Transmission of bacterial wilt resistance by First Division Restitution (FDR) 2n pollen via 4x×2x crosses in potatoes

Summary Resistance to bacterial wilt (Pseudomonas solanacearum) found in tuber-bearing Solanum species was transferred into a diploid potato breeding population. Simultaneous selections were made for agronomic characters, production of first division restitution (FDR) 2n pollen, and resistance. Diploid resistant genotypes were identified via inoculation with a virulent isolate (CIP-204) of race 3 of Pseudomonas solanacearum. These resistant diploid genotypes were crossed to susceptible tetraploid potatoes. An investigation was made to assess whether resistant diploid genotypes transmit resistance to bacterial wilt, which is a quantitatively inherited trait, to tetraploid potatoes via FDR 2n pollen. Tetraploid seedlings from 4x×2x crosses were inoculated with the same isolate CIP-204, and the percentage of surviving seedlings was scored. Some 4x×2x families from resistant diploid genotypes demonstrated a high level of survival rate. The transmission of bacterial wilt resistance was achieved by the use of FDR 2n pollen. It was speculated that a female x male interaction effect on the survival rate of the evaluated progeny may exist. Selecting proper 4x and 2x parents would be important for obtaining a higher frequency of transmission of resistance to bacterial wilt in the progeny.     

Induction of haploids and aneuhaploids in colchicine-induced tetraploid Solanum chacoense Bitt.

Summary The species Solanum chacoense BITT. (2n=2x=24) is a tuber-bearing, self-incompatible species which is important both for breeding and for genetic research. It crosses readily with most other tuber-bearing Solanum species including the common potato S. tuberosum (2n=4x=48). Gametophytic incompatibility hampers research in and utilization of this species. Doubling the chromosome number by colchicine makes it a self-compatible autotetraploid. By crossing selfed progeny of 4x-S. chacoense with a number of haploid-inducing diploid Solanum species a high yield of different dihaploid S. chacoense individuals (2n=24) could be obtained from one originally diploid clone: S. chacoense CPC 1153. More than 160 haploids showing a large variability were identified. The average haploid frequency was 53.7 per 100 berries. Most hybrid plants (70–100%) from four 4x × 2x crosses studied were tetraploid. The frequency of triploid hybrids was low (0–10%). Haploid-inducing capacity of fifteen male parents used in this study varied from 0–141 haploids per 100 berries. Careful examination of 156 haploids revealed 15 viable aneuhaploids (2n=25, 26, and 27), i.e. 9.4%. The potential value of these aneuhaploids is discussed.About 50% of the haploids were sufficiently male fertile to use them in crossing. A few of them set berries after selfing. Five aneuhaploids including the one with 27 chromosomes were successfully crossed as females with a diploid hybrid clone.     

Production of intraspecific aneuploids in the genus Solanum. III. Intraspecific aneuploids

