Wild relatives can contribute to the improvement of fruit quality in pepino (Solanum muricatum)

Narrow intraspecific variation for sugar content in pepino (Solanum muricatum Aiton) hinders the development of new pepino cultivars with improved fruit quality. However, some wild related species have high soluble solids concentration (SSC) and titratable acidity (TA). Generation means analysis was used to estimate genetic parameters for SSC, TA and ascorbic acid concentration (AAC) in two families of interspecific crosses between a S. muricatum parent (Sm-32) and one accession of each of the wild species Solanum caripense (Sc-4) and Solanum tabanoense (St-1). In both families, the additive effect [d] was the only significant parameter for SSC, the alleles of wild species contributing positively to increasing SSC values. For TA, genetic effects associated with additivity [d] and dominance [h] were detected in both families. Additionally, in Sm-32 × Sc-4 nonallelic interactions associated with dominance ([j] and [l]) were significant. For this trait, alleles from the wild species contribute additively to a high value of the character, but are recessive to those of the cultivated S. muricatum. No genetic variation was found for AAC. Broad-sense heritabilities for SSC and TA were intermediate (0.40 to 0.50). Positive significant genetic correlations (around 0.68 in both families) were found between SSC and TA. The results obtained in backcrosses of interspecific hybrids to S. muricatum, together with the estimates of genetic advance after selection, indicate that recovery of individuals with higher SSC and TA and adequate SSC/TA ratio can be achieved in a few backcrosses. In conclusion, this work indicates that wild species S. caripense and S. tabanoense are worthy sources of variation for improving pepino fruit quality. This revised version was published online in July 2006 with corrections to the Cover Date.     

Utilization of genetic resources for the introduction and adaptation of exotic vegetable crops: The case of pepino (Solanum muricatum)

Summary Pepino (Solanum muricatum), a vegetatively propagated plant from the Andean region used for its edible fruits, has been identified as a potential crop for greenhouse cultivation in Mediterranean regions. However, attempts for introducing it have been unsuccessful, either because of the low yield, poor fruit quality, or both. Screening of germplasm under Mediterranean conditions showed that sources of variation for high yield existed in the cultivated genepool and that wild species S. caripense and S. tabanoense could contribute to a considerable improvement of soluble solids content (SSC) of pepino. Progenies obtained after crossing genetically distant (AFLP-based) parental clones were heterotic for yield, and allowed the selection of clones with an improved combination of yield and SSC. As a result of this intraspecific programme, two new improved cultivars (‘Turia’ and ‘Valencia’), which outperform the rest of cultivars available, have been selected for cultivation in Mediterranean conditions. The interspecific programme involved the selection of clones with high yield and SSC in backcross generations. The results show that introgression of genes from the wild species can contribute to improve the SSC of pepino. The backcross programme is in an advanced stage and new cultivars derived from the interspecific crosses are expected to be released in a near future. New prospects for the future in pepino breeding include the genetic transformation, the establishment of a genetic map, and the use of the genomic information from related Solanaceae important crops. All the information obtained is illustrative for the breeding for adaptation of vegetatively propagated crops.     

Interspecific hybridization between Solanum melongena and Solanum macrocarpon: study of the F1 hybrid plants

Summary Egg plant is one of the most common vegetable crop grown in India and other parts of the world. The cultivated egg plant Solanum melongena is found to be susceptible to the shoot and fruit borer (Leucinodes orbonalis). Whereas the species Solanum macrocarpon is resistant. In order to incorporate the resistance of shoot and fruit borer to the cultivated egg plant, an interspecific hybridization between Solanum melongena and Solanum macrocarpon and reciprocals were carried out. The hybrids were found to be sterile, the investigations revealed that the failure of seed set in hybrids is due to the ovule abortion. In order to overcome the sterility, the colchicine has been applied to the interspecific hybrids.     

Enhancement of diploid Solanum chacoense Bitt. using adapted clones of Solanum phureja Juz. & Buk.

