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.     

Rapid estimation of ploidy levels in in vitro-regenerated interspecific Arachis hybrids and fertile triploids

Summary A significant correlation among chromosome number, chloroplast number and pollen grain size was observed using interspecific hybrids (2n=30,60), Arachis hypogaea (2n=40), A. stenosperma (PI 338280) (2n=20), A. batizocoi (PI 468329) (2n=20) and A. villosulicarpa Hoehne (PI 336984) (2n=20), representing four ploidy groups. Both chloroplast number in guard cells and pollen grain size were found to be positively correlated with ploidy, and provided a reliable method to distinguish diploid, triploid, tetraploid and hexaploid plants from each other regardless of their taxonomic backgrounds. These methods in combination with root-tip chromosome counts enabled us to confirm ploidy determinations on all three dissue layers, L₁, L₂ and L₃ (guard cells, microsporocytes and root meristematic cells, respectively), and to detect the chimeric nature of some colchicine-treated tissue culture-derived plants. Screening pollen grains by size and the detection of highly stainable and viable, 30-chromosome pollen grains have enhanced the use of triploid hybrids in peanut breeding programs.     

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.     

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.     

A synthetic hexaploid (2n = 42) oat from the cross of Avena strigosa (2n = 14) and domesticated A. magna (2n = 28)

A synthetic hexaploid oat was produced by chromosome doubling of a sterile triploid hybrid between cultivated Avena strigosa (2n = 14) cv. Saia and a domesticated form of A. magna (2n = 28). The synthetic hexaploid was intermediate between its parents in panicle shape and lemma color, similar to the tetraploid parent in spikelet structure, and to the diploid parent in having a single, albeit partially shriveled seed per spikelet, and low protein content. By the third generation, plants with yellowish lemmas, mostly two seeds per spikelet and better filled grains had been selected. Rust resistance of the diploid parent was retained in the synthetic hexaploid, but not tolerance to barley yellow dwarf virus disease (BYDV). Chromosome associations at meiosis in the triploid hybrid was low, with over 60% of them being univalents. Bivalent association was the rule in the synthetic hexaploid with an occasional one or two quadrivalents. Regular meiosis with 21 bivalents was observed in further generations. The preferential pairing of homologous chromosomes in the synthetic hexaploid was probably contributed by the A. strigosa genome which exhibits this tendency in artificial allopolyploid situations. Selection of yellow lemma color and two seeds per spikelet suggests that the genes controlling these traits are located on the chromosomes involved in quadrivalents in the synthetic hexaploid. The potential and limitations of utilizing the synthetic hexaploid in oat research and breeding are briefly discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Pollen grain size in four ploidy levels of genusAvena

Summary Pollen grain size in many genera positively correlated with chromosome number. In this study, oat (Avena) pollen grain size was examined in diploids (2×=14, one species, for accessions), tetraploids (4×=28, five species, 20 accessions), hexaploids (6×=42, one species, eight cultivars), and in 10 octoploid (8×=56) accessions. Mature anthers ready to dehisce pollen were sampled from one to six plants per accession, and pollen grains were squeezed out of the anther with tweezers. Oat pollen grains are slightly elliptical, and the length of the major axis was found to be highly correlated with the minor axis width (r=0.94^**). Pollen grain length, 39.3 m for diploids, 41.3 m for tetraploids, 47.0 m for hexaploids, and 48.8 m for octoploids, was positively correlated (r=0.86^**) with ploidy level. No genomic or species effects appeared to influence this trait. Only tetraploidAvena vaviloviana accession PI 412767 produced two distinct class sizes of pollen grains, 99% normal (43.0 m) and 1% large (52.7 m).     

Production of 2n pollen of Asiatic hybrid lilies by nitrous oxide treatment

Tetraploid varieties of lilies have superior agronomic traits such as large flowers and resistance to physiological disorders. In the present study, we attempted to induce 2n pollen of Asiatic hybrid lilies by arresting the meiotic process with nitrous oxide (N₂O) gas. To determine which meiotic stage is optimal for induction of 2n pollen, plants with attached buds at different meiotic stages were treated with N₂O for 24 h in a pressure-tolerant cylinder. A few 2n pollen grains were induced using plants with anthers in prophase I, whereas mixed pollen grains of differing size were produced using plants undergoing meiotic metaphase predominantly in anthers. Although normal lily pollen grains are elliptical, nitrous oxide exposure induced giant pollen grains that appeared spherical. Flow cytometry analysis showed that the giant pollen grains were diploid. When mixed pollen that included normal and giant pollen was crossed to tetraploid cultivars, the resulting seedlings were tetraploid and aneuploid, indicating that the giant pollen grains were diploids that could generate tetraploid seedlings through fusion to diploid eggs supplied from a tetraploid female parent. Thus, treatment with N₂O is useful for the production of 2n lily pollen and may provide a new approach for tetraploid lily breeding.     