Summary The triploids recovered from 4x×2x crosses in three Solanum species were very vigorous and although few seeds/fruit were obtained when the triploids were crossed to diploids, the extensive crossing programme produced sufficient seed of four species cph, chc, pnt and tar. The average seed set for the 3x-2x crosses was an extremely low 3.5 seeds/fruit.Approximately 90% of the progeny of the 3x–2x crosses were aneuploids with chromosome numbers of 2n=25–29. The frequency of the aneuploids in the three species that were studied was chc 93%, pnt 94% and tar 92%. The aneuploids of chc and tar were extremely vigorous and fertile and they were used as females in crosses to the marker stocks. The aneuploids of pnt were vigorous, but the crossability barrier of pnt prevented their use in crosses to the marker stocks. A number of the aneuploids produced seed upon being selfed, but the ability to produce self seed may be related to the pseudo-compatibility of the parental clones. In only 7 aneuploids was there an indication that the self fertility was due to overcoming the self-incompatibility barrier as a result of competition-interaction of the aneuploid gamete.Plants in the families from the triploid-diploid crosses had a tremendous amount of variation in morphological characteristics (leaf shape, size and color; berry shape, color and degree or verrucose spotting: and plant habit and vigor). A large portion of variation exhibited in these families was due to the normal genetic segregation of the heterozygous parents. It was impossible to distinguish the aneuploids from their diploid sibs especially those having only one or two additional chromosomes because they were as vigorous and fertile as the diploids.There were some preliminary indications of the existence of distinct morphological characteristics among the aneuploids (separate petals, long berries, and extreme verrucose berries). However, there was no indication that these traits were due to the dosage effect of the extra chromosome. If it should be determined that this was true, it would be extremely useful in associating genes with chromosomes and determining the phenotypic effects due to the presence of an additional chromosome.Cooperative investigations of the College of Agriculture and Life Sciences, University of Winconsin and the Agricultural Research Service, U.S. Department of Agriculture, Madison, Wisconsin. Supported in part by grants from the Research Committee of the Graduate School, University of Wisconsin.First and third author respectively, formally Graduate Research Assistant Department of Horticulture, University of Wisconsin, Madison, and Geneticist Agriculture Research Service, U.S. Department of Agriculture, University of Wisconsin, Madison. Present address: Department of Genetics. University of Wisconsin, Madison, Wisconsin 53706 and Head Breeding and Genetics Department, International Potato Center, Lima, Peru.     

Evaluation of resistance to verticillium wilt in diploid,wild potato interspecific hybrids

Summary Verticillium wilt (V. albo-atrum Reinke & Berthold or V. dahliae Kleb) threatens potato (Solanum tuberosum L.) production in most growing areas of the world. Genetic resistance offers the most cost-effective and environmentally-sound control measure. However, there is a dearth of genetic and breeding information on resistance to verticillium wilt in potato, because of obscure parentage of some standard cultivars and the complex segregation at the tetraploid level. The wide range of genetic variability in wild relatives of potatoes can be potentially useful as a source of disease resistance, such as verticillium wilt resistance. Six diploid, wild, interspecific Solanum hybrids involving resistant x resistant and susceptible x resistant crosses, were assayed for verticillium wilt resistance under greenhouse conditions to evaluate their potential as sources of verticillium wilt resistance. The cross between S. gourlayi Oka. and S. chacoense Bitt. and its reciprocal had the most resistant progenies based on mean colony counts. No simple mode of inheritance can be proposed based on the observed segregation ratios. However, the continuous distributions observed on verticillium wilt disease response among hybrid families indicate that inheritance of resistance may be polygenic and complex. In addition, skewness of colony count distributions toward the resistance parents were characteristic of all resistant x susceptible crosses suggesting that resistance may be dominant. By contrast, the susceptible x susceptible cross showed a more normal distribution. Overall, the cross between S. gourlayi and S. chacoense showed the most potential as a source of verticillium wilt resistance.     

Inheritance of early blight resistance in diploid potatoes

Early blight (Alternatia solani) is a fungal disease in hot and humid environments, which causes leaf, stem and tuber lesions. Early blight resistance should be incorporated into potato cultivars because the fungicide spraying is an expensive solution for developing countries. The diploid cultivated species Solanum tuberosum group Phureja and group Stenotomum are sources of resistance alleles. The elucidation of the inheritance for early blight resistance must help to decide what could be the best breeding procedure to improve this diploid germplasm and transfer the resistance to the tetraploid level. Three experiments were carried out under controlled and field conditions to determine the heritability of this trait using nested and diallel mating designs with haploid, species and haploid-species hybrids. The narrow-sense heritability estimates were relatively high (0.64–0.78). This means that additivity was the most important type of gene action for determining resistance to early blight at the diploid level. The results suggested that diploid parents showing highest levels of resistance, throughout the cycle of disease development, can be used in 4x×2x crosses to obtain resistant tetraploid progenies to this fungal disease.     