Summary F₁ hybrids between Solanum phureja and S. chacoense were studied in a new approach to enhance wild diploid potato species using adapted S. phureja clones as the female parent. S. chacoense parents had few tubers, whereas S. phureja parents had been previously selected for tuberization under long days. Twenty hybrid families were evaluated for haulm vigor, vine maturity, stolon development, frequency of tuberization, tuber set, tuber yield and average tuber weight under naturally occurring increasing or decreasing photoperiods in spring and fall growing seasons in Morocco. The hybrid plants were more vigorous, later maturing with thicker stolons in the spring season. Frequency of tuberization and tuber set did not vary with season. Tuber yield and average tuber weight were higher in fall; the maximum yield was 2.4 and 4.6 kg per hill under spring and fall conditions, respectively. Remarkable variation was found both between and within hybrid families for all traits in this first cycle of enhancement. The selection of vigorous, fertile clones incorporating the S. chacoense genome is possible by using S. phureja as an adapted germplasm source.     

Intra- and interspecific transmission of androgenetic competence in diploid potato species

Summary In an attempt to determine the transmission of androgenetic competence, 10 families resulting from intra- and interspecific hybrids including three reciprocal hybrids were examined in anther culture. Hybrid families were generated between competent clones of Solanum phureja and incompetent clones of S. phureja, S. microdontum and S. berthaultii. S. phureja clones PP5 and A95 (derived by androgenesis of a 2n microspore of PP5) were found to be consistently competent for androgenesis. Androgenetic competence was observed to segregate in all hybrid families with some highly responsive and some unresponsive genotypes in all families. A total of 9,465 cultured anthers have yielded 936 embryoids and 91 plants, including 29 monoploids. The cytoplasm of species lacking competence appeared to have greater influence on the expression of androgenesis in intraspecific than in interspecific hybrids. Expression of androgenesis varied among half-sib hybrid families indicating that competence for androgenesis was influenced by the parents lacking competence. The anther culture data on a backcross between a highly responsive hybrid and its unresponsive parent indicated that competence may be under control of a single dominant gene.     

Evaluation of globe artichoke and related germplasm for resistance to artichoke latent virus

Summary A total of 280 globe artichoke (Cynara scolymus L.) clones were assessed for reaction to naturally occurring artichoke latent virus (ALV) in field trials. Also, the reaction of 17 seed progenies of crosses which produced ALV-resistant clones in the field, 20 self progenies of parental genotypes common to these crosses, self progenies of 17 artichoke cultivars, and 7 accessions ofC. cardunculus andC. syriaca were evaluated in the glasshouse following mechanical inoculation with ALV. The local and systemic presence of ALV, and the relative virus content in infected plants were determined by ELISA.All cultivars and most of the otherC. scolymus material tested (clones and seed progenies) proved to be highly susceptible to ALV. Ten progenies were less susceptible to mechanical inoculation as shown by the moment the virus could be detected, and a slow rate of accumulation of viral antigen. Some plants in two seed progenies possessed apparently a high resistance as could be concluded from the failure of ALV to spread or to multiply in plant.Accessions ofC. cardunculus andC. syriaca were also highly susceptible to ALV. However, an interspecific hybridC. cardunculus x C. syriaca presented the highest level of resistance, the plants remained free of local and systemic infection. Ratios of resistant to susceptible plants in self progenies of resistant plants suggested that this resistance was dominant.     

The breeding of the cultivated potato species solanum x juzepczukii and S. x curtilobum II. The resynthesis of S. x juzepczukii and S. x curtilobum

Summary The possibility of combining anew the genomes of wild and cultivated progenitors of triploid S. x juzepczukii and pentaploid S. x curtilobum by following the known evolutionary pathway of these species was investigated.The resynthesis of S. x juzepczukii was easy, and a wide range of synthetic forms was bred. Among these were forms with higher frost resistance (-5°C) than has been found in natural S. x juzepczukii. The total tuber glycoalkaloid content of several synthetic hybrids was lower than or as low as that of natural clones. Most synthetic hybrids were more vigorous than natural S. x juzepczukii and produced about the same types of tubers as are found in the natural range of variation. The best diploid parents were found in the species S. goniocalyx.The attempt to resynthesize pentaploid S. x curtilobum has not been successful but tetraploid plants were obtained in the process. An explanation for the occurrence of tetraploids resulting from triploid x tetraploid and/or diploid crosses is offered.The newly bred tetraploids contain at least one genome from S. acaule (possibly two) and hybridize easily with ssp. andigena. They thus provide a means for the transfer of S. acaule germ plasm into the tetraploid cultivated gene pool which would profit from the frost resistance of S. acaule.     