Formation of 2n gametes in durum wheat haploids: Sexual polyploidization

Allopolyploidy, resulting from interspecific and intergeneric hybridization accompanied by sexual doubling of chromosomes, has played a major role in the evolution of crop plants that sustain humankind. The allopolyploid species, including durum wheat, bread wheat, and oat, have developed a genetic control of chromosome pairing that confers on them meiotic regularity (diploid-like chromosome pairing), and hence reproductive stability, and disomic inheritance. Being natural hybrids, they enjoy the benefits of hybridity as well as polyploidy that make them highly adaptable to diverse environments. Despite the complexities of sexual reproduction, it is widespread among plants and animals. Sexual polyploids are highly successful in nature. Sexual polyploidization is far more efficient than somatic chromosome doubling. Sexual polyploidization effected by functioning of unreduced (2n) gametes in the parental species or in their hybrids has been instrumental in producing our grain, fiber, and oilseed crops. Evidence is presented for the occurrence of sexual polyploidization in durum haploids. ThePh1-induced failure of homoeologous pairing is an important factor in the formation of first division restitution(FDR) nuclei and 2n gametes. The evolutionary and breeding significance of sexual polyploidization is discussed. It is emphasized that three factors, viz.,sexual reproduction, allopolyploidy, and genetic control of chromosome pairing,jointly constitute a perfect recipe for cataclysmic evolution in nature. 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.     

Genotypic control of chromosome pairing in amphiploids involving the cultivated oat Avena sativa L.

Summary A genotype of the diploid species Avena longiglumis (Cw 57) has been shown to modify the genetic control of diploid-like chromosome pairing in the cultivated oat, A. sativa (2n=6x=42) leading to increased homoeologous chromosome pairing in 4x hybrids between the two species (Rajhathy & Thomas, 1974). The Cw 57 genotype has a similar effect in increasing homoeologous chromosome pairing in amphiploids combining diploid and hexaploid genomes including associations between alien chromosomes and their corresponding pairs in hexaploid species. The effect of the Cw 57 genotype is probably in altering the specificity of chromosome pairing in the early stages of meiosis. The use of the Cw 57 genotype to induce homoeologous chromosome pairing as a technique for the transfer of desirable alien variation into the cultivated oat is discussed.     

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.     

The production of 2n pollen in rose

Based on the size differences found between haploid and diploid pollen produced by diploid and tetraploid rose cultivars, respectively, 2n pollen producers were identified in a population of 53 diploid hybrids from a cross between a dihaploid rose, derived from the haploidization of a tetraploid modern cultivar and the diploid species R. wichuraiana. Frequency of 2n pollen producers was estimated in 2002, 2003 and 2004. Highly variable frequencies were found i) within population; ii) during years of observation (between years and between different months in the same year). The variation of 2n pollen production could be related to environmental fluctuations. A cytological analysis of male meiosis was carried out in 10 hybrids randomly chosen. Among meiotic abnormalities leading to 2n pollen formation, triads (containing a 2n microspore at one pole and two n microspores at the other) resulting from abnormal spindle geometry were frequently observed. The mode of 2n pollen formation is genetically equivalent to a First Division Restitution (FDR) mechanism. FDR 2n pollen transmits a high percentage of the heterozygosity from the diploid parent – 2n pollen producer-to the tetraploid offspring.     

