the female progenitor is unknown, but various cultivated tetraploid Triticum wheats exist today. Genetic and archaeological evidence suggests that durum wheat (T. turgidum ssp. durum) may be the female progenitor. In previous studies, however, F₁ hybrids of durum wheat crossed with Ae. tauschii consistently had low levels of fertility. To establish an empirical basis for the theory of durum wheat being the female progenitor of bread wheat, we crossed a durum wheat cultivar that carries a gene for meiotic restitution with a line of Ae. tauschii. F₁ hybrids were produced without using embryo rescue techniques. These triploid F₁ hybrids were highly fertile and spontaneously set hexaploid F₂ seeds at the average selfed seedset rate of 51.5%. To the best of our knowledge, this is the first example of the production of highly fertile F₁ hybrids between durum wheat and Ae. tauschii. The F₁ and F₂ hybrids are both similar morphologically to bread wheat and have vigorous growth habits. Cytological analyses of F₁ male gametogenesis showed that meiotic restitution is responsible for the high fertility of the triploid F₁ hybrids. The implications of these findings for the origin of bread wheat are discussed.     

Origin,evolution and genome differentiation in Guizotia abyssinica and its wild species

Summary Genome affinities were analyzed at meiosis in C-banded metaphase-I cells of wheat x Ae. Sharonensis hybrid plants. The results showed that the most frequent type of pairing occurred between chromosomes of the A and D genomes in all plants, as well as in cells with different numbers of associations. These findings clearly indicated that Ae. Sharonensis can be excluded as the donor of the B genome of wheat.     

Production of intergeneric hybrid between dwarfing polish wheat (<Emphasis Type="Italic">Triticum polonicum</Emphasis> L.) and <Emphasis Type="Italic">Aegilops tauschii</Emphasis> Cosson. with reference to wheat origin

Dwarfing polish wheat is a dwarfing accession of Triticum polonicum L. from Xinjiang of China. In the present study, the artificial hybridization between dwarfing polish wheat and two accessions of Aegilops tauschii Cosson. (AS60 and AS65) was carried out, and the F₁ hybrids were obtained successfully without using embryo rescue techniques for the first time. The crossabilities of hybrids T. polonicum × Ae. tauschii (AS60) and T. polonicum × Ae. tauschii (AS65) were 1.67% and 0.60% respectively. Only the hybrids of T. polonicum × Ae. tauschii (AS60) germinated well, and 24 F₁ hybrid plants were obtained. All the F₁ hybrid plants grew vigorously, and the morphological traits were similar to bread wheat. The F₁ plants had some obvious traits inherited from T. polonicum and Ae. tauschii and were completely sterile. Chromosome pairing in the hybrid was characterized by a large number of univalents, with an average of 20.56 and 0.22 bivalents per PMC, and no ring bivalents and multivalents were observed. Furthermore, the potential value of the F1 hybrids between T. polonicum and Ae. tauschii for studying wheat origin and breeding are discussed. The article is published in the original.     

Genome relationships between Hordeum vulgare L. and H. bulbosum L.

