Complex origins of breadfruit (Artocarpus altilis, Moraceae): implications for human migrations in Oceania

Breadfruit (Artocarpus altilis, Moraceae), a traditional starch crop in Oceania, has enjoyed legendary status ever since its role in the infamous mutiny aboard the H.M.S. Bounty in 1789, yet its origins remain unclear. Breadfruit's closest relatives are A. camansi and A. mariannensis. DNA fingerprinting data (AFLP, amplified fragment length polymorphisms) from over 200 breadfruit cultivars, 30 A. camansi, and 24 A. mariannensis individuals were used to investigate the relationships among these species. Multivariate analyses and the identification of species-specific AFLP markers indicate at least two origins of breadfruit. Most Melanesian and Polynesian cultivars appear to have arisen over generations of vegetative propagation and selection from A. camansi. In contrast, most Micronesian breadfruit cultivars appear to be the result of hybridization between A. camansi-derived breadfruit and A. mariannensis. Because breadfruit depends on humans for dispersal, the data were compared to theories on the human colonization of Oceania. The results agree with the well-supported theory that humans settled Polynesia via Melanesia. Additionally, a long-distance migration from eastern Melanesia into Micronesia is supported.     

Development of Triticum turgidum subsp. durum - Aegilops longissima amphiploids with high iron and zinc content through unreduced gamete formation in F1 hybrids

Four different interspecific hybrids involving three different accessions of Aegilops longissima Schweinf. & Muschl. with high grain iron and zinc content and three Triticum turgidum L. subsp. durum (Desf.) Husn. cultivars with low micronutrient content were made for durum wheat biofortification and investigated for chromosome pairing, fertility, putative amphiploidy, and micronutrient content. The chromosome pairing in the 21-chromosome F1 hybrids (ABSl) consisted of 0-6 rod bivalents and occasionally 1 trivalent. All the F1 hybrids, however, unexpectedly showed partial but variable fertility. The detailed meiotic investigation indicated the simultaneous occurrence of two types of aberrant meiotic divisions, namely first-division restitution and single-division meiosis, leading to regular dyads and unreduced gamete formation and fertility. The F2 seeds, being putative amphiploids (AABBSlSl), had nearly double the chromosome number (40-42) and regular meiosis and fertility. The F1 hybrids were intermediate between the two parents for different morphological traits. The putative amphiploids with bold seed size had higher grain ash content and ash iron and zinc content than durum wheat cultivars, suggesting that Ae. longissima possesses a better genetic system(s) for uptake and seed sequestration of iron and zinc, which could be transferred to elite durum and bread wheat cultivars and exploited.     

The breadfruit from colombia-a detailed chemical analysis

Economic Botany - A detailed chemical analysis of the breadfruit (Artocarpus altilis) is presented, followed by an analysis of the fat and a partial amino acid analysis of the globular proteins and...     

Transfer of resistance to the beet cyst nematode (Heterodera Schachtii Schm.) from Sinapis alba L. (white mustard) to the Brassica napus L. gene pool by means of sexual and somatic hybridization

Summary Sexual and somatic hybrid plants have been produced between Sinapis alba L. (white mustard) and Brassica napus L. (oil-seed rape), with the aim to transfer resistance to the beet cyst nematode Heterodera schachtii Schm. (BCN) from white mustard into the oil-seed rape gene pool. Only crosses between diploid accessions of S. alba (2n = 24, S_a1S_a1) as the pistillate parent and several B. napus accessions (2n = 38, AACC) yielded hybrid plants with 31 chromosomes. Crosses between tetraploid accessions of S. alba (2n = 48, S_a1S_a1S_a1S_a1) and B. napus were unsuccessful. Somatic hybrid plants were also obtained between a diploid accession of S. alba and B. napus. These hybrids were mitotically unstable, the number of chromosomes ranging from 56 to more than 90. Analysis of total DNA using a pea rDNA probe confirmed the hybrid nature of the sexual hybrids, whereas for the somatic hybrids a pattern identical to that of B. napus was obtained. Using chloroplast (cp) and mitochondrial (mt) DNA sequences, we found that all of the sexual F₁ hybrids and somatic hybrids contained cpDNA and mtDNA of the S. alba parent. No recombinant mtDNA or cpDNA pattern was observed. Three BC₁ plants were obtained when sexual hybrids were back-crossed with B. napus. Backcrossing of somatic hybrids with B. napus was not successful. Three sexual hybrids and one BC₁ plant, the latter obtained from a cross between a sexual hybrid and B. napus, were found to show a high level of BCN resistance. The level of BCN resistance of the somatic hybrids was in general high, but varied between cuttings from the same plant. Results from cytological studies of chromosome association at meiotic metaphase I in the sexual hybrids suggest partial homology between chromosomes of the AC and S_a1 genomes and thus their potential for gene exchange.     