Failure in an interspecific cross between Solanum khasianum Clarke and Solanum mammosum L.

Summary Attempts to obtain a successful interspecific cross between Solanum khasianum Clarke and Solanum mammosum L. were unsuccessful, and the probable reasons for this failure were investigated. A reduced number of viable pollen grains and low receptivity of the stigmas of S. mammosum L. and heterostyly in both species are the probable reasons for the failure of the attempted interspecific crosses between S. khasianum Clarke and S. mammosum L.     

Isolation and characterisation of somatic hybrids of diploid Solanum tuberosum and Solanum brevidens and the use of amylose-free starch mutation for detection of introgression

Summary Somatic hybrids of diploid amylose-free (amf) Solanum tuberosum and diploid S. brevidens were made by Poly-Ethylene-Glycol (PEG) or electrofusion methods. For the isolation of interspecific hybrids the use of selection markers (kanamycin and hygromycin resistance) was useful but not essential. In this 2x+2x interspecific combination 4x and 6x somatic hybrids were obtained. Seed set was the best in 4x×4x (S. tuberosum) backcrosses, but seed germination was the best in 6x×4x combinations, using in vitro germination of unripe seeds harvested 25 days after pollination. A high degree of pollen stainability (30–40%) was observed in 7 tetraploid hybrids and very low in all hexaploids. After iodine staining, the recessive amf marker was expressed by a red colour instead of blue, visible in starch-containing cells like columella cells of root tips, (micro)tubers or microspores. As expected, complementation was observed in starch-containing cells of the fusion hybrids. Segregation of the amf marker was clearly observed in microspores of 4x and 6x hybrids. Segregation ratios in the 4x hybrids showed variable recombination frequencies. In the backcross progeny of hexaploid F12-5 with a tetraploid amf mutant one amylose-free recombinant among 67 plants was found, indicating the occurrence of meiotic recombination in the megaspore mother cells.     

Transmission of Verticillium wilt resistance to tetraploid potato via unilateral sexual polyploidization

Summary Verticillium wilt is a serious disease of potato and is caused by the soil-borne fungi Verticillium dahliae and V. albo-atrum. No major cultivar is resistant to this disease. Two diploid interspecific potato clones, C287 and C545, were previously identified with consistently high levels of Verticillium wilt resistance and are thought to have the same genotype for the loci that confer resistance to V. dahliae stem colonization. The purpose of this study was to determine whether resistance to V. dahliae stem colonization could be transferred to the tetraploid level in potato via unilateral sexual polyploidization (USP). Progenies in eight families obtained by crossing C287 and C545 to two tetraploid breeding clones, S438 and S440, and the cultivar ‘Atlantic’ were planted in a V. dahliae-infested field and a field without a significant V. dahliae infestation. Resistance was evaluated relative to C545 and C287. There were differences among families for stem colonization and yield. No differences between the mean stem colonization of C545 and C287 progeny were detected. Family differences due to the tetraploid parents indicate that they contributed resistance to the progeny. Stem colonization data from this experiment were consistent with the proposed complementary two-gene model for Verticillium wilt resistance in the diploid parents. Unilateral sexual polyploidization is an effective method for transferring V. dahliae stem colonization resistance to the tetraploid level.     