Interrelationships between H. vulgare (2x=14) and H. bulbosum (2x=14; 4x=28) were estimated on the basis of the karyotypes and the pairing behaviour of the chromosomes in diploid, triploid and tetraploid hybrids obtained with the aid of embryo culture. — A comparison of the karyotypes of the two species revealed similarities as well as differences. It was concluded that at least 4 or more of the chromosomes were similar in morphology and probably closely related. — Diploid and tetraploid hybrids are rarely obtained and their chromosome numbers tend to be unstable whereas triploid hybrids (1 vulgare + 2 bulbosum genomes) were stable and relatively easy to produce. In the diploid hybrid only 40% of the meiotic cells contained 14 chromosomes while the numbers ranged from 7 to 16 in other cells. All hybrids exhibited pairing between the chromosomes of the two species. Diploid hybrids had a mean of 5.0 and a maximum of 7 bivalents per cell in those cells having 14 chromosomes. Triploid hybrids from crosses between 2x H. vulgare and 4x H. bulbosum exhibited a mean of 1.5 and a maximum of 5 trivalents per cell. In a hexaploid sector found following colchicine treatment of a triploid the mean frequencies of chromosome associations per cell were: 5.5_I+8.0_II+0.7_III+3.7_IV+0.3_V+0.4_VI. One unstable 27 chromosome hybrid obtained from crosses between the autotetraploid forms had a mean of 1.1 and a maximum of 4 quadrivalents per cell. The chromosome associations observed in these hybrids are consistent and are taken as evidence of homoeologous pairing between the chromosomes of the two species. Interspecific hybridization between these two species also reveals that chromosome stable hybrids are only obtained when the genomes are present in a ratio of 1 vulgare2 bulbosum. Based upon the results obtained, the possibility of transferring genetic characters from H. bulbosum into cultivated barley is discussed.     

Natural variation for fertile triploid F1 hybrid formation in allohexaploid wheat speciation

The tempo, mode, and geography of allopolyploid speciation are influenced by natural variation in the ability of parental species to express postzygotic reproductive phenotypes that affect hybrid fertility. To shed light on the impact of such natural variations, we used allohexaploid Triticum aestivum wheats’ evolution as a model and analyzed the geographic and phylogenetic distributions of Aegilops tauschii (diploid progenitor) accessions involved in the expression of abnormality and fertility in triploid F₁ hybrids with Triticum turgidum (tetraploid progenitor). Artificial-cross experiments and chloroplast-DNA-based evolutionary analyses showed that hybrid-abnormality-causing accessions had limited geographic and phylogenetic distributions, indicative that postzygotic hybridization barriers are underdeveloped between these species. In contrast, accessions that are involved with fertile triploid F₁ hybrid formation have wide geographic and phylogenetic distributions, indicative of a deep evolutionary origin. Wide-spread hybrid-fertilizing accessions support the theory that T. aestivum speciation occurred at multiple sites within the species range of Ae. tauschii, in which existing conditions enabled natural hybridization with T. turgidum. Implications of our findings on how natural variation in the ability of Ae. tauschii to express those postzygotic reproductive phenotypes diversified and contributed to the speciation of T. aestivum are discussed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Analysis of metaphase I chromosome association in species of the genus Aegilops

Summary Metaphase-I chromosome associations in every diploid and polyploid species of the genus Aegilops were studied using C-banding in order to analyse the cytogenetic behaviour of the whole complement as well as of specific genomes in different polyploid species. Differences were observed in the frequency of associations per cell among different species of the same ploidic level and even between species sharing the same genomic constitution. Differences were also found between different genomes within the same polyploid species and between the same genome when present in several diploid and polyploid species. Several factors proposed as having an influence on the frequency of metaphase-I associations, such as chromosome morphology, C-heterochromatin content, genetic control and genome interactions, are discussed. Most of the polyploid Aegilops species showed a diploid-like behaviour at metaphase I although multivalents involving homoeologous associations were occasionally observed in Ae. biuncialis, Ae. juvenalis and Ae. crassa(6x); therefore, the Aegilops diploidising genetic system is not equally effective in all polyploid species.     

BAC libraries of Triticum urartu, Aegilops speltoides and Ae. tauschii, the diploid ancestors of polyploid wheat

Triticum urartu, Aegilops speltoides and Ae. tauschii are respectively the immediate diploid sources, or their closest relatives, of the A, B and D genomes of polyploid wheats. Here we report the construction and characterization of arrayed large-insert libraries in a bacterial artificial chromosome (BAC) vector, one for each of these diploid species. The libraries are equivalent to 3.7, 5.4 and 4.1 of the T. urartu, Ae. speltoides, Ae. tauschii genomes, respectively. The predicted levels of genome coverage were confirmed by library hybridization with single-copy genes. The libraries were used to estimate the proportion of known repeated nucleotide sequences and gene content in each genome by BAC-end sequencing. Repeated sequence families previously detected in Triticeae accounted for 57, 61 and 57% of the T. urartu, Ae. speltoides and Ae. tauschii genomes, and coding regions accounted for 5.8, 4.5 and 4.8%, respectively.     

