Genetic diversity within cultivated teas based on nucleotide sequence comparison of ribosomal RNA maturase in chloroplast DNA

The wild tuber-bearing Solanumspecies represent a genetic pool of enormous diversity, embracing high variability for many agronomic traits. S. kurtzianum (ktz), S. chacoense(chc) and S. ruiz-lealii (rzl) are wild diploid self-incompatible relatives of the common potato, Solanum tuberosumssp. tuberosum, with resistance to adverse biotic and abiotic factors. Pollen-pistil/style compatibility relations and seed set in intra- and interspecific crosses among ktz, chc and rzl were investigated. Pre- and post-zygotic hybridization barriers were detected in the three species, with variable incidence because the percentage of genotypic combinations that produced seeds was15–30% and 7–28% in intra and interspecific crosses, respectively. In pollen-pistil/style incompatible combinations, either pollen grains did not germinate or pollen tube growth was arrested at different sites of the style, but mainly in its upper third. Unilateral incompatibility was detected in both intra-and interspecific crosses, as well as bilateral incompatibility in certain combinations. The incompatibility relations observed could be explained by a genetic model with dominant genes in pistils that interact with corresponding dominant genes in pollen, that are independent of the S-locus. The complex arrangement of compatibility relationships found could not be directly related to taxonomic hypothesis or geographic distribution. These results suggest that divergence between the three species has occurred recently, and that internal barriers of reproductive isolation have not been an important factor in their evolution. This revised version was published online in July 2006 with corrections to the Cover Date.     

The importance of Endosperm Balance Number in potato breeding and the evolution of tuber-bearing Solanum species

Summary Endosperm failure is considered the primary reason for the lack of success in intra-and interspecific crosses. The Endosperm Balance Number (EBN) hypothesis is a unifying concept for predicting endosperm function in intraspecific, interploidy, and interspecific crosses. In the EBN system, every species has an effective ploidy (EBN), which must be in a 2:1 maternal to paternal ratio in the endosperm for crosses to succeed. The knowledge of EBN is very useful in the transfer of genes from exotic germplasm, and in the development of new breeding schemes in potato. The paper describes the strategies for introducing 2x(1EBN), 2x(2EBN), 4x(2EBN) and 6x(4EBN) germplasm into the cultivated 4x(4EBN) potato gene pool. A new methodology for producing 4x(4EBN) and 2x(2EBN) chromosome addition lines is also discussed. EBN has evolutionary importance in the origin of tuber-bearing Solanums. The role of the EBN in the origin of diploid and polyploid potato species, and as a barrier for hybridization and speciation of sympatric species within the same ploidy level is demonstrated. The origin of 3x and 5x cultivated tuber-bearing Solanums may also be explained using the EBN concept. EBN has been reported to exist in other plant species: alfalfa, beans, blueberries, rice, soybeans, squashes, tomato, forage legumes, grasses, ornamentals and Datura stramonium. This indicates that EBN may have broad application and could be useful for germplasm transfer and breeding in other crop species.     

黑籽南瓜种间杂交研究

通过调整播期克服了南瓜属种间花期不遇的问题,以美洲南瓜、中国南瓜、印度南瓜3个种共12个材料与黑籽南瓜进行了种间杂交试验。黑籽南瓜为母本不结实,以其他3个种为母本则均可结实。美洲南瓜×黑籽南瓜及印度南瓜×黑籽南瓜杂交获得的果实内均无种子。获得了18株中国南瓜×黑籽南瓜F1植株。种间杂种F1植株雄蕊退化,以其作为母本以中国南瓜或黑籽南瓜为父本进行回交,可结实但无种子。中国南瓜×黑籽南瓜F1形态为双亲中间型偏父本,其抗病性、抗虫性不及父本黑籽南瓜而倾向于母本中国南瓜。     

Effects of <Emphasis Type="Italic">Solanum demissum</Emphasis> chromosomes on crossability in the backcross progeny to <Emphasis Type="Italic">Solanum tuberosum</Emphasis>