A tetraploid hybrid plant from 4x × 2x crosses in Vitis and its origin

To study the origin of unreduced (2n) gametes in diploid Vitis cultivars, we surveyed the occurrence of tetraploid hybrid seedlings from 40 interploid crosses with five tetraploid and seven diploid cultivars. A total of 250 seedlings from the interploid crosses were established through embryo culture and by seed sowing. In 20 2x × 4x crosses, no tetraploid hybrid seedlings were derived from 8,902pollinations. In 20 4x × 2x crosses, two tetraploid hybrid seedlings were obtained from 8,057 pollinations. Investigation of isozyme genotypes of the two tetraploid seedlings using three variable enzyme systems indicated that one of the two seedlings resulted from the union of a diploid egg with a 2n male gamete and that failure of second meiotic division resulted in the formation of the 2n male gamete. The percentage of giant pollen grains in `Muscat Bailey A', a pollen parent of the tetraploid seedling, was relatively high(about 5.9%) and 10.9% of the giant pollen grains germinated on agar medium. These results suggested that 2n pollen of diploid cultivars are useful for breeding tetraploid grape. This revised version was published online in August 2006 with corrections to the Cover Date.     

Introgressive hexaploid oats from the Avena abyssinica (♀) × A. sativa hybrid: performance, grain lipids and proteins

A fertile hexaploid oat plant was obtained after several generations of selection for seed set and plant type from a colchicine-produced decaploid hybrid, Avena abyssinica(2n = 4x = 28, AABB) × A. sativa (2n = 6x = 42, AACCDD). The selected line proved to be stably fertile and in many characteristics equal or superior to the hexaploid parent. The grain protein fractions showed two qualitative differences from those of the pollen parent and several differences from the maternal parent. The fractionating extraction used was new for oats. The fatty acid composition of grains of the hybrid derivative was similar to that of the pollen parent, but different from that of the maternal parent. The maternal parent (A. abyssinica) had a relatively high 16:0 fatty acid content (ca. 20.5 mol%) compared with the level of the hexaploid parent and the hybrid derivation (ca. 17.5 mol% each) in field-grown grain. However, in grain produced in the greenhouse, the hexaploids had ca. 20.5 mol% of 16:0 fatty acid and a decrease in 18:1 fatty acid, whereas seed of the A. abyssinica parent showed only a slight increase (ca.21.5 mol%) in 16:0 fatty acid. These and other responses statistically significant may be due to adaptation to temperature conditions being wider in the hexaploids than in the East-African A. abyssinica. A new method of grain lipid extraction was introduced and showed good reproducibility. The derived hexaploid oat can be crossed with A. sativa for breeding purposes and due to its early maturity might also have direct use in northern or high-altitude cultivation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Introgression of crown rust resistance from diploid oat Avena strigosa into hexaploid cultivated oat A. sativa by two methods: direct crosses and through an initial 2x·4x synthetic hexaploid

New sources of resistance to crown rust, Puccinia coronata f. sp. avenae (Eriks.), the major fungal disease of cultivated oat, Avena sativa L. (2n = 6x = 42), are constantly needed due to frequent, rapid shifts in the virulence pattern of the pathogen. Crown rust resistance identified in the diploid oat A. strigosa (Schreb.) (2n = 2x = 14) accession CI6954SP was transferred into cultivated oat using two methods: direct cross of the diploid to a hexaploid cultivar facilitated by embryo rescue, and initial cross of the diploid to a wild tetraploid oat to make a synthetic hexaploid for subsequent crossing to a hexaploid cultivar. Two tetraploids, a crown rust resistant A. murphyi (Ladiz.) accession P12 and a susceptible A. insularis (Ladiz.) accession INS-1, were used in the 2x·4x crosses. Resistant backcross-derived lines were recovered by both methods. Although the 2x·4x synthetic method did not require the laborious discovery and rescue of an infrequent initial hybrid embryo of the direct cross, the direct cross method provided more rapid backcross recovery of plants with high fertility, full transmission of resistance, and desired plant and seed phenotypes. A suppressor effect, present initially but segregating in backcrosses, appeared to come from the CI6954SP donor and is the same as, or analogous to, suppression by an oat line with the crown rust resistance gene Pc38. No resistance from A. murphyi P12 was detected in advanced generations when it was introduced either as a component of a synthetic hexaploid or in direct crosses to A. sativa, indicating suppression of its resistance in interploidy combinations. That the dominant resistance gene transferred from CI6954SP and a gene Pc94 introgressed earlier from a different A. strigosa accession may be the same or quite similar to one another is indicated by their in-common specificity to suppression of resistance expression, susceptibility to a newly recovered rust isolate, and close linkage to the molecular marker SCAR94-2. The introgressed resistance genes from the different sources, even if the same, may have different cultivar genomic introgression sites, which would allow tests of dosage effects on resistance expression.     

