A taxonomìc revision of Hordeum sect. Critesion

Eight taxa are treated, viz. Hordeum pubiflorum ssp. pubiflorum (2n=14), ssp. halophilum (comb. nov.) (2n=14), H. comosum (2n=14), H. jubatum ssp. jubatum (2n = 28), H. jubatum ssp. intermedium (2n = 28), H. lechleri (2n=42), H. procerum (2n=42), and H. arizonicum (2n=42). Information about morphological variation, cytology, and genome relationship, ecology and distribution is presented.     

Genome relationships in the genus Dasypyrum (Gramineae)

To elucidate the genome relationships in the genus Dasypyrum and the ancestry of tetraploid D. breviaristatum, two cytotypes of D. breviaristatum and D. villosum were reciprocally crossed with one another. Chromosome pairing at the first metaphase of meiosis and fertility were examined in the F1 hybrids and the parental plants. The mean pairing configuration and mean arm pairing frequency in D. villosum-D. breviaristatum (2x) hybrids were 11.12I + 1.44II per cell and 0.107, respectively, and they were almost completely sterile. In D. breviaristatum (4x)-D. breviaristatum (2x) hybrid, up to seven trivalents were formed, and the mean pairing configuration was 3.38I + 3.20II + 3.74III + 0.005IV per cell. The mean arm pairing frequency and relative affinity calculated in that F1 hybrid were 0.915 and 0.641, respectively. Seven bivalents and seven univalents were characteristically formed in D. villosum-D. breviaristatum (4x) hybrids. Based on the present results, we clearly concluded that the genome of diploid D. breviaristatum is distantly related to the genome V of D. villosum, and that these two species have different basic genomes. We, therefore, proposed the symbol Vb for the haploid genome of diploid cytotype of D. breviaristatum. Moreover, we concluded that tetraploid D. breviaristatum is an autotetraploid with doubled sets of the genomes homologous with that of diploid D. breviaristatum, and we proposed the genome constitution VbVb for the haploid genome set of tetraploid cytotype of D. breviaristatum. Furthermore, from the chromosome pairing in the F1 hybrids involving Moroccan and Greek accessions, it was suggested that complicated rearrangements of chromosome structure have occurred in tetraploid D. breviaristatum in its natural populations across the entire distribution area.     

Genotypic variation in tetraploid wheat affecting homoeologous pairing in hybrids with Aegilops peregrina.

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 (TIMO1). 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.     

Interspecific hybridization with Hordeum depressum (Poaceae)

Hordeum depressum is a distinct, reproductively isolated, tetraploid species. The study of meiosis in hybrids with the closest congeneric relatives revealed intermediate to low pairing with most species. The exceptions are Hordeum brachyantherum ssp. californicum and H. arizonicum. H. depressum , probably share one highly similar genome with ssp. californicum and perhaps two with H. arizonicum . However, it is difficult to draw firm conclusions based only on meiotic figures, since the pairing may be disturbed by high autosyndetic pairing between the H. depressurn genomes.     

INTERSPECIFIC HYBRIDIZATION IN HAPLOPAPPUS AND ITS BEARING ON CHROMOSOME EVOLUTION IN THE BLEPHARODON SECTION

Jackson , R. C. (U. Kansas, Lawrence.) Interspecific hybridization in Haplopappus and its bearing on chromosome evolution in the Blepharodon section. Amer. Jour. Bot. 49 (2) : 119–132. Illus. 1962.—Cytological analyses of interspecific hybrids between H. gracilis (n = 2) and H. spinulosus ssp. australis (n = 8) indicate that ssp. australis is a segmental allotetraploid, derived from past hybridization between 2 taxa with chromosome numbers of n = 4. Analysis of hybrids between H. gracilis (n = 2) and H. ravenii (n = 4), a previously undescribed species, has shown that the chromosome segments of these 2 species are almost completely homologous. Differential contraction is suggested as the explanation for the disappearance in late pachytene of presumed non-homologous segments which were evident in some cells at early pachytene. The pairing relationship of gracilis and ravenii chromosomes at pachytene and later prophase I stages of meiosis indicates that gracilis has evolved from ravenii by an aneuploid reduction process similar to that described for Crepis. The close morphological relationship of the 2 species adds further support to this proposition. Data from the cytological analysis of both interspecific hybrids indicate that x = 4 is the basic chromosome number for the Blepharodon section of Haplopappus.     