A balance of maternal and paternal genetic factors, conceptually named the endosperm balance number (EBN), is required for normal endosperm development in interspecific crosses in potato. We previously found that Solanum demissum (D), a hexaploid wild species widely used in potato breeding, has a slightly lower EBN than S. tuberosum (T). To explore the genetic nature of the EBN, the berry-setting rate, seed number/berry, and seed weight were evaluated in BC₁ [(D?×?T)?×?T] plants, each possessing different portions of the S. demissum chromosomes, by reciprocal crosses with D and T, and a quantitative trait locus (QTL) analysis was performed. At least 99 S. demissum-derived QTLs were detected, of which 29 were associated with differential responses to D and T. Three QTLs were possibly co-localized on chromosomes 7A and 10D1, while the remaining 23 QTLs were independently located. The QTLs in the three S. demissum homoeologous chromosomes exhibited three types of interaction: (1) positive, (2) negative, and (3) one positive and one negative effect on the same trait. We found that several major genes, one of which was localized in the S. demissum chromosome 9A, and many minor genes controlled the crossability of BC₁ plants. The QTLs responsible for the differential responses to D and T were different between the BC₁ plants used as male and female parents, indicating that different genes control the male and female EBNs. Consequently, we conclude that the EBN is represented by the sum of various genetic effects controlled by a large number of genes.     

Variation in pollen and ovule fertility in varieties of cassava,and the effect of interspecific crossing on fertility

Records of the set of seeds obtained from intraspecific cassava crosses, from interspecific crosses between other Manihot species and cassava, and from backcrosses to cassava of three generations of interspecific hybrids are used to assess the variation in pollen and ovule fertility. It is concluded that the capacity of cassava varieties to set seeds has been reduced since they have evolved from non-tuberous wild forms and have been propagated vegetatively. Fertility was found to be very variable, and the capacity of the pollen of a variety to promote seed-set was not related to the fertility of the variety's female flowers. This situation would be expected if reduction in the fertilities of the two sexes were caused either by loss of balance in the genes controlling the functioning of the male gametophyte, or by unbalance of those concerned with the functioning of the female gametophyte or the tissues which nurture the embryo. Additional causes of sterility, probably including meiotic irregularities, appeared to operate in the F₁ interspecific hybrids, but such factors were probably not important after the first backcross generation. Manihot melanobasis, a wild form which is normally propagated by seed and whose relationship to cassava should be regarded as subspecific, contributed factors which enhanced the fertility of its hybrids with cassava. This form could be used in cassava breeding as a donor of seed fertility, but use could also be made of some existing varieties which still possess a moderately high capacity to set seed. It is desirable to select these as one parent when making difficult crosses.This paper is based almost entirely on the records of the late R. F. W. Nichols, who preceded the writer as cassava breeder with the East African Agriculture & Forestry Research Organisation, at Amani, Tanganyika.     

Identification of barriers to interspecific crosses in the genus Trifolium

Summary A study of pre- and post-fertilisation barriers after interspecific crosses of diploid and tetraploid Trifolium pratense L. and wild species T. alpestre L., T. medium L. and T. sarosiense Hazsl. was aimed at finding of a promising cross combination for obtaining hybrids. The growth of pollen tubes was arrested in interspecific crosses mainly when T. pratense was at a diploid level. To investigate the post-fertilisation barriers in detail, the hybrid embryo viability was traced by two clearing treatments of immature seeds: (1) using chloral hydrate (which proved to be most appropriate); and (2) a mixture of benzyl benzoate and dibutly phthalate. In interspecific combinations T. pratense (4×) × either T. alpestre or T. sarosiense, enlargement of immature seeds occurred, but no hybrid embryo was traced. Of the wild species used as a male parent for crosses, T. medium was the only exception from the point of view of fertilisation. Globular, heart and the early torpedo stages of hybrid embryos were observed 7 days after pollination (DAP) but only when T. pratense was at a tetraploid level. When T. pratense (2×, 4×) was used as a male parent for interspecific crosses with T. alpestre, T. medium and T. sarosiense, strong defects in various stages of embryogenesis were observed, particularly wrinkled and narrowing embryo sacs caused by an expansion of endothelial cells. We conclude with the following finding: (1) to make crosses only in one direction with T. pratense as a female parent and T. medium as a male; (2) to use tetraploid plants of T. pratense; (3) and to excise hybrid embryos at an early torpedo stage, about 7 DAP.     