Variability of ribosomal DNA sites inFestuca pratensis, Lolium perenne, and their intergeneric hybrids, revealed by FISH and GISH

This study focuses on the variability of chromosomal location and number of ribosomal DNA (rDNA) sites in some diploid and autotetraploidFestuca pratensis andLolium perenne cultivars, as well as on identification of rDNA-bearing chromosomes in their triploid and tetraploidF. pratensis ×L. perenne hybrids. The rDNA loci were mapped using fluorescence in situ hybridization (FISH) with 5S and 25S rDNA probes, and the origin of parental genomes was verified by genomic in situ hybridization (GISH) withL. perenne genomicDNAas a probe, andF. pratensis genomic DNA as a block. FISH detected variation in the number and chromosomal location of both 5S and 45S rDNA sites. InF. pratensis mostly additional signals of 5S rDNA loci occurred, as compared with standardF. pratensis karyotypes. Losses of 45S rDNA loci were more frequent inL. perenne cultivars and intergeneric hybrids. Comparison of theF. pratensis andL. perenne genomes approved a higher number of rDNA sites as well as variation in chromosomal rDNA location inL. perenne. A greater instability ofF. pratensis-genome-like andL. perenne-genome-like chromosomes in tetraploid hybrids was revealed, indicating gains and losses of rDNA loci, respectively. Our data indicate that the rDNA loci physically mapped on chromosomes 2 and 3 inF. pratensis and on chromosome 3 inL. perenne are useful markers for these chromosomes in intergenericFestuca ×Lolium hybrids.     

<Emphasis Type="Italic">Aegilops-Secale</Emphasis> amphiploids: chromosome categorisation,pollen viability and identification of fungal disease resistance genes

The aim of this study was to assess the potential breeding value of goatgrass-rye amphiploids, which we are using as a “bridge” in a transfer of Aegilops chromatin (containing, e.g. leaf rust resistance genes) into triticale. We analysed the chromosomal constitution (by genomic in situ hybridisation, GISH), fertility (by pollen viability tests) and the presence of leaf rust and eyespot resistance genes (by molecular and endopeptidase assays) in a collection of 6× and 4× amphiploids originating from crosses between five Aegilops species and Secale cereale. In the five hexaploid amphiploids Aegilops kotschyi × Secale cereale (genome UUSSRR), Ae. variabilis × S. cereale (UUSSRR), Ae. biuncialis × S. cereale (UUMMRR; two lines) and Ae. ovata × S. cereale (UUMMRR), 28 Aegilops chromosomes were recognised, while in the Ae. tauschii × S. cereale amphiploid (4×; DDRR), only 14 such chromosomes were identified. In the materials, the number of rye chromosomes varied from 14 to 16. In one line of Ae. ovata × S. cereale, the U-R translocation was found. Pollen viability varied from 24.4 to 75.4%. The leaf rust resistance genes Lr22, Lr39 and Lr41 were identified in Ae. tauschii and the 4× amphiploid Ae. tauschii × S. cereale. For the first time, the leaf rust resistance gene Lr37 was found in Ae. kotschyi, Ae. ovata, Ae. biuncialis and amphiploids derived from those parental species. No eyespot resistance gene Pch1 was found in the amphiploids.     

Karyotypes,nucleoli, and amphiplasty in hybrids between Hordeum vulgare L. and H. bulbosum L.