Ploidy Manipulation and Introgression of Resistance to Alternaria Helianthi from Wild Hexaploid Helianthus Species to Cultivated Sunflower (H. annuus L.) Aided by Anther Culture

The present investigation discusses the scope for transferring of resistance to leaf spot disease incited by Alternaria helianthi from two hexaploid wild species (H. tuberosus and H. resinosus) to diploid cultivated sunflower. Interspecific hybrids produced between sunflower and these two hexaploid species were partially fertile with tetraploid chromosome status. Backcrosses of these interspecific hybrids with cultivated sunflower resulted in the formation of sterile triploid plants. To overcome the problem of sterility and facilitate backcrosses with cultivated sunflower, anther culture of the tetraploid interspecific hybrids was carried out to bring down their chromosome number to diploid status. Anthers from both interspecific hybrids were cultured on basal Murashige and Skoog media supplemented with varying concentrations of organics and the growth regulators benzyladenine and naphthaleneacetic acid. Anthers of interspecific hybrids involving H. resinosus responded well and regenerated through an embryogenic route at a frequency of 98.7%. But in interspecific hybrids with H. tuberosus, anthers formed callus and subsequently regenerated shoots through an organogenic pathway. DNA ploidy analysis of anther culture plants of interspecific hybrids derived from H. tuberosus crosses was carried out to identify plants with desired diploid status. In vitro screening of parents, interspecific hybrids and anther culture plantlets against A. helianthi showed resistance in 68.5% of the anther culture plants of interspecific hybrids from H. tuberosus and in 24.3% of the plants derived from interspecific hybrids involving H. resinosus.     

Screening and selection of eggplant and wild related species for resistance to <Emphasis Type="Italic">Leveillula taurica</Emphasis>

Screening for resistance to powdery mildew of Solanum melongena and wild related species was made in the field under natural infection conditions. A total of 172 accessions originating from several geographical parts of the world were tested. Single plant selection for resistance was carried out and open-pollination was used. Most S. melongena accessions were susceptible or highly susceptible to the disease. By S₀ to S₃ selection, an increase in the overall powdery mildew resistance level of S. melongena population was obtained and four S. melongena lines possessing a high level of resistance were obtained. Among the wild Solanum species, S. laciniatum and S. nigrum showed to be non-host plants of L. taurica. S. quinquangolare showed no symptoms of powdery mildew, S. linnaeanum, S. aculeatissimum, S. aviculare, S. pseudocapsicum were highly resistant, S. spinosissimum was resistant, S. gilo, S. capsicoides were susceptible or highly susceptible, and plants of S. sisymbriifolium showed a widely variable disease reaction. Four S. melongena resistant lines were obtained: PAVEG 10187 S₃, PAVEG 10196 S₃, P.I. 230279 S₃ and P.I. 419198 S₃. These S. melongena lines together with the resistant wild species could be used for genetic studies, classical breeding programs and biotechnological applications.     

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.     

Resistance to late blight (Phytophthora infestans) in ten wild Solanum species

Nineteen accessions of the tuber-bearing species Solanum berthaultii, S. chacoense, S. leptophyes, S. microdontum, S. sparsipilum, S. sucrense, S. venturii, S. vernei and S. verrucosum were tested for their resistance to late blight in two years of field experiments. Plants were artifically inoculated with zoospores of race 1.2.3.4.5.7.10.11 and the development of the disease was followed. Resistance ratings, calculated as the areas under the disease progress curves (ADPC), demonstrated a high resistance in all accessions except in S. sparsipilum, S. leptophyes and their interspecific hybrid. Segregations suggest that major genes for resistance are present in S. sucrense and S. venturii, and may also play a role in S. verrucosum. It is not yet certain wether the resistance of the other accessions is comparable to the partial and durable resistance of S. tuberosum cultivars like Pimpernel, as inheritance and mechanism have yet to be established. However, segregations suggesting the presence of single major genes with complete dominance were not found in these other accessions. Tuber initiation in the field occurred in only one accession, S. tuberosum ssp. andigena, and maturity of the clones was not related to their resistance. In the other accessions maturity types could not be assessed, as the clones require short day conditions for tuber initiation.     