Intergeneric hybridization between Hordeum and Hordelymus (Poaceae)

Intergeneric hybridization between the monotypic Eurasian genus Hordelymus and species of Hordeum was performed. Hybrids were obtained in nine combinations and the meiotic pairing was analyzed in four combinations (6 families). Hordelymus europaeus is probably an alloploid with two different genomes. The very low pairing in the hybrids with H. jubatum, H. brachyantherum , and H. pusillum indicates that these species do not have any genomes in common with Hordelymus. The hybrids with H. depressum have an intermediate pairing which suggests that there is a certain homoeology in one genome.     

Studies on genome relationship and species-specific PCR marker for Dasypyrum breviaristatum in Triticeae

Dasypyrum breviaristatum and nine related species in Triticeae were analyzed using the random amplified polymorphic DNA (RAPD) technique, in order to understand the genetic relationship and to develop species specific markers. The genome relationship dendrogram shows that D. breviaristatum and D. villosum could not be grouped together, indicating that D. breviaristatum was unlikely to be directly derived from D. villosum, while D. breviaristatum was closest to Thinopyrum intermedium, which implied that they might have similar breeding behaviors when introducing their chromatins into wheat. A D. breviaristatum genome specific RAPD product of 1182bp, was cloned and designated as pDb12H. Sequence analysis revealed that pDb12H was strongly homologuos to a long terminal repeat (LTR) Sabrina retrotransposon newly reported in Hordeum. The pDb12H was converted into a PCR based marker, which allows effectively monitoring the D. breviaristatum chromatin introgression into wheat. Fluorescence in situ hybridization (FISH) suggested that pDb12H was specifically hybridized throughout all D. breviaristatum chromosomes arms except for the terminal and centromeric regions, which can be used to characterize wheat -D. breviaristatum chromosome translocation. The genomes repetitive element will also be useful to study gene interactions between the wheat and alien genomes after the polyploidization.     

Connection between rod bivalents and incomplete meiotic association at NORs in Hordeum marinum Huds

The hypothesis of a connection between rod bivalent formation and incomplete meiotic association at NORs of SAT-chromosomes of H. marinum is supported. PMCs of H. marinum ssp. gussoneanum (2x), two diploid ssp. marinum × ssp. gussoneanum (2x) hybrids and two ssp. gussoneanum (4x) × Secale cereale hybrids at metaphase I (M-I) were analyzed by in situ hybridization. The probe pTa71 labelled rDNA sites at NORs of a single pair of homologous or near-homologous SAT-chromosomes of H. marinum in each material. In the three diploids, M-I was regular with ring bivalents and one or a few rods (av.13.52 bound arms cell(-1) ). More rod bivalents than the expected one out of seven, i.e. 30, 67 and 89% included rDNA-carrying chromosomes. Corresponding bound short arm frequencies were 0.89, 0.72 and 0.52, while long arms and arms of other chromosomes presented complete or near- complete association. The two heterogenomic hybrids had a less regular M-I (av. 8.04 bound arms cell(-1) ) including 20% rDNA-carrying rods with bound arm frequencies of 0.29 in short and 0.87 in long arms. Positions of chromosome associations were established in all 150 rDNA-carrying bivalents. In 77 bivalents with short arm associations, 4% of these occurred proximally to, none at, and 96% distally to rDNA sites, i.e. in satellites. In 143 bivalents with long arm associations, 83% occurred at interstitial and 17% at terminal positions. The observations combine increased frequency of rDNA-carrying rods with decreased frequency of association at NOR regions of SAT-chromosomes. The basis for the relationship is discussed.     

Hystrix patula与Pseudoroegneria libanotica属间杂种的细胞学研究

为研究猬草Hystrixpatula的染色体组组成,进行了H.patula与Pseudoroegnerialibanotica的人工杂交,获得杂种F1,观察了亲本和杂种F1花粉母细胞减数分裂染色体配对行为。杂种F1染色体配对较高,84%的细胞形成7个或7个以上二价体,其构型为6.08Ⅰ+7.48Ⅱ,C-值为0.69。结果表明,H.patula含有St染色体组。     

Determination of the frequency of wheat-rye chromosome pairing in wheat x rye hybrids with and without chromosome 5B

Genomic in-situ hybridization (GISH) was used to determine the amount of wheat-rye chromosome pairing in wheat (Triticum aestivum) x rye (Secale cereale) hybrids having chromosome 5B present, absent, or replaced by an extra dose of chromosome 5D. The levels of overall chromosome pairing were similar to those reported earlier but the levels of wheat-rye pairing were higher than earlier determinations using C-banding. Significant differences in chromosome pairing were found between the three genotypes studied. Both of the chromosome-5B-deficient hybrid genotypes showed much higher pairing than the euploid wheat hybrid. However, the 5B-deficient hybrid carrying an extra chromosome 5D had significantly less wheat-rye pairing than the simple 5B-deficient genotype, indicating the presence of a suppressing factor on chromosome 5D. Non-homologous/non-homoeologous chromosome pairing was observed in all three hybrid genotypes. The value of GISH for assessing the level of wheat-alien chromosome pairing in wheat/alien hybrids and the effectiveness of wheat genotypes that affect homoeologous chromosome pairing is demonstrated.     