Chromosome measurements were carried out in Hordeum vulgare, H. bulbosum, and their diploid, triploid, and tetraploid hybrids. The chromosomes were classified by using relative values, and thus karyotypes were established. For comparison of these karyotypes both relative and absolute values were used. It was concluded that differential amphiplasty occurred, whereas neutral amphiplasty could not be demonstrated. In the hybrids the relative length of the parts of the chromosomes (long arm, short arm, satellite) was not changed in comparison with these lengths in the pure species. The karyotypes of both species had considerable similarities. From comparing the mean absolute genome lengths, it was, however, concluded that in the pure species, as well as in all hybrid types, the chromosomes of H. vulgare were longer than those of H. bulbosum. In the diploid and tetraploid hybrids the mean genome lengths were shorter than those in the pure species and the triploid hybrids. The differential amphiplasty was such that the secondary constriction of chromosome 6 of H. bulbosum, did not show up in the hybrids. This could be related to the suppression of nucleolar formation in the genome of H. bulbosum, because the maximum number of nucleoli in root tip cells equalled the number of satellite chromosomes. Finally it was found that the pattern of nucleolar fusion in diploid and triploid hybrids deviated from the expectation. The results were discussed in relation to chromosomal disturbances that occurred in the hybrid tissues and that resulted in elimination of chromosomes and other effects.     

Introgression of wheat DNA markers from A, B and D genomes in early generation progeny of Aegilops cylindrica Host × Triticum aestivum L. hybrids

Introgression from allohexaploid wheat (Triticum aestivum L., AABBDD) to allotetraploid jointed goatgrass (Aegilops cylindrica Host, CCDD) can take place in areas where the two species grow in sympatry and hybridize. Wheat and Ae. cylindrica share the D genome, issued from the common diploid ancestor Aegilops tauschii Coss. It has been proposed that the A and B genome of bread wheat are secure places to insert transgenes to avoid their introgression into Ae. cylindrica because during meiosis in pentaploid hybrids, A and B genome chromosomes form univalents and tend to be eliminated whereas recombination takes place only in D genome chromosomes. Wheat random amplified polymorphic DNA (RAPD) fragments, detected in intergeneric hybrids and introgressed to the first backcross generation with Ae. cylindrica as the recurrent parent and having a euploid Ae. cylindrica chromosome number or one supernumerary chromosome, were assigned to wheat chromosomes using Chinese Spring nulli-tetrasomic wheat lines. Introgressed fragments were not limited to the D genome of wheat, but specific fragments of A and B genomes were also present in the BC1. Their presence indicates that DNA from any of the wheat genomes can introgress into Ae. cylindrica. Successfully located RAPD fragments were then converted into highly specific and easy-to-use sequence characterised amplified regions (SCARs) through sequencing and primer design. Subsequently these markers were used to characterise introgression of wheat DNA into a BC1S1 family. Implications for risk assessment of genetically modified wheat are discussed.     

OBSERVATIONS OF CHROMOSOMES IN CIRCAEA (ONAGRACEAE)

Observations of meiotic and/or mitotic chromosomes for 16 specific, infraspecific, and hybrid taxa from 56 populations are presented; chromosome numbers for 11 taxa are reported for the first time. All samples reported have 2n = 22, with the exception that 3 of 12 collections of C. × intermedia have at least some individuals that are triploid. North American diploid collections of this hybrid are heterozygous for one reciprocal translocation; only 1 of 5 European diploid collections also exhibited a translocation. Meiosis in four naturally occurring Asian hybrids is also analyzed: C. × ovata (C. cordata × C. mollis), C. × decipiens (C. erubescens × C. lutetiana) and C. x× dubia (C. cordata × C. eurbescens) possess a single translocation; C. × mentiens (C. alpina × C. eurbescens) is structurally homozygous. With the exception of the translocation, chromosome pairing in these hybrids is normal. Early diakinesis chromosomes possess densely staining centric regions and diffusely staining arms that are subequal in length.     