Introgression of genes into cultivated Brassica napus through resynthesis of B. napus via ovule culture and the accompanying change in fatty acid composition

Fifteen lines of Brassica napus were resynthesized via ovule culture through 24 interspecific crosses between four Brassica oleracea and three Brassica campestris accessions. The degree of success in the interspecific crosses was significantly influenced by maternal genotypes. The interspecific hybrid production rate (HPR) varied with combinations from 0 to 76.9%, with a mean HPR of 24.7% for the crosses with B. campestris as the female parent and 6.9% for the crosses with B. oleracea as female parent. Twenty‐four crosses between seven natural and six resynthesized B. napus gave, on average, 10.3 seeds per pod, and ranged from 1.2 to 22.0 seeds per pod, depending on genotypes of both parents. Resynthesized lines of B. napus showed high erucic acid content and variable content of linolenic acid, ranging from 3.4% to 9.9%. The fatty acid composition in hybrid seeds between natural and resynthesized B. napus was dominated by the embryo genotypes; an additive mode was shown for erucic acid and positive over‐dominance for linolenic acid content.     

Interactions of BA, GA3, NAA, and surfactant on interspecific hybridization of Lycopersicon esculentum × L. chilense

Time consuming and expensive tissue culture type techniques are currently used for achieving Lycopersicon esculentum × L. chilense interspecific hybrids. The objective of this study was to determine if the number of viable hybrid seed produced directly from this wide cross could be improved by optimizing female/male parent selection when treated with various hormones in the presence or absence of a surfactant (Triton X-100). Individual or combination treatments of BA(6-benzylaminopurine), GA₃ (gibberellic acid), and/or NAA(α-naphthaleneacetic acid) were applied (both with and without surfactant)to the ovaries of two L. esculentum female parents (Fla 7613 and89S) one day following pollination by nineL. chilense accessions. Hormone applications over 1-, 3-, and 5-dayperiods were made and all treatments were compared to a distilled water control. Across all experiments 21 viable hybrids from 1920 fruits (1.1%) were obtained from2128 pollinations. There were five consistent trends observed across three experiments. First, treatments utilizing NAA produced more hybrids than any other treatment hormone. Second, eight of the total 21 hybrids resulted from five consecutive days of hormone treatments (P-value ≤0.01), and six of those eight were realized when NAA was involved. Third, the use of Triton X-100 in combination with the hormone treatments significantly reduced the number of fruits/pollination (P-value ≤ 0.01), but did not significantly reduce the number of overall viable hybrids (P-value > 0.4). Fourth, the L. chilense male parent LA2759 significantly produced the most interspecific hybrids (P-value ≤ 0.04),seven hybrids from 295 fruits, compared to the male parent LA130 which did not produce any hybrids (from 293 fruits) -other accessions were intermediate. Fifth, nearly ten times more hybrids were produced from crosses with Fla 7613 than with 89S. These experiments provide a foundation for future studies to discern specific concentrations, timing and frequency of application of NAA and possibly, GA₃, to efficiently produce interspecific hybrids of L. esculentum × L. chilense. This revised version was published online in August 2006 with corrections to the Cover Date.     

Nicotiana tabacum L. as a source of cytoplasmic male sterility in interspecific cross with N. alata Link & Otto

A new source of cytoplasmic male sterility (CMS) in tobacco with interspecific origin is reported. In traditional selection wild tobacco species have been used as donors of cytoplasm. In the present study the cultivated species Nicotiana tabacum L. (n = 24) is a source of CMS. It was used as female parent and N. alata (n = 9) was involved as a pollinator. The F1 hybrid of this cross was completely sterile. Tissue culture method was applied to restore the female fertility. Regenerants obtained from the 5th passage were successfully pollinated with N. tabacum and seed-containing capsules were formed. All BC1P1 plants were male sterile. They possessed normally developed corollas, three-loculed or deformed pistils, and 1–2 stamens modified into secondary pistils. In some plants stamenless flowers were observed. Male sterility of BC1P1 was preserved in BC2P1–BC7P1 progenies confirming its cytoplasmic nature. This revised version was published online in July 2006 with corrections to the Cover Date.     