Genomic relationships between hexaploid Helianthus resinosus and diploid Helianthus annuus (Asteraceae)

Genus Helianthus comprises diploid and polyploid species. An autoallopolyploid origin has been proposed for hexaploid species but the genomic relationships remain unclear. Mitotic and meiotic studies in annual Helianthus annuus (2n = 2x = 34) and perennial Helianthus resinosus (2n = 6x = 102) as well as the F₁ hybrids between both species were carried out. Chromosome counting confirmed the hybrid origin of the latter plants and their tetraploid condition. Bivalents in hybrids ranged from 12 to 28 ( $ \bar{x} $  = 20.8). Univalents, trivalents and quadrivalents were also observed. Meiotic products comprised dyads, triads and normal tetrads and pollen grains were heterogeneous in size. These observations suggest the occurrence of 2n pollen in addition to the expected n. Genomic in situ hybridization (GISH) of total H. annuus DNA on H. resinosus chromosomes rendered weak but uniform signals; similar hybridization pattern was observed using three other annual species. Hybridization with H. annuus probe performed on root tip cells of F₁ H. annuus × H. resinosus hybrids revealed 17 chromosomes with a strong hybridization signal. GISH in hybrid meiocytes distinguished chromosomes from parental species and revealed autosyndetic pairing of H. resinosus chromosomes, allosyndetic pairing in bivalents, trivalents and quadrivalents, and the presence of univalents derived from parents, H. annuus and H. resinosus. Results obtained from classical and molecular cytogenetics do not support H. annuus as a direct ancestor of H. resinosus. The occurrence of allosyndetic pairing and the relatively high fertility of the F₁ hybrids point to the possibility that useful genes could be transferred from H. resinosus to cultivate sunflower, although the effective rate of recombination has not been evaluated. GISH method proved effective to recognize parental chromosomes in H. annuus × H. resinosus progeny.     

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.     

Molecular cytogenetic characterization and disease resistance observation of wheat-Dasypyrum breviaristatum partial amphiploid and its derivatives

A wheat-Dasypyrum breviaristatum partial amphiploid and its derivatives were analyzed by molecular cytological observation and tested for disease resistance in order to evaluate the potential use of the D. breviaristatum for wheat improvement. A fertility-improved partial amphiploid, TDH-2, was produced from the selfing population of Triticum aestivum cv. Chinese spring (CS)-D. breviaristatum amphiploid. Based on the results obtained from genomic in situ hybridization (GISH) and seed protein electrophoresis, we found the presence of fourteen D. breviaristatum chromosomes and the absence of D genome in TDH-2, indicating that the genomic composition of TDH-2 was AABBV(b)V(b). GISH analysis on BC(1)F(4) progenies of TDH-2xwheat demonstrated that alien D. breviaristatum chromosomes or segments were frequently transmitted. A survey of diseases resistance revealed that powdery mildew resistance from D. breviaristatum was totally expressed, however, the expression of stripe rust resistance from D. breviaristatum was dependent on the wheat background. The comparison of polymerase chain reaction (PCR), which was carried out using molecular marker SCAR(1400) linked to Pm21 D. villosum-derived powdery mildew resistance gene, suggested that D. breviaristatum possessed new resistance gene(s) different from that in D. villosum. The present study showed that the partial amphiploid TDH-2 and its derivatives could serve as novel sources for transfer of disease resistance genes to wheat.     

Cytoplasmic and genomic effects on meiotic pairing in brassica hybrids and allotetraploids from pair crosses of three cultivated diploids

Interspecific hybridization and allopolyploidization contribute to the origin of many important crops. Synthetic Brassica is a widely used model for the study of genetic recombination and "fixed heterosis" in allopolyploids. To investigate the effects of the cytoplasm and genome combinations on meiotic recombination, we produced digenomic diploid and triploid hybrids and trigenomic triploid hybrids from the reciprocal crosses of three Brassica diploids (B. rapa, AA; B. nigra, BB; B. oleracea, CC). The chromosomes in the resultant hybrids were doubled to obtain three allotetraploids (B. juncea, AA.BB; B. napus, AA.CC; B. carinata, BB.CC). Intra- and intergenomic chromosome pairings in these hybrids were quantified using genomic in situ hybridization and BAC-FISH. The level of intra- and intergenomic pairings varied significantly, depending on the genome combinations and the cytoplasmic background and/or their interaction. The extent of intragenomic pairing was less than that of intergenomic pairing within each genome. The extent of pairing variations within the B genome was less than that within the A and C genomes, each of which had a similar extent of pairing. Synthetic allotetraploids exhibited nondiploidized meiotic behavior, and their chromosomal instabilities were correlated with the relationship of the genomes and cytoplasmic background. Our results highlight the specific roles of the cytoplasm and genome to the chromosomal behaviors of hybrids and allopolyploids.     