Chromosome numbers and meiotic analysis in the pre-breeding of <Emphasis Type="BoldItalic">Brachiaria decumbens</Emphasis> (Poaceae)

A total of 44 accessions of Brachiaria decumbens were analysed for chromosome count and meiotic behaviour in order to identify potential progenitors for crosses. Among them, 15 accessions presented 2n = 18; 27 accessions, 2n = 36; and 2 accessions, 2n = 45 chromosomes. Among the diploid accessions, the rate of meiotic abnormalities was low, ranging from 0.82% to 7.93%. In the 27 tetraploid accessions, the rate of meiotic abnormalities ranged from 18.41% to 65.83%. The most common meiotic abnormalities were related to irregular chromosome segregation, but chromosome stickiness and abnormal cytokinesis were observed in low frequency. All abnormalities can compromise pollen viability by generating unbalanced gametes. Based on the chromosome number and meiotic stability, the present study indicates the apomictic tetraploid accessions that can act as male genitor to produce interspecific hybrids with B. ruziziensis or intraspecific hybrids with recently artificially tetraploidized accessions.     

PCR-RFLP of the ribosomal DNA internal transcribed spacers (ITS) provides markers for the A and B genomes in<Emphasis Type="Italic"> Musa</Emphasis> L.

Musa acuminata Colla (AA genomes) and Musa balbisiana Colla (BB genomes) are the diploid ancestors of modern bananas that are mostly diploid or triploid cultivars with various combinations of the A and B genomes, including AA, AAA, BB, AAB and ABB. The objective of this study was to identify molecular markers that will facilitate discrimination of the A and B genomes, based on restriction-site variations in the internal transcribed spacers (ITS) of the nuclear ribosomal RNA genes. The ITS regions of seven M. acuminata and five M. balbisiana accessions were each amplified by PCR using specific primers. All accessions produced a 700-bp fragment that is equivalent in size to the ITS of most plants. This fragment was then digested with ten restriction enzymes (AluI, CfoI, DdeI, HaeIII, HinfI, HpaII, MspI, RsaI, Sau3AI and TaqI) and fractionated in 2% agarose gels, stained with ethidium bromide and visualized under UV light. The RsaI digest revealed a single 530-bp fragment unique to the A genome and two fragments of 350-bp and 180-bp that were specific to the B genome. A further 56 accessions representing AA, AAA, AAB, AB and ABB cultivars, and synthetic hybrids, were amplified and screened with RsaI. All accessions with an exclusively A genome showed only the 530-bp fragment, while accessions having only the B-genome lacked the 530-bp fragment but had the 350-bp and 180-bp fragments. Interspecific cultivars possessed all three fragments. The staining intensity of the B-genome markers increased with the number of B-genome complements. These markers can be used to determine the genome constitution of Musa accessions and hybrids at the nursery stage, and, therefore, greatly facilitate genome classification in Musa breeding.Communicated by H.F. Linskens     

Genomic diversity in aneudiploid (2n = 38) and diploid (2n = 40) Glycine tomentella revealed by cytogenetic and biochemical methods.

Of the 15 perennial species of the subgenus Glycine Willd., G. tomentella Hayata is unique in that it has four cytotypes (2n = 38, 40, 78, and 80) and a wide range of geographical distributions. The objective of this study was to uncover the genomic diversity among accessions of aneudiploid (2n = 38) and diploid (2n = 40) G. tomentella based on crossability rate, hybrid seed and seedling viability, meiotic chromosome pairing of F1 hybrids, and seed protein and protease inhibitor profiles. Aneudiploid and diploid G. tomentella accessions were divided into two (D1 and D2) and three [D3(A,B,C), D4, and D5] groups, respectively, based on previous isozyme studies. Crossability rate, intergenomic hybrid viability, degree of chromosome pairing, total seed protein profiles, and trypsin and chymotrypsin inhibitor banding patterns confirmed the isozyme grouping with minor disagreements. A consistent variation was not observed among the aneudiploid accessions in any method of analysis used in this study. Similarly, cytogenetic analysis and the total seed protein profiles did not show dissimilarity among the accessions from Papua New Guinea (PNG; the D3 group) and north of Mitchell River in Northern Queensland [N.Qld(n)]. However, trypsin and chymotrypsin inhibitor analysis revealed that the PNG accessions were distinctly different from N.Qld(n) accessions. The D4 and D5 group accessions were clearly distinguishable by both cytogenetic and biochemical methods. Thus, this study indicates the presence of four genomic groups among G. tomentella (2n = 38, 40) accessions, including the aneudiploids D1 and D2 in one group and diploids in three groups (D3, D4, and D5). These findings will be useful in further genome analysis and add to our present understanding of the biosystematics of the genus Glycine.     

Meiotic behavior in nonaploid accessions of Brachiaria humidicola (Poaceae) and implications for breeding

Brachiaria humidicola is a grass adapted to seasonally swampy grasslands in Africa; two cultivars, 'common' and Llanero, are widely used in Brazilian pastures. New cultivars are in great demand in order to diversify current production systems to achieve improved quality and yield. Cytological analyses of 55 accessions of this species available from the Embrapa Beef Cattle germplasm collection revealed that 27 are apomictic and have 2n = 54 chromosomes. Chromosome pairing as bi- to nonavalent associations at diakinesis indicated a basic chromosome number in this species of x = 6, as found in other closely related Brachiaria species. Thus, these 27 accessions are nonaploid (2n = 9x = 54). Abnormalities were found in the meiosis of these accessions, at variable frequencies. The most common abnormalities were those related to irregular chromosome segregation, which led to unbalanced gamete formation; but chromosome stickiness, cell fusion, and absence of cytokinesis were also recorded. Although some accessions have a low frequency of meiotic abnormalities, ensuring potentially good pollen viability, these cannot be used in hybridization due to a lack of sexual accessions with the same ploidy level.     

Comparative cytogenetics of hamsters of the genus Calomyscus

Karyotypes of Calomyscus from different regions of Turkmenistan, Iran, and Azerbaijan were studied using chromosome banding (G- and C-banding) and analyses of meiosis in laboratory hybrids. Extensive variation in the diploid number and the number of autosomal arms (FNa) was revealed (2n = 30, FNa = 44; 2n = 32, FNa = 42; 2n = 44, FNa = 46; 2n = 44, FNa = 58; 2n = 37, FNa = 44; 2n = 50, FNa = 50; 2n = 52, FNa = 56). Centric and tandem fusions and heterochromatin changes were identified as the major modes of karyotype evolution in this group. Natural hybrids between individuals with different karyotypes were recorded, and regular chromosome pairing in meiosis was observed in laboratory hybrids. Fluorescence in situ hybridization with a 353-bp BspRI complex tandem repeat indicated that chromosomal repatterning occurred recently within the genus. There is no unequivocal evidence suggesting the role of chromosomal change in the speciation of the populations of Calomyscus examined.     

Diversity of Breadfruit (<Emphasis Type="Italic">Artocarpus altilis</Emphasis>, Moraceae) Seasonality: A Resource for Year-Round Nutrition<Superscript>1</Superscript>

Diversity of Breadfruit ( Artocarpus altilis , Moraceae) Seasonality: A Resource for Year-Round Nutrition. Breadfruit, Artocarpus altilis (Parkinson) Fosberg, is an important but underutilized staple crop cultivated throughout Oceania and much of the wet tropics. Indigenous peoples have selected and cultivated cultivars with different fruiting seasons to effectively extend fruit production for most of the year. In the present study, the seasonality of 219 breadfruit accessions originating from 17 Pacific island groups, the Seychelles, the Philippines, and Indonesia, and now growing in the National Tropical Botanical Garden’s (NTBG) Kahanu Garden, Hana, Hawaii, were evaluated. The predominant season of male flower production for most cultivars was from May to September, and fruit was produced most frequently between August and January. However, there were differences in the duration of the fruiting season from year to year and among cultivars. Over the 10-year period, 14 cultivars did not reliably produce fruit; most of this group were ‘ulu afa’ trees collected from Tokelau. About 24 cultivars exhibited very little seasonality and produced fruit throughout the year. The rest of the cultivars could be clustered into seasonality groups with characteristic fruiting patterns. Comparison of literature values indicates that unlike Hawaii, the breadfruit season in most locations begins around the date the sun reaches zenith prior to the summer months and extends throughout the summer months. Five cultivars asexually propagated from the NTBG collection growing in Kiribati (1°28′ N) exhibit similar fruiting patterns as in Hawaii, except that they begin 2–3 months earlier. These data predict that cultivars with complementary fruiting seasons in Hawaii may experience a temporal shift in their seasonality but will maintain their compatibility when cultivated in different locations and could enable year-round fruit production in many regions.     

Cytological investigations of the interspecific hybrids of Nicotiana tabacum L. x N. glauca Grah

Interspecific amphihaploid and amphidiploid hybrids between Nicotiana glauca Grah. (2n = 24) and N. tabacum L. (2n = 48) cultivars BY 103 and K 326 were analysed. F1 amphihaploids (2n = 36) were viable and completely self- and cross-sterile, and mostly univalents were present during meiosis (with pairing range from 0 to 5). In some meiocytes, meiotic irregularities were observed, such as sporadic chromatin bridges and formation of restitution nuclei. The resultant F1 hybrids were easily converted to amphidiploids (2n = 72) via colchicine treatment of seedlings. The number of univalents and the frequency of PMCs containing unpaired chromosomes indicated that amphidiploids N. tabacum cv. BY 103 or K 326 x N. glauca represented quite a high pairing category. However, they were male sterile because pollen mother cells were arrested at the tetrad stage. The termination of development of PMCs, and consequently male sterility, are very rare in this kind of tobacco hybrids.     

Discrimination of American Cranberry Cultivars and Assessment of Clonal Heterogeneity Using Microsatellite Markers

Cranberries (Vaccinium macrocarpon Ait.) are an economically important fruit crop derived from a North American native species. We report the application of 12 simple sequence repeats (SSR) or microsatellite markers to assess the genetic diversity of cranberry cultivars. We studied 164 samples of 21 different cranberry cultivars, 11 experimental hybrids, and 6 representative accessions of wild species. Genetic cluster analysis, based on 117 SSR alleles, differentiated the major cranberry cultivars. However, some cranberry cultivar subclone variants and mislabeled samples were observed. Consensus genetic profiles identified the most likely clonal representatives of several important cranberry cultivars (e.g., “Ben Lear,” “Howes,” and “Stevens”). The markers were further used to confirm putative parents of several hybrid progenies. The long-term goal of our studies is to identify, preserve, and utilize unique genetic materials to breed improved cranberries. Attaining this goal will help growers maintain sustainability under changing economic and environmental conditions.     

High throughput analysis of grape genetic diversity as a tool for germplasm collection management

Using 20 SSR markers well scattered across the 19 grape chromosomes, we analyzed 4,370 accessions of the INRA grape repository at Vassal, mostly cultivars of Vitis vinifera subsp. sativa (3,727), but also accessions of V. vinifera subsp. sylvestris (80), interspecific hybrids (364), and rootstocks (199). The analysis revealed 2,836 SSR single profiles: 2,323 sativa cultivars, 72 wild individuals (sylvestris), 306 interspecific hybrids, and 135 rootstocks, corresponding to 2,739 different cultivars in all. A total of 524 alleles were detected, with a mean of 26.20 alleles per locus. For the 2,323 cultivars of V. vinifera, 338 alleles were detected with a mean of 16.9 alleles per locus. The mean genetic diversity (GDI) was 0.797 and the level of heterozygosity was 0.76, with broad variation from 0.20 to 1. Interspecific hybrids and rootstocks were more heterozygous and more diverse (GDI?=?0.839 and 0.865, respectively) than V. vinifera cultivars (GDI?=?0.769), Vitis vinifera subsp. sylvestris being the least divergent with GDI?=?0.708. Principal coordinates analysis distinguished the four groups. Slight clonal polymorphism was detected. The limit between clonal variation and cultivar polymorphism was set at four allelic differences out of 40. SSR markers were useful as a complementary tool to traditional ampelography for cultivar identification. Finally, a set of nine SSR markers was defined that was sufficient to distinguish 99.8% of the analyzed accessions. This set is suitable for routine characterization and will be valuable for germplasm management.     

Assessment of gene flow from a herbicide-resistant indica rice (Oryza sativa L.) to the Costa Rican weedy rice (Oryza sativa) in Tropical America: factors affecting hybridization rates and characterization of F1 hybrids

Herbicide-resistant rice cultivars allow selective weed control. A glufosinate indica rice has been developed locally. However, there is concern about weedy rice becoming herbicide resistant through gene flow. Therefore, assessment of gene flow from indica rice cultivars to weedy rice is crucial in Tropical America. A field trial mimicking crop–weed growing patterns was established to assess the rate of hybridization between a Costa Rican glufosinate-resistant rice line (PPT-R) and 58 weedy rice accessions belonging to six weedy rice morphotypes. The effects of overlapping anthesis, morphotype, weedy accession/PPT-R percentage, and the particular weedy accession on hybridization rates were evaluated. Weedy rice accessions with short overlapping anthesis (4–9 days) had lower average hybridization rates (0.1%) than long anthesis overlapping (10–14 days) accessions (0.3%). Hybridization also varied according to weedy rice morphotype and accession. Sativa-like morphotypes (WM-020, WM-120) hybridized more readily than intermediate (WM-023, WM-073, WM-121) and rufipogon-like (WM-329) morphotypes. No hybrids were identified in 11 of the 58 accessions analyzed, 21 accessions had hybridization rates from 0.01% to 0.09%, 21 had rates from 0.1% to 0.9%, and 5 had frequencies from 1% to 2.3%. Another field trial was established to compare the weedy rice-PPT-R F₁ hybrids with their parental lines under noncompetitive conditions. F₁ hybrids had a greater phenotypic variation. They had positive heterosis for vegetative trait and reproductive potential (number of spikelets and panicle length) traits, but negative heterosis for seed set. This study demonstrated the complexity of factors affecting hybridization rates in Tropical America and suggested that the phenotype of F₁ hybrids facilitate their identification in the rice fields.     

Features of recombination of nuclear genome in back crossed offspring of barley-wheat hybrids Hordeum vulgare L. (2n=14) x Triticum aestivum L. (2n+42) with the use of SSR-analysis

The backcross progenies of the barley-wheat hybrids Hordeum vulgare L. (2n = 14) x Triticum aestivum L. (2n = 42) and two alloplasmic lines derived from them were studied using microsatellite markers of barley and wheat. The F1 hybrids and first backcross plants BC1 contained the genetic material of both cultivated barley and the cultivars of common wheat involved in developing of these hybrid genotypes. The genomes of BC3, BC4, and alloplasmic lines contained no microsatellite markers of the cultivated barley, whereas chromosomes of each homeologous group of common wheat were identified. In chromosomes of backcross progenies BC3, BC4, and alloplasmic lines yielded by backcrosses of hybrids and various common wheat cultivars, microsatellite markers of the parental wheat cultivars were shown to undergo recombination.     

“Erussica”, the intergeneric fertile somatic hybrid developed through protoplast fusion between Eruca sativa Lam. and Brassica juncea (L.) Czern.

Summary Hypocotyl calli-derived protoplasts of two cultivars of Brassica juncea (2n=36), a major oil-seed crop, were fused with normal as well as -irradiated mesophyll protoplasts of Eruca sativa (2n=22). The irradiation of the Eruca fusion partner increased the plating efficiency as well as the morphogenic potentiality of the fusion products over the normal fusion. Fertile plants could be regenerated from such fusion products. Analysis of 63 out of 181 plants regenerated showed that, indeed, 11 somatic hybrids (2n=58) and 10 partial somatic hybrids (chromosome number ranged between 50 and 56) had been obtained. Pollen viability (0%–82.9%) and seed set (0%–50%) of the hybrids indicated them to be useful for future studies.     

Chromosomal relationships among cultivars of Citrus reticulata Blanco, its hybrids and related species

Karyotypes of 93 individuals belonging to 18 accessions of mandarins, mandarin hybrids and two related species were analysed with the fluorochromes CMA and DAPI, to identify marker chromosomes. The karyotypes revealed highly differentiated banding patterns and could be classified in four groups (I–IV) according to the presence/absence of chromosomes with three bands (type A) or with two bands (one proximal and one terminal, type B, or both terminal, type C). The accessions of group I exhibited the simplest and homozygous karyotypes (lacking chromosome types A, B and C), represented by `Sunki' and `Cleopatra'. Group II (lacking chromosome types A and B) included three accessions of Mediterranean mandarins and `Cravo' mandarin, all of them with very similar and almost completely homozygous karyotypes. All other karyotypes of groups II and III (lacking chromosome type A) were heterozygous for one or more chromosome pairs and most of them seemed to be hybrid derivatives from non-mandarin accessions. Group IV (with chromosome types A and B) was represented only by two heterozygous hybrids (`Murcott' and `King'). The karyotype of most hybrids agrees with one of the possible combinations resulting from chromosome types segregation from their putative ancestor karyotypes, but at least `Orlando' seemed to be a more complex hybrid. Comparing with banding patterns of other Citrus species, those of group I and the Mediterranean mandarins (group II) are the best candidates to represent C. reticulata (sensu Swingle) as a true species.     

Chromosome markers and alterations in mitotic cells from interspecific Citrus somatic hybrids analysed by fluorochrome staining

Mitotic cells from Rough lemon (Citrus jambhiri Lush.), Ohta ponkan (C. reticulata Blanco) and two somatic hybrid plants obtained from protoplast fusion were analysed by double staining with chromomycin A₃ (CMA) and 4′-6-diamidino-2-phenylindole. Only CMA-positive bands were observed in metaphasic chromosomes. The two parental karyotypes (2n=2x=18) were heteromorphic, yielding some marker chromosomes that could be identified in the somatic hybrids. One of the somatic hybrids had 2n=37 chromosomes, and the possible extra chromosome was distinguishable. The second somatic hybrid was tetraploid (2n=4x=36), with one of the chromosomes bearing a putative structural alteration. Furthermore, aneusomaty and some mitotic abnormalities were also observed in this latter plant. Such irregularities are reported for the first time for citrus somatic hybrids, and their possible causes and implications are discussed. Received: 23 December 1996 / Revision received: 21 May 1997 / Accepted: 16 June 1997     

Constitutive heterochromatin DNA polymorphisms in diploid Medicago sativa ssp. falcata.

A Giemsa C-banding technique was used to study the amount and location of constitutive heterochromatin in diploid (2n = 2x = 16) Medicago sativa ssp. falcata (L.) Arcangeli. Most accessions had the standard C-banding pattern with centromeric bands on all the chromosomes and a prominent heterochromatic band at the nucleolar organizer regions (NOR) of the satellited (SAT) chromosomes. However, we observed in various accessions that constitutive heterochromatic C-bands can exist at the telomeric ends of all the chromosomes. Interstitial bands occurred on the short arms of all chromosomes except for chromosome 3 and on the long arms of chromosomes 1, 2, 3, and 6, only. Rearranged chromosomes such as isochromosomes have been observed for the short arms of chromosomes 2 and 6. This is the first report on the existence of C-banding polymorphisms and the detection of putative isochromsomes in the chromosomes of diploid ssp. falcata which could have contributed to the variation observed in cultivated alfalfa.     

Pennisetum squamulatum: is the predominant cytotype hexaploid or octaploid?

Apomixis is a mode of asexual reproduction where maternal clones are produced through seeds. Consequently, genetic segregation is prevented in hybrid progenies. Pennisetum squamulatum has been used to transfer apomixis into the related sexual species Pennisetum glaucum by the introgression of an apospory-specific genomic region (ASGR)-carrier chromosome. Crosses between P. glaucum and P. squamulatum or Pennisetum purpureum have been relatively easy to make even though P. squamulatum has been reported to have a different basic chromosome number than the other 2 species (9 vs. 7) and to be hexaploid (2n = 6x = 54). Our extensive examination of one accession had shown a chromosome number of 2n = 56. In order to determine if there was a variation among accessions, we counted the number of chromosomes in 5 accessions of P. squamulatum using centromeric and 18S-5.8S-26S rDNA probes as molecular cytological markers. Our results showed that P. squamulatum is most likely octaploid with a basic chromosome number of 7 (2n = 8x = 56) and may belong to the secondary gene pool of Pennisetum. Moreover, a morphologically similar ASGR-carrier chromosome that confers apomixis was observed in all accessions.