Crossability relationships of andean potato varieties of three ploidy levels

Summary The formation of triploids in 4x×2x crosses was studied in primitive potato varieties from the Andes of Peru and Bolivia. Triploid frequencies were as high as 80–100% in some crosses, although the actual number of seeds was low. The triploid block varied according to the tetraploid parent, and was never entirely absent. The likelihood of triploid formation was studied in 4x×2x crosses where pollen from both tetraploids and diploids of known genotype was used. Even though the diploids were chosen for their non-production of 2n gametes, as detected in pollen grain size, 66% of the hybrids were tetraploid. The amount and probability of gene-flow between the tetraploid and diploid gene-pools, either directly through 2n gametes or through triploid bridges was examined. Most of the 20 cultivars of S. x chaucha produced seed in crosses with tetraploids and diploids, although the fertility of the latter crosses was four times higher. The female fertility of some of the triploids thus allows them to exchange genes with the tetraploids and diploids, although gene-flow occurs more rapidly through direct transfer from diploids to tetraploids via 2n gametes.     

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.     

Comparison of haploid Tuberosum-Solanum chacoense versus Solanum phureja-haploid Tuberosum hybrids as staminate parents of 4x–2x progenies evaluated under distinct crop management systems

The relative value of haploid Tuberosum-Solanum chacoense (TCH) vs. Solanum phureja-haploid Tuberosum (PTH) hybrids as male parents in 4x–2x crosses was estimated under two distinct crop management systems. The first experiment (E#1) was carried out with supplementary irrigation (three times a week) at Hancock-Wisconsin (WI) with 27 families derived from a sub-set of crosses involving 11 tetraploid (4x) cultivars and 10 diploid (2x) clones (3 TCH; 6 PTH; and one TCH-PTH hybrid clone). A second experiment (E#2) was conducted at Rhinelander-WI with the same group of families but it was only rainfed with no supplementary irrigation being provided during the entire course of the assay. For comparison, a complete set of 4x and 2x clones (used as parents) was also planted in each location. Three traits were evaluated in both locations: total tuber yield (TTY), haulm maturity (HM), and plant vigor (PV). Altogether, 18 out of 27 and 10 out of 27 families outyielded the corresponding 4x parents at E#1 and E#2, respectively. Significant differences among all 4x–2x families and among families of the TCH group were observed for TTY at both locations. No difference was found for TTY among families of PTH and [TCH–PTH] groups at each location. A total of 75% and 50% of the TCH families outyielded their 4x parents at E#1 and E#2, respectively. The percentage of families from the PTH group that outyielded their 4x parents at E#1 and E#2 was 83.3% and 25%, respectively. The TCH group had on average lower HM scores than PTH (at both locations) indicating some segregation for earliness. However, the HM values of the 4x–2x families were, in general, higher than those observed for the 4x parents. For PV, the PTH group mean was significantly higher than TCH group only in E#1. The PV values of the 4x–2x families were higher than the 4x-parent group at both locations. Our results indicated that TCH hybrids might be as good parents as PTH to increase the TTY of 4x–2x progenies. In addition, the TCH families displayed a slight higher level of performance under more stressful growing conditions than PTH hybrids as indicated by the assay at Rhinelander. Therefore, selection of genetic materials with potential broad range of adaptation seems to be feasible with both hybrids via the 4x–2x breeding scheme. 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.     