The transfer of the synaptic gene (sy-2) from 1EBN Solanum commersonii Dun. to 2EBN germplasm

Summary Crosses of a synaptic mutant (sy-2) of 2x(1EBN) S. commersonii Dun. with 2x(2EBN) S. chacoense Bitt. resulted in the production of 3x(2EBN) hybrids. Low levels of 2n egg production in these hybrids allowed subsequent crosses to be made with 4x(4EBN) cultivars, resulting in 5x(4EBN) S. commersonii-S. chacoense-Group Tuberosum hybrids. Seed set in these crosses averaged 0.22 seeds per fruit. Fifty-four unspotted seeds were produced in 1225 crosses of the 5x(4EBN) hybrids with 2x(2EBN) Group Phureja haploid extraction clones carrying dominant seed spot markers. These seeds yielded a total of 15 confirmed haploids having a diploid (2n=2x=24) or near diploid chromosome number. One haploid clone recovered displayed a synaptically abnormal phenotype similar to the original S. commersonii synaptic mutant. Preliminary test crosses of these haploids indicated that all haploids produced were 2EBN.This method for the transfer of alleles across EBN levels holds promise for the recovery of both qualitative and quantitative traits from 1EBN species. In these recovered, cross-compatible forms, clones with desired characteristics may be selected and easily incorporated into breeding programs.     

Incompatibility in diploid and tetraploid crosses of Cucumis sativus and Cucumis metuliferus

The African horned cucumber (Cucumis metuliferus Naud.; 2x = 2n = 24) contains genes that can confer resistance to many important cucumber (C. sativus L.; 2x = 2n = 14) pests [e.g., root-knotnematode, Meloidogyne incognita (Kofoid & White) Chitwood]. Cucumber is highly susceptible to this root-knot nematode species, and a recent screening of C. sativus accessions in the U.S. National Plant Germplasm collection did not identify sources of resistance. Thus,autotetraploids of Cucumis sativus and C. metuliferus were created to recover fertile resistant interspecific progeny. Autotetraploids were obtained at the highest rate when seeds were immersed in 0.5% colchicine for a period of 6 to 8 hrs. Treatment durations less than 6 hrs produced few tetraploids, and durations of 10 hrs or more were lethal to seeds or developing seedlings. Crosses between C. sativus and C. metuliferus were made using diploid and tetraploid lines in all possible combinations, including reciprocals. Fruit development occurred in crosses when diploid and tetraploid C. sativus were used as the female parent. However, seeds developed only in fruit of C. sativus (4n) ×C. metuliferus (2n) crossings. Seeds from these crosses, however,were flat and not viable. No fruit development occurred in crosses whereC. metuliferus was used as the female parent. This revised version was published online in August 2006 with corrections to the Cover Date.     

Parental effects on the performance of cultivated × wild species hybrids in potato

Valuable genetic diversity in diploid wild Solanum species can be accessed through crosses to haploids (2n = 2×) of the tetraploid cultivated potato, Solanum tuberosum. Haploid-wild species hybrids segregate for the ability to tuberize in the field. In addition, they vary in male fertility, vine size, stolon length, and tuber size. In this study, three haploids were crossed with nine diploid wild Solanum species and 27 hybrid families were evaluated in the field for two years. The proportion of male fertile hybrid clones varied depending on the wild species parent. A large effect of the female parent was detected for vine size, stolon length, tuber size, percent tuberization, and percent plants selected for agronomic quality. An exceptional haploid (US-W4) was identified for the production of agronomically desirable haploid-wild species hybrids. In hybrids derived from US-W4, differences among wild species parents were observed for agronomic quality. Superior hybrids were produced by S. berthaultii and S. microdontum. Reciprocal crosses were evaluated for a subset of families. When the wild species was used as the female parent, male fertility was restored, but tuberization and tuber size were reduced. Careful selection of both haploid and wild species parents can result in a large proportion of fertile, agronomically desirable hybrid offspring.     

Optimal pollination conditions for seed set after a self-pollination,an intraspecific cross and an interspecific cross of marrow-stem kale (Brassica oleracea var. acephala)

Summary The effects of different pollination techniques, with and without emasculation and delayed pollination, were investigated to find the conditions for maximum seed set after self-pollination and intraspecific and interspecific crosses of Brassica oleracea var. acephala. The results indicated that the pollination conditions achieving maximum seed set vary with the type of pollination. After controlled self-pollination, the best seed set occurs in bud 3 to bud 10. For the intraspecific cross, the youngest flower and the oldest bud produced the largest number of developed ovules but bud pollination was productive to bud 8. The yields from these two pollination types were best when the female parent was not emasculated. In the interspecific cross with B. campestris cv. Marco the best results came from the youngest flowers and the oldest buds subjected to the standard practice of pollinating directly after emasculation. Possible reasons for these effects are discussed.     

多毛番茄主要生物学特性研究

为了更好利用番茄野生资源,以多毛番茄为材料,对其生物学特性及与栽培番茄杂交坐果情况进行研究。结果表明：多毛番茄种子较小,无毛,具有较强的发芽能力;叶片较小,小叶排列紧密,叶绿素含量与普通番茄差别较小;茎较细、节间较长;花色鲜艳,柱头呈凹状,伸出花药,低温条件下花粉具有较强萌发能力;与栽培番茄杂交时,以栽培番茄做母本,具有坐果能力,反之不坐果。     

RFLP analysis of cytoplasmic male-sterile lines of pigeonpea [Cajanus cajan (L.) Millsp.] developed by interspecific crosses

Total DNA from three putative cytoplasmic male sterile (CMS) progenies derived from crosses between the wild species Cajanus sericeus and the cultivated species Cajanus cajan, five C. cajan, one accession of C. sericeus and two genetic male sterile lines of pigeonpea were compared for their RFLP patterns using maize mitochondrial DNA (mtDNA) specific probes. Three putative cytoplasmic male sterile (CMS) progenies from the multiple cross genome transfer of pigeonpea lines (CMS 7–1, CMS 12–3, and CMS 33–1) showed hybridization patterns identical to that of C. sericeus when DNA was digested with EcoRI and HindIII and probed with maize mtDNA clones. The results suggested that these putative CMS progenies have the mitochondria of the female wild species parent. The hybridization patterns of the three male parental lines used in the development of the CMS progenies were similar in all the restriction enzyme-probe combinations except HindIII-atp6. The genetic male sterile lines, MS Prabhat and QMS 1 differed from each other in their hybridization pattern. The genomic DNA hybridization pattern of HindIII digested DNA from ICPL 87 differed from the other pigeonpea lines when probed with the maize mtDNA clones. The cluster analysis of the hybridization data suggested the occurrence of variation in the mitochondrial genome even among the cultivated species. This revised version was published online in July 2006 with corrections to the Cover Date.     

胚珠培养获得秘鲁番茄与栽培番茄种间杂种具有非温敏抗根结线虫基因Mi-3

以含有Mi-3抗根结线虫基因的秘鲁番茄(Lycopersicon peruvianum)LA2823为父本,栽培番茄（Lycopersicon esculentum)高代优良自交系为母本,授粉后20～30d摘取幼果。将从幼果中分离出的直径大于2mm的胚珠放置在营养分化培养基上培养。2～4周后胚芽、胚根分别由两端伸出。培养的胚珠还可以通过体细胞胚发生途径产生胚芽。获得的种间杂种对根结线虫的抗性表型与野生种父本完全相同,在植物学性状上亦与其父本秘鲁番茄更相近。以栽培番茄为母本进行回交,成熟果实与母本相似,但其种子即使放置在MS培养基上也不能萌发。对回交种子胚败育现象进行了探讨。     

Breeding potential of Solanum tuberosum–S. commersonii pentaploid hybrids: fertility studies and tuber evaluation

Twenty-one (near) pentaploid hybrids between sexually incompatible Solanum commersonii [2x(1EBN)] and cultivated S. tuberosum [4x(4EBN)] were characterized for tuber traits and fertility. A number of genotypes resembled the S. tuberosum type in terms of stolon length and eye depth and produced tubers under long day conditions. Tuber yields were not as high as expected, probably due to lack of adaptation of the S. commersonii genome to the environmental conditions in southern Italy (on average 325 g·pt⁻¹ and 285 g·pt⁻¹ in 2003 and in 2004, respectively). Compared to 2004, in the summer of 2003, characterized by extremely high temperatures, hybrids gave a higher tuber yield than the S. tuberosum control, suggesting that in our environmental conditions the wild S. commersonii genome, rather than resulting in heterosis for tuber yield, provides better adaptation to harsh environments. Although aneuploidy has often been associated with reduced fertility, several hybrids were fertile in crosses with S. tuberosum when used as female parents. In particular, the average berry set and number of seeds per berry were 38.2% and 31.8%, respectively. Based on significant relationships between ploidy levels and all yield and fertility data measured, the presence of extra chromosome affected the parameters considered: tuber yield in 2003 and 2004, percentage of fruit set, number of seeds per berry and number of seeds per pollinated flower. Regression analysis also indicated that “Residuals” were significant for all parameters measured. Therefore, additional factors (e.g. the genetic makeup of hybrids) may be key to fertility and yield.     

Genotypic diversity enhances recovery of hybrids and fertile backcrosses of Phaseolus vulgaris L. × P. acutifolius A. Gray

Summary Homozygous and heterozygous Phaseolus vulgaris and P. acutifolius parental genotypes were hybridized to determine if genotypic diversity would aid gene transfer between these species. Certain P. vulgaris female parents resulting from diverse intraspecific crosses increased the frequency of species hybrids obtained. From a total of 19 self-sterile hybrids, 20 backcross-1 (BC₁) lines (P. vulgaris recurrent parent) were produced from 4 partially female-fertile hybrids, each of which had a heterozygous P. vulgaris female parent. Heterozygous P. acutifolius male parents did not influence the frequency at which interspecific hybrids could be produced but apparently improved female-fertility. Fertility of the F₁BC₁ generation was highly variable. The frequency of fertile individuals increased in each subsequent backcross generation (F₁BC₂ and F₁BC₃).Purdue University Agricultural Experiment Station Publication 9946.     

Triple fusion of the primary endosperm nucleus as a cause of interspecific cross-incompatibility in Avena

Summary Using five diploid, two tetraploid and three hexaploid Avena species (x=7) 78 reciprocal crosses were made. Of these 74 were successfull, though the frequency of seed set varied greatly. The development of hybrid seeds differed widely between cross combinations, especially in reciprocal crosses. Hybrid seeds were grouped into four types, (1) normal kernels, well developed and germinative (designated as D+); (2) shrivelled-empty kernels, which do not germinate (E–); (3) small viable kernels (Rd+) and (4) small inviable kernels (Rd–). Some rare intermediate types were found in certain crosses.The results are well interpreted in terms of a hypothesis of polar-nuclei activation in which the strength of the activating action of the male nucleus, and the reaction of the female nucleus at double fertilization are expressed as activating value(AV) and response value(RV), respectively. The degree of seed failure is closely related to the activation index(AI) of the polar nuclei, AV/2RV (or × 100). In a selfed plant the activation index is always 1/2=0.5 (or50%), and this usually results in the formation of normal seeds. If AI deviates from 50% the endosperm often stops developing or degenerates.Based on the degree of abnormality of hybrid seeds, activating values of the ten species were arbitrarily assigned from 0.4 in A. ventricosa (2x) to 3.0 in A.sterilis (6x). For comparison A.strigosa (2x) was used as a standard and set at 1.0. Among the 74 succesful interspecific crosses the AI of the polar nuclei varied widely, from 7 to 375% Generally speaking, AI values of less than 20, 20–30, 30–80, and more than 80% show the R-, Rd+, D+ and E-kernel types, respectively. In other words, if the AV of the pollen parent is < 40 or >160% that of the maternal parent hybrid seeds fail to develop. Thus in double fertilization of Avena or probably angiosperms, the triple fusion of two polar nuclei with a secondary male nucleus is apparently a sexual isolation mechanism, being a barrier to interspecific hybridization.Professor Emeritus of Kyoto University, Faculty of Agriculture, Kyoto.     

Fertility of Brachiaria ruziziensis in interspecific crosses with Brachiaria decumbens and Brachiaria brizantha: meiotic behaviour,pollen viability and seed set

Summary Colchicine induced tetraploid (2n=4x=36) Brachiaria ruziziensis were used as female parent in crosses with apomictic tetraploid species (2n=4x=36) Brachiaria decumbens and Brachiaria brizantha. Tetraploid B. ruziziensis pollinated with B. decumbens set significantly more seed than selfed or crossed with B. brizantha. The crossability between B. ruziziensis and B. decumbens is also better than between B. ruziziensis and B. brizantha. In addition, hybrid seedlings obtained in crosses involving B. brizantha are more frequently lethal. All the viable F₁ hybrids are tetraploid with 36 chromosomes. Meiotic chromosome behaviour suggest that the three species belong to the same genomic group and therefore the same agamic complex. Chromosome associations at metaphase I do not allow to identify fertile and sterile hybrids. The interspecific hybrids averaged a lower fertility than their female parent, but some hybrids were more fertile than their apomictic male parent.