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1.
The Endosperm Balance Number (EBN) of over 80 species and subspecies of the tuber-bearing Solanums and their close non-tuber-bearing relatives representing 13 taxonomic series has been determined, with no species assigned to more than one EBN level. Among North American species, most diploids are 1EBN, most tetraploids are 2EBN and all hexaploids are 4EBN; however, among South American species most diploids are 2EBN, most tetraploids are 4EBN and again all hexaploids are 4EBN. Thus species may be isolated from others of the same ploidy level by EBN differences, e.g., 4×(2EBN) from 4×(4EBN), while other species differing in ploidy but having the same EBN may be intercrossed, e.g., 4×(2EBN) and 2×(2EBN). Chromosome doubling or 2n gametes can be used to make a lower EBN species compatible with a higher EBN species. These findings also explain the major crossing difficulties previously inherent in the use of North American species in potato improvement. They also have direct implications for potato improvement, barring the occurrence of other incompatibility barriers. Any 4×(4EBN) cultivar is endosperm compatible and thus will cross with 4×(4EBN) and 6×(4EBN) species. The 2×(2EBN) haploids of 4×(4EBN) cultivars likewise will hybridize with 2×(2EBN) and 4×(4EBN) species. All 2×(1EBN) species are crossable with 2×(2EBN) haploids through 2n gametes or chromosome doubling.Cooperative investigation of the U.S. Department of Agriculture, Agriculture Research Service, and the Wisconsin Experiment Station.  相似文献   

2.
Three wild potato species with different ploidies and Endosperm Balance Numbers (EBN) were crossed in a complete diallel design and the development of the embryo and endosperm as well as the type of seeds produced were analyzed. The compatible crosses – intraspecific intra-EBN and interspecific intra-EBN – produced more than 89% plump seeds, whereas in the incompatible crosses – intraspecific inter-EBN and interspecific inter-EBN – more than 85% of the seeds were