AUTOTETRAPLOIDY IN HAPLOPAPPUS SPINULOSUS HYBRIDS: EVIDENCE FROM NATURAL AND SYNTHETIC TETRAPLOIDS

Haplopappus spinulosus (Asteraceae) is a herbaceous, perennial weed common throughout the western Great Plains of North America and includes both diploid and tetraploid populations. A number of populations in southeastern Colorado were analyzed cytogenetically and morphologically for two reasons. First, initial observations during a routine population survey showed they were morphologically intermediate between the diploid subspecies glaberrimus and spinulosus, suggesting they might have arisen via hybridization. Second, cytological examination revealed that they were tetraploid. Because there was indication of hybrid origin, it was of interest to determine whether the populations were behaving as autopolyploids or segmental allopolyploids. The distinction between these two polyploid types is not easily made since both are likely to form multivalents at meiosis, but equations derived from a model proposed by Jackson and Hauber (1982) have made it possible to determine statistically whether a tetraploid individual is behaving meiotically as an autotetraploid. Meiotic configuration frequencies at diakinesis were determined for each tetraploid plant sampled, and observed frequencies were compared to those expected for an autotetraploid having the same maximum number of chiasmata per bivalent, chiasma frequency and chromosome number. In general, the meiotic behavior of the tetraploids was no different from that expected for autoploids. The initial hypothesis that the populations were derived from hybridization was tested by a detailed cytogenetic and morphological study of the presumed parental subspecies, F, hybrids, and natural putative hybrids. The evidence supports the hypothesis that the natural autotetraploids arose from the hybridization of ssp. glaberrimus and ssp. spinulosus.     

Morphological, yield, cytological and molecular characterization of a bread wheat X tritordeum F1 hybrid

The morphological, yield, cytological and molecular characteristics of bread wheat X tritordeum F₁ hybrids (2n =6x = 42; AABBDH^ch) and their parents were analysed. Morphologically, these hybrids resembled the wheat parent. They were slightly bigger than both parents, had more spikelets per spike, and tillered more profusely. The hybrids are self-fertile but a reduction of average values of yield parameters was observed. For the cytological approach we used a double-target fluorescencein situ hybridization performed with total genomic DNA fromHordeum chilense L. and the ribosomal sequence pTa71. This technique allowed us to confirm the hybrid nature and to analyse chromosome pairing in this material. Our results showed that the expected complete homologous pairing (14 bivalents plus 14 univalents) was only observed in 9.59% of the pollen mother cells (PMCs) analysed. Some PMCs presented autosyndetic pairing of Hch and A, B or D chromosomes. The average number of univalents was higher in the wheat genome (6.8) than in the Hch genome (5.4). The maximum number of univalents per PMC was 20. We only observed wheat multivalents (one per PMC) but the frequency of trivalents (0.08) was higher than that of quadrivalents (0.058). We amplified 50 RAPD bands polymorphic between the F₁ hybrid and one of its parents, and 31 ISSR polymorphic bands. Both sets of markers proved to be reliable for DNA fingerprinting. The complementary use of morphological and yield analysis, molecular cytogenetic techniques and molecular markers allowed a more accurate evaluation and characterization of the hybrids analysed here.     