GENOME ANALYSIS OF AGROPYRON REPENS × AGROPYRON CRISTATUM SYNTHETIC HYBRIDS

Hexaploid A. repens, 2n = 42, and diploid A. cristatum, 2n = 14, were hybridized and gave rise to two 28-chromosome reciprocal hybrids. Approximately 1% of hand-emasculated florets of both parent species produced viable hybrid seed following controlled pollination. Early embryo abortion prevented greater hybrid seed set on A. repens, whereas failure of fertilization appeared to be the major cause of poor hybrid seed set on A. cristatum. Reciprocal differences in hybrid vegetative and spike morphology were striking. The A. repens × A. cristatum hybrid was vigorous, highly rhizomatous, and bore abundant spikes whose morphology was intermediate between that of the parent species. A. cristatum × A. repens hybrids were weak, non-rhizomatous with frequently-malformed spikes. Mean chromosome associations of 0.10 I, 20.10 II, and 0.43 IV were observed in 134 metaphase-I cells of A. repens. Subsequent meiotic stages were regular except for occasional laggards and bridges at anaphase I and II. Metaphase-I chromosome associations averaged 0.07 I and 6.97 II in 124 A. cristatum cells. Chromosome pairing in the hybrids was highly variable and averaged 11.45 I, 7.58 II, 0.44 III, and 0.02 IV per cell in 187 cells interpreted. From 5 to 14 laggards appeared in every hybrid cell at anaphase I. Bridges were observed in approximately 25% of the anaphase-I cells. Similar irregularities were observed at anaphase II. Pollen viability was estimated as 3%, and the hybrids failed to set viable seed. On the basis of chromosome pairing in the species itself and in the hybrids, A. repens was designated as a segmental autoallohexaploid with a genome formula of the type A₁A₁A₂A₂BB. Although A. repens and A. cristatum chromosomes paired occasionally, the genomes of the 2 species were essentially non-homologous. Some of the interpretational difficulties of genome analysis were discussed.     

Identification and genetic characterization of an <Emphasis Type="Italic">Aegilops tauschii</Emphasis> ortholog of the wheat leaf rust disease resistance gene <Emphasis Type="Italic">Lr1</Emphasis>

Aegilops tauschii (goat grass) is the progenitor of the D genome in hexaploid bread wheat. We have screened more than 200 Ae. tauschii accessions for resistance against leaf rust (Puccinia triticina) isolates, which are avirulent on the leaf rust resistance gene Lr1. Approximately 3.5% of the Ae. tauschii accessions displayed the same low infection type as the tester line Thatcher Lr1. The accession Tr.t. 213, which showed resistance after artificial infection with Lr1 isolates both in Mexico and in Switzerland, was chosen for further analysis. Genetic analysis showed that the resistance in this accession is controlled by a single dominant gene, which mapped at the same chromosomal position as Lr1 in wheat. It was delimited in a 1.3-cM region between the restriction fragment length polymorphism (RFLP) markers ABC718 and PSR567 on chromosome 5DL of Ae. tauschii. The gene was more tightly linked to PSR567 (0.47 cM) than to ABC718 (0.79 cM). These results indicate that the resistance gene in Ae. tauschii accession Tr.t. 213 is an ortholog of the leaf rust resistance gene Lr1 of bread wheat, suggesting that Lr1 originally evolved in diploid goat grass and was introgressed into the wheat D genome during or after domestication of hexaploid wheat. Compared to hexaploid wheat, higher marker polymorphism and recombination frequencies were observed in the region of the Lr1 ortholog in Ae. tauschii. The identification of Lr1^Ae, the orthologous gene of wheat Lr1, in Ae. tauschii will allow map-based cloning of Lr1 from this genetically simpler, diploid genome.Hong-Qing Ling and Jiwen Qiu have contributed equally to this work     

Meiosis and pollen mitosis in diploid and triploid Colocasia antiquorum Schott.

The relationship between diploid and triploid forms of Colocasia antiquorum Schott. was assessed through comparative meiotic and pollen mitotic studies. Owing to poor spreading of the chromosomes of both materials, karyological observations on pachytene nuclei were limited to a few chromosomes. Among the two nucleolar chromosomes and a metacentric, telochromomere-bearing chromosome of the diploid, the latter and one of the nucleolar chromosomes characterized by a heteropycnotic short arm were identified in both bivalent and trivalent associations in the triploid. The homologues in these cases were homomorphic and intimately paired. Two types of heteromorphic bivalents exhibiting partial pairing of homomorphic segments were also recorded in the triploid. Among the 14 bivalents of the diploid at diakinesis, two were nucleolus-associated. In the triploid, chromosomal associations at diakinesis included trivalents (2 to 9), bivalents and univalents, and the chiasma frequency per paired chromosome was lower than in the diploids. In 21.6 percent of the PMCs at this stage intragenomic pairing of one or two chromosomes was observed. Post-diakinesis stages in the diploid were regular while in the triploid they were marked by various irregularities in a majority of the cells. However, fertility (stainability), size and divisional frequency of pollen in both materials were remarkably similar. Chromosome numbers in pollen nuclei in the triploid ranged from 8 to 25. Based on these data an autopolyploid origin for the triploid Colocasia and a lower base number than the gametic chromosome number for this genus are advanced.     

Genome differentiation in Aegilops. 3. Evolution of the D-genome cluster

 Six polyploid Aegilops species containing the D genome were studied by C-banding and fluorescence in situ hybridization (FISH) using clones pTa71 (18S-5.8S-26S rDNA), pTa794 (5S rDNA), and pAs1 (non-coding repetitive DNA sequence) as probes. The C-banding and pAs1-FISH patterns of Ae. cylindrica chromosomes were identical to those of the parental species. However, inactivation of the NOR on chromosome 5D with a simultaneous decrease in the size of the pTa71-FISH site was observed. The N^v and D^v genomes of Ae. ventricosa were somewhat modified as compared with the N genome of Ae. uniaristata and the D genome of Ae. tauschii. Modifications included minor changes in the C-banding and pAs1-FISH patterns, complete deletion of the NOR on chromosome 5D^v, and the loss of several minor 18S-5.8S-26S rDNA loci on N^v genome chromosomes. According to C-banding and FISH analyses, the D^cr1 genome of Ae. crassa is more similar to the D^v genome of Ae. ventricosa than to the D genome of Ae. tauschii. Mapping of the 18S-5.8S-26S rDNA and 5S rDNA loci by multicolor FISH suggests that the second (X^cr) genome of tetraploid Ae. crassa is a derivative of the S genome (section Emarginata of the Sitopsis group). Both genomes of Ae. crassa were significantly modified as the result of chromosomal rearrangements and redistribution of highly repetitive DNA sequences. Hexaploid Ae. crassa and Ae. vavilovii arose from the hybridization of chromosomal type N of tetraploid Ae. crassa with Ae. tauschii and Ae. searsii, respectively. Chromosomal type T1 of tetraploid Ae. crassa and Ae. umbellulata were the ancestral forms of Ae. juvenalis. The high level of genome modification in Ae. juvenalis indicates that it is the oldest hexaploid species in this group. The occurrence of hexaploid Ae. crassa was accompanied by a species-specific translocation between chromosomes 4D^cr1 and 7X^cr. No chromosome changes relative to the parental species were detected in Ae. vavilovii, however, its intraspecific diversity was accompanied by a translocation between chromosomes 3X^cr and 3D^cr1. Received July 24, 2001 Accepted October 1, 2001     

Pairing and recombination at meiosis of Brassica rapa (AA) × Brassica napus (AACC) hybrids

Interspecific crosses contribute significantly to plant evolution enabling gene exchanges between species. The efficiency of interspecific crosses depends on the similarity between the implicated genomes as high levels of genome similarity are required to ensure appropriate chromosome pairing and genetic recombination. Brassica napus (AACC) is an allopolyploid, resulting from natural hybridization between Brassica rapa (AA) and Brassica oleracea (CC), both being diploid species derived from a common ancestor. To study the relationships between genomes of these Brassica species, we have determined simultaneously the pairing and recombination pattern of A and C chromosomes during meiosis of AAC triploid hybrids, which result from the interspecific cross between natural B. napus and B. rapa. Different AAC triploid hybrids and their progenies have been analysed using cytogenetic, BAC-FISH, and molecular techniques. In 71% of the pollen mother cells, homologous A chromosomes paired regularly, and usually one chromosome of each pair was transmitted to the progeny. C chromosomes remained mainly univalent, but were involved in homoeologous pairing in 21.5% of the cells, and 13% of the transmitted C chromosomes were either recombined or broken. The rate of transmission of C chromosomes depended on the identity of the particular chromosome and on the way the hybrid was crossed, as the male or as the female parent, to B. napus or to B. rapa. Gene transfers in triploid hybrids are favoured between A genomes of B. rapa and B. napus, but also occur between A and C genomes though at lower rates.     

The formation of diploid and triploid hybrids of female grass carp × male blunt snout bream and their 5S rDNA analysis

Background

Hybridization is a useful strategy to alter the genotypes and phenotypes of the offspring. It could transfer the genome of one species to another through combing the different genome of parents in the hybrid offspring. And the offspring may exhibit advantages in growth rate, disease resistance, survival rate and appearance, which resulting from the combination of the beneficial traits from both parents.

Results

Diploid and triploid hybrids of female grass carp (Ctenopharyngodon idellus, GC, Cyprininae, 2n?=?48)?×?male blunt snout bream (Megalobrama amblycephala, BSB, Cultrinae, 2n?=?48) were successfully obtained by distant hybridization. Diploid hybrids had 48 chromosomes, with one set from GC and one set from BSB. Triploid hybrids possessed 72 chromosomes, with two sets from GC and one set from BSB.The morphological traits, growth rates, and feeding ecology of the parents and hybrid offspring were compared and analyzed. The two kinds of hybrid offspring exhibited significantly phenotypic divergence from GC and BSB. 2nGB hybrids showed similar growth rate compared to that of GC, and 3nGB hybrids significantly higher results. Furthermore, the feeding ecology of hybrid progeny was omnivorous.The 5S rDNA of GC, BSB and their hybrid offspring were also cloned and sequenced. There was only one type of 5S rDNA (designated type I: 180 bp) in GC and one type of 5S rDNA (designated type II: 188 bp) in BSB. However, in the hybrid progeny, diploid and triploid hybrids both inherited type I and type II from their parents, respectively. In addition, a chimera of type I and type II was observed in the genome of diploid and triploid hybrids, excepting a 10 bp of polyA insertion in type II sequence of the chimera of the diploid hybrids.

Conclusions

This is the first report of diploid and triploid hybrids being produced by crossing GC and BSB, which have the same chromosome number. The obtainment of two new hybrid offspring has significance in fish genetic breeding. The results illustrate the effect of hybridization and polyploidization on the organization and variation of 5S rDNA in hybrid offspring.

     

The pattern of zygotene and pachytene pairing in allotetraploid Aegilops species sharing the D genome

Chromosome pairing behaviour of the allotetraploid Aegilops species sharing the D genome, Ae. crassa (DDMM), Ae. cylindrica (DDCC) and Ae. ventricosa (DDNN), was analyzed by electron microscopy in surfacespread prophase-I nuclei. Synaptonemal-complex analysis at zygotene and pachytene revealed that synapsis in the allotetraploids was mostly between homologous chromosomes, although a few multivalents were also formed. Only homologous bivalents were observed at metaphase-I. It is concluded that the mechanism controlling bivalent formation in these species acts mainly at zygotene by restricting pairing to homologous chromosomes, but also acts at pachytene by preventing chiasma formation in homoeologous associations. These observations are discussed in relation to mechanisms of diploidization of polyploid meiosis.