Breaking the crossability barriers between disomic tetraploid Solanum acaule and tetrasomic tetraploid S. tuberosum

Summary A combination of compatible second pollinations and embryo rescue was applied for systematic production of true tetraploid hybrids from crosses between disomic tetraploid Solanum acaule and tetrasomic tetraploid potato, S. tuberosum. Several genotypes of tetraploid potatoes were pollinated with S. acaule, and the compatible second pollinations were made on the following day, with a genotype of S. phureja, IvP 35 to promote fruit development. Embryo rescue was carried out in 21 families, 14 to 27 days after the first pollination. A total of eight plants were obtained from the embryo rescue and their chromosome numbers were counted in the root tips. Three of the eight plants were identified as tetraploid, and five others as diploid. Morphology, isozyme banding patterns, and pollen stainability, as well as potato spindle tuber viroid (PSTVd) resistance, indicated the hybrid nature of the three plants. This is the first report of successful tetraploid hybrid production between disomic tetraploid S. acaule (4x) and tetrasomic tetraploid potatoes. Seed set from the crosses between one of hybrids and diploid potatoes indicated workable levels of both male and female fertility for introgression of valuable genes from S. acaule into the cultivated potato gene pool. The methodology used may be applied to other disomic tetraploid tuber-bearing Solanum species and with some modifications also to distantly related solanaceous species and genera.     

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.     

Evidence for second division restitution as the basis for 2n + n maternal chromosome transmission in a sugarcane cross

Chromosome doubling in megaspores of sugarcane was first reported over 70 years ago and was seen primarily in interspecific crosses between Saccharum officinarum and Saccharum spontaneum. Since then, there has been robust debate on the mechanism(s) responsible for this doubling with hypotheses ranging from second division restitution (SDR), post-meiotic restitution to fusion of the two innermost cells of the megaspore tetrad. In the present study, we sought to answer this question using molecular marker data from a cross which demonstrated 2n + n chromosome transmission between an S. officinarum clone (IJ76-514; 2n transmission) and a commercial hybrid (Q165 ; n transmission). Molecular marker data from four other bi-parental crosses were used to form a “base-line” for expected segregation ratios (n + n transmission). Comparison of segregation ratios showed that 2n transmission from the S. officinarum clone could be identified by the increased segregation ratios, a characteristic of both SDR and tetrad cell fusion. Simulation was then undertaken based on current understandings of sugarcane meiosis, to determine if the observed segregation ratios matched those simulated for SDR/tetrad cell fusion. Analysis of the simulation confirmed that most and perhaps all progeny appear to have been formed from a mechanism akin to SDR/tetrad cell fusion. The genetic consequences of this are discussed.     

Adaptation to day length and yield stability of families from 4x×2x crosses in potato

Summary The objectives of this research were to evaluate the response to day length and the yield stability of environments of 4x families from 4x×2x crosses in potato. Twenty families from crosses between 4x female parents with different genetic backgrounds and 2x parents, which formed 2n pollen by first division restitution and were selected under short days, were evaluated in four locations in Peru (short days) and during two years at Rhinelander, USA (long days). Results from the USA indicate that the 4x progeny from 4x×2x crosses were very late maturing and poor yielding.Further, vine maturity, tuber size and tuber number are highly correlated with the percentage of Tuberosum germplasm in the parentage. Results from Peru indicate that most 4x×2x families had high tuber yield, stability for yield and high dry matter content. The lack of correlation between rank order of the parental value, based on progeny testing, of the male and female parents among short day locations and between short and long day environments indicates that selection of parents must be done in each location in which the final product, either a family or a clone, will be used.International Potato Center (CIP)     

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.     

Manipulation of the 2n-gametes frequencies in Solanum pollen

Besides the normal gametes, some 2x potato genotypes produce 2n-gametes as well. In interploidy crosses (2x×4x, 4x×2x) 2n-gametes are necessary, whilst in intercrossing at the diploid level they are undesirable.By means of a set of simply constructed sieves, pollen of several 2n-pollen producing progenitors have been sifted. It was possible to increase the 2n-pollen content of low 2n graded pollen to interesting levels. No decrease in in vitro pollen vitality due to the treatment could be demonstrated. The in vivo pollen vitality of the aqueous pollen suspension was satisfactory to very good under cool weather conditions.     