光对棉铃虫和烟青虫杂交的影响

【目的】力求建立一种准确鉴定室内棉铃虫Helicoverpa armigera(Hübner)、烟青虫H.assulta(Guenée)及其杂交种的分子技术,探讨棉铃虫和烟青虫杂交的可能。【方法】室内暗期设置0.5 lx黑光灯和白炽灯,测定不同配对模式下3日龄处女棉铃虫和烟青虫的杂交率;筛选可区分室内建立的棉铃虫、烟青虫及杂交种家族的微卫星位点;于架设有黑光灯的温室内释放棉铃虫和烟青虫混合种群,利用微卫星分子标记技术检测子代微卫星位点大小。【结果】两种蛾类在任何一种配对模式下均可杂交,混合种群处理杂交率为2.92%,且均为烟青虫雌蛾与棉铃虫雄蛾配对交配;黑光灯、白炽灯和黑暗条件下杂交率无显著性差异;筛选出的Har SSR1、Har SSR9和Har SSR10在两种蛾类上的等位基因片段大小不一样;两年温室实验共鉴定360头子代,混合种群中未检测到杂交后代。【结论】交配笼中棉铃虫和烟青虫能进行种间杂交,弱光不能提高二者杂交率;混合种群处理只发现一种杂交配对模式,说明了棉铃虫雄蛾的交配竞争能力更强;成功利用微卫星分子标记技术鉴定室内建立的棉铃虫、烟青虫及杂种家族,提供了一种简单准确的分子鉴定手段;两年温室实验未检测到杂交种,说明2种蛾类之间存在着交配前生殖隔离机制。     

Genome differentiation in Magonoliaceae as revealed from meiotic pairing in interspecific and intergeneric hybrids

The cross compatibility within and between Yulania Spach and Michelia L.(Magnoliaceae) is relatively good and various such hybrids,obtained by conventional artificial hybridization,are available.The aim of the present study was to determine the extent of genome differentiation between the species involved in these crosses through the observation of chromosome pairing during meiosis in pollen mother cells (PMCs) of the hybrids.Chromosome pairing behavior was studied in five species (2n =38) and two interspecific hybrids of Michelia,eight species (2n =38,76 and 114) and 10 interspecific hybrids of Yulania,and three intergeneric hybrids between Michelia and Yulania.The results showed that chromosome pairing was normal with bivalent formation in diploid parental species and in interspecific hybrids.In addition to bivalents,multivalents were encountered in polyploid parental species and polyploid interspecific hybrids.In the intergeneric hybrids between a tetraploid Yulania and two diploid Michelia,19 chromosomes,most likely originating from Michelia,were unable to synapse from zygotene to metaphase I.Meiotic chromosome pairing indicated a high degree of homology between species within Michelia and Yulania and less homology between the genomes of these two genera.The differentiation of morphological characters and the distinctness of natural distribution also support the conclusion that these two genera are likely independent monophyletic groups.This suggests that the two genera were split at early evolution of Magnoliaceae and the overlapping characteristics in external morphology and internal structures of the two genera may be the result of parallel evolution or ancient common ancestry.     

Intergeneric hybridization and C-banding patterns inHordelymus (Triticeae,Poaceae)

Crosses ofHordelymus europaeus (2n = 4x = 28) with four genera in theTriticeae were attempted. Adult hybrids were obtained in combinations withHordeum bogdanii (2x),Hordeum depressum (4x), andSecale cereale (2x). The meiotic pairing was very low in the hybrids withH. bogdanii andSecale cereale (0.12 and 0.30 chiasmata/cell, respectively), whereas high pairing (9.90 chiasmata/cell) was found in hybrids withH. depressum due to autosyndetic pairing ofH. depressum chromosomes. The chromosome complement ofHordelymus europaeus comprised 16 metacentrics, 8 submetacentrics, and 4 SAT-chromosomes. The Giemsa C-banding patterns of the chromosomes were characterized by small to minute bands at no preferential positions. It is hypothesized thatHordelymus europaeus may genomically be closest related toTaeniatherum andPsathyrostachys spp.     

Genome relationships between Hordeum procerum (6x) and H. lechleri (6x)

Investigations on the meiotic behaviour of chromosomes in interspecific hybrids (2n=6x=42) between Hordeum lechleri (6x) and H. procerum (6x) and in their component haploids have been utilized to assess the nature of pairing and the extent of genome homology between the two species. In the F₁ hybrids an average of 25 (60%) chromosomes associated at metaphase I, mostly as bivalents. A majority (60%) of the pollen mother cells (PMCs) in H. procerum haploids (2n=3x=21) displayed 21 univalents and even in the remainder, a maximum of two rod bivalents were formed resulting in an average of 0.52 bivalents per cell. In haploids of H. lechleri (2n=3x=21) however, 30% of chromosomes pair. The sum of the chromosomal associations in the component haploids represents only 17% of the complement, far below the observed frequency (60%) in the hybrids. Thus, the pairing displayed in hybrids between H. lechleri and H. procerum was mostly allosyndetic and suggestive of two genomes being common in these species.In haploid H. procerum 1/3 of the PMCs displayed a tripolar organisation of chromosomes leading to triad and hexad formation after divisions I and II respectively. The significance of hexad formation in the trihaploid H. procerum and a possible suppression of homoeologous pairing in H. procerum haploids are discussed.