New sources of resistance of cowpea (Vigna unguiculata) toStriga gesnerioides,a parasitic angiosperm

Summary Thirty-seven accessions of cowpea and yard-long bean were assessed for resistance toStriga gesnerioides. Cowpea plants were grown using anin vitro method, then inoculated with young seedlings ofS. gesnerioides produced from seed from three West African countries. Resistance was assessed by comparing the number and size ofS. gesnerioides tubercles on these accessions with those on a known susceptible cowpea, cv. Blackeye. Two cowpea landraces, APL-1 and 87-2, were completely resistant toS. gesnerioides from Burkina Faso, Mali and Cameroon and partially resistant toS. gesnerioides from Niger. Complete resistance was expressed either as a hypersensitive response of infected root tissues or as a severely retarded development of successful infections. All other accessions, including three samples of yard-long bean were susceptible toS. gesnerioides. The original 87-2 plants segregated for resistance and susceptibility. However, uniformly resistant progeny were obtained by producing seed from vegetatively propagated clones of single resistant 87-2 plants. Resistance of APL-1 and 87-2 toS. gesnerioides was confirmed in pot and field trials. Neither of these cowpeas were resistant toAlectra vogelii. Varieties APL-1 and 87-2 provide additional sources of resistance to most races ofS. gesnerioides, including a newly discovered virulent race from Benin.Abbreviations ICRISAT International Crops Research Institute for the Semi-Arid Tropics - IITA International Institute of Tropical Agriculture - SAFGRAD Semi-Arid Food Grain Research and Development     

Comparison of diploid and tetraploid potato families derived from Solanum phureja x dihaploid S. tuberosum hybrids and their vegetatively doubled counterparts

Summary To determine whether in potatoes the tetraploid level is preferable to the diploid level, especially regarding tuber yield, four diploid (2n=2×=24) Solanum phureja x dihaploid S. tuberosum hybrid parents and their vegetatively doubled, tetraploid (2n=4×=48) counterparts were intermated, which resulted in two F₁ hybrid families at both levels of ploidy. The parents and clones of the F₁ families and their offspring were used in crosses in such a way that in addition Sib₁, Sib₂, F₁×Sib₁, BC₁ and Sib₁×Sib₁ families were produced. Of the first clonal generation of the 12 2 x families and their 12 counterpart 4 x families two tubers per clone were planted in three replications in a field experiment at Sturgeon Bay in 1969; of the parents six tubers were planted in each replication. Data were recorded on 16 characters, including plant height at four different times.The ANOVA's showed significant clone effects within each family for all characters. Computed from all family means as well as from the family means per ploidy level, differences due to family were also significant for all characters except one.As at the 2 x level and at the 4 x level of ploidy the mean phenotypic correlations between characters were of similar magnitude, it is concluded that they are independent of ploidy level.With the exception of eye depth, the mean coefficient of variation was greater at the 2 x than at the 4 x level of ploidy, indicating the greatest response to selection for those characters at the 2 x level. From the differences in family means between the 4 x and the 2 x level of ploidy it was apparent that the 4 x families generally had significantly taller plants, later maturity, fewer tubers, higher mean tuber weight, more tuber yield and more dry matter yield than their counterpart 2 x families. In contrast, the 4 x parents had on the average shorter plants, lower mean tuber weight, much lower tuber yield and lower dry matter yield than their 2 x counterparts.The phenotypic correlation and Spearman's rank correlation between the family means of the 2 x and the 4 x level of ploidy were positive for almost all characters and significant for nearly half of them.From the results it is concluded that 1. in potatoes the 4 x level of ploidy is preferable to the 2 x level, and 2. the performance of 4 x families is predictable from the performance of their counterpart 2 x families.Based on results mentioned in the literature and on the present results, a continued use of S. tuberosum dihaploids in potato breeding needs to be dissuaded.     

Molecular characterization of the progeny of <Emphasis Type="Italic">Solanum etuberosum</Emphasis> identifies a genomic region associated with resistance to potato leafroll virus

Potato leafroll virus (PLRV; Genus Polerovirus; Family Luteoviridae) is one of the most important virus pathogens of potato worldwide and breeders are looking for new sources of resistance. Solanum etuberosum Lindl., a wild potato species native to Chile, was identified as having resistances to PLRV, potato virus Y, potato virus X, and green peach aphid. Barriers to sexual hybridization between S. etuberosum and cultivated potato were overcome through somatic hybridization. Resistance to PLRV has been identified in the BC₁, BC₂ and BC₃ progeny of the somatic hybrids of S. etuberosum (+) S. tuberosum haploid × S. berthaultii Hawkes. In this study, RFLP markers previously mapped in potato, tomato or populations derived from S. palustre (syn S. brevidens) × S. etuberosum and simple sequence repeat (SSR) markers developed from tomato and potato EST sequences were used to characterize S. etuberosum genomic regions associated with resistance to PLRV. The RFLP marker TG443 from tomato linkage group 4 was found to segregate with PLRV resistance. This chromosome region has not previously been associated with PLRV resistance and therefore suggests a unique source of resistance. Synteny groups of molecular markers were constructed using information from published genetic linkage maps of potato, tomato and S. palustre (syn. S. brevidens) × S. etuberosum. Analysis of synteny group transmission over generations confirmed the sequential loss of S. etuberosum chromosomes with each backcross to potato. Marker analyses provided evidence of recombination between the potato and S. etuberosum genomes and/or fragmentation of the S. etuberosum chromosomes.     

Synthesis and preliminary characterization of a new species (amphidiploid) in Cucumis

A successful interspecific hybridization between cucumber (Cucumis sativus L., 2n = 14) and a wild it Cucumis species, C. hystrix Chakr. (2n = 24) was made via embryo rescue. Hybrid plants (2n = 19; 7 from cucumber and 12 from C. hystrix) were sterile, but morphologically uniform. Self-pollination and backcrossing of F₁ hybrid plants to either parent confirmed presence of both male- and female-sterility that were likely caused by lack of homology and improper pairing during meiosis. While the multiple-branching habit, densely brown hairs (on corolla and pistil), orange-yellow corolla, and ovate fruit of F₁ hybrid plants were similar to that of the C. hystrix parent, the appearance of the first pistillate flower was more similar to that of C. sativus parent. The diameter and internode length of the stem, and the shape and size of leaves and flowers were intermediate when compared to the parents. The chromosome number in the hybrid was doubled through somaclonal variation during embryo culture and regeneration process to restore the fertility. Pollen grains were released and fruits with viable seeds matured on fertile, synthetic amphidiploid plants. The results from flow cytometry indicated that, on average, 7.3% of the morphologically unique regenerants had the 4C DNA content of 2.35 pg relative to the 2C DNA content of the original F₁ hybrid at 1.17 pg and, therefore, were likely chromosome-doubled F₁ hybrids (2n = 38). Nutrition alanalysis indicated that the synthetic species had higher protein (0.78%)and mineral (0.35%) content compared to the normal pickling cucumber(0.62% and 0.27%, respectively), and could be considered a new Cucumis crop having a special place in the future agriculture. Preliminarily evaluation indicates that C. hystrix possesses a high level of root-knot nematode resistance, and that this resistance is partially expressed in the interspecific F₁ and chromosome-doubled F₁. This and the fact that the fruit morphology of the fertile amphidiploid differs during the growing season (e.g., short and long fruit) suggest that it could be useful in broadening the germplasm base of cucumber. This revised version was published online in July 2006 with corrections to the Cover Date.     

Parthenogenetic monohaploids (2n=x=12) from Solanum tuberosum L. and S. verrucosum Schlechtd. and the production of homozygous potato diploids

Summary Dihaploid and dihaploid derived clones of Solanum tuberosum and diploid genotypes of S. verrucosum produced 85 viable monohaploids by female parthenogenesis. All were induced using diploid S. phureja clones, homozygous for embryo spot, as pollinator. Frequency of S. tuberosum monohaploids per 100 berries was rather constant in three successive years (14, 17 and 17 respectively). No male and female fertility was found in flowering monohaploids.Colchicine-induced chromosome doubling yielded homozygous s. tuberosum diploids with low pollen quality but good seed fertility.Two diploid self-incompatible species (S. multidissectum and S. berthaultii) produced no monohaploids. The presence of genes for female parthenogenesis in some dihaploids is discussed.