Variation for crossability in a reciprocal,interspecific cross involvingHordeum vulgare andH. lechleri

Summary Several crossing series including the hexaploid (2n=6x=42), South American speciesHordeum lechleri and diploid (2n=2x=14) cultivated barley (H. vulgare) were performed. Barley functioned better as the paternal than as the maternal parent in all cases. Viable offspring were only obtained from one hybrid combination when barley was used as the maternal parent. There was an environmental influence on the success of crosses. A high seed set was counteracted by a lower germination frequency. The outcome is that different crossing series give similar results. As a result of chromosome elimination and in a few cases duplication of especially the barley chromosomes, the chromosome numbers of the offspring (239 plants) varied from 2n=21 (trihaploids ofH. lechleri) to 2n=30. Fifty-five % of the plants were euploid with the number expected for a hybrid (2n=28). The frequency of hyperploids, euploids, hypoploids (2n=22–27) and trihaploids varied by year, locality, type (winter/spring) of barley,H. lechleri population, and crossing direction.     

The occurrence of 2n-pollen and the origin of sexual polyploids in dihaploid roses (Rosa hybrida L.)

Haploidisation by in situ parthenogenesis of 4x R. hybrida resulted in the production of some dihaploid roses (2n=2x=14) able to produce viable pollen. A cytological study of microsporogenesis revealed that, although the first meiotic reductional division occurred normally, the second (equational) division was characterised by frequent abnormalities which concerned spindle formation and led to unreduced gametes of First Division Restitution (FDR) type. Analysis of the hybrid progeny of a parthenogenetically derived male fertile dihaploid plant revealed a selective advantage of the 2n-pollen, especially in the case of an hybridization with a tetraploid female parent. Moreover, crosses carried out among dihaploid partners always resulted in hybrids with ploidy levels≥ 3x. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Structural hybridity in the series Etuberosa of the genus Solanum and its bearing on crossability

Summary Three non-tuberous Solanum species of series Etuberosa-S. brevidens (brd), S. etuberosum (etb) and S. fernandezianum (frn)-upon selfing displayed compatible berry set but seed set varied and on an average was below normal. Also in crosses between species berry set was unhampered (apart from the crosses with short-styled frn as pollen plant), but the interspecific hybrid plants showed various degrees of sterility and a low average seed set upon selfing. Cytological investigations revealed that in pure species-plants semi-sterility regularly occurred and was invariably associated with multivalent associations (up to decavalents). This may account for the decreased seed set found after selfing and intercrossing. Meiosis in F₁-hybrids was even more irregular, 75% of the pmc's showing multivalents involving from 4 to as many as 22 chromosomes in various complicated configurations in which even numbers of chromosomes were predominant. However, anaphase I separation was numerically balanced in more than 75% of the cells. Pollen stainability was very low in the hybrids.It is concluded that heterozygous translocations occur within each species and that the genomes of the species are differentiated by means of gross structural changes. In view of this and of the high chromosome pairing affinity the genome symbols E ₁, E ₂ and E ₃ are proposed for etb, brd and frn, respectively.Crossability of the Etuberosa species with tuberous S. pinnatisectum (pnt) was slightly more successful (average 2.7 seeds/berry) than that of Etuberosa species hybrids with S. pinnatisectum (average 1.7 seed/berry). In the hybrids studied-etb × pnt, frn × pnt and (brd × etb) × pnt-a high frequency of univalents was the rule (range 6–16 per cell). In addition rod bivalents (4–8 per cell) and a low frequency of trivalents occurred (0–3 in the biparental hybrids and 1–4 in the trispecific hybrid).Utilization of the valuable characters of Etuberosa for potato breeding is dependent on the degree of gene exchange between Etuberosa chromosomes and chromosomes of tuberous species. In the diploid hybrids such exchange, though limited, may take place through bivalents and trivalents. As shown in this article such diploid hybrids are non-crossable because of complete male and female sterility. Only after doubling the number of chromosomes the hybrids are fertile and can be used for further crosses.