Transfer of high kernel weight and high protein from wild tetraploid wheat (Triticum turgidum dicoccoides) to bread wheat (T. aestivum) using homologous and homoeologous recombination

Summary Wheat pentaploids were produced by hybridizing a high kernel weight (1000 grain wt=56 g), high protein (25.4%) line of wild tetraploid wheat (Triticum turgidum dicoccoides) as male parent, with the three hexaploids (T. aestivum) — normal Chinese Spring and its two homoeologous pairing mutants, ph _1b and ph ₂. The pentaploids were crossed as female parents to the two commercial hexaploid cultivars Warigal and Barkaee and 42-chromosome stable plants selected from the F₁ of the pentaploid x hexaploid crosses.Mean protein content of certain F₃ lines from all six pentaploid x hexaploid crosses was significantly higher than Chinese Spring and the respective commercial hexaploid parent (p<0.005) indicating high protein had been transferred from the tetraploid to the hexaploid level.Kernel weight amongst certain F₃ lines of the three pentaploids x Barkaee was significantly (p<0.0005) higher than either Chinese Spring or Barkaee, indicating the transfer also of high kernel weight from the tetraploid to the hexaploid level. However kernel weight was not significantly increased over Warigal in any F₃ lines of its crosses with the three pentaploids.High levels of homoeologous chromosome pairing in the ph-mutant pentaploids, plus evidence for significant modification of the composition of high-molecular weight (HMW) glutenin subunits of grain protein in certain F₃ derivatives of the ph-mutant pentaploid x hexaploid, crosses indicates that the ph-mutant-derived lines may possess novel (intergenome) genetic recombination, at least for high protein, and possibly kernel weight.     

Bilateral polyploidization inDactylis glomerata L. subsp.lusitanica: occurrence,morphological and genetic characteristics of first polyploids

Summary InDactylis glomerata L. subsp.lusitanica, triploid and tetraploid plants were obtained by bilateral sexual polyploidization in crosses between diploid parents known to produce 2n gametes. The polyploid and diploid progeny were compared for allozyme diversity (allele number and heterozygosity), phenological (pollen fertility, inflorescence emergence date), cellular (stomatic cell size) and morphological characters (vegetative biomass, seed weight, total seed number per plant, panicle number and seed number per panicle length) in four contrasting growing conditions. Stomatal cell size and the total number of alleles were significantly lower in diploids than in polyploids which sometimes recovered more than two distinct alleles at a locus via 2n gametes possessing heterozygous loci. Pollen fertility was similar in diploids and tetraploids, but significantly lower in the triploids. Seed weight was significantly higher in polyploids, although the average number of seeds per panicle length and the total number of seeds was higher in the diploid sibs. No phenological differences occurred among ploidy levels whereas such differences were observed inDactylis in all the natural areas of sympatry between diploids and their related polyploids. Our results provide evidence for an instantaneous effect of polyploidization on certain traits and suggest that natural selection would act subsequently on several other traits to allow the establishment of new polyploids.     

The production and behaviour of hexaploid hybrids between Hordeum vulgare and H. Bulbosum

Summary To produce hexaploid (or other polyploid) hybrids, diploid or tetraploid Hordeum vulgare was crossed with hexaploid or octoploid H. bulbosum, and perennial triploid hybrids between the two species were treated with colchicine. The crosses did not yield viable plants: seedset was low, the seed aborted and embryo culture was unsuccessful. The colchicine treatments geve rise to plants in which hexaploid chromosome numbers were observed. At the hexaploid level chromosomal instability occurred, resulting in chromosome elimination.The colchicine-treated triploid hybrids showed in the first years after the treatment better fertility after open flowering than untreated plants, but the level of fertility remained very low. The offspring consisted of haploid, diploid and approximately triploid plants like H. vulgare, tetraploid and approximately tetraploid plants like H. bulbosum, and plants with hybrid morphology and unstable chromosome number, which were highly sterile. Thus the crossing barrier between H. vulgare and H. bulbosum could not be broken down at higher ploidy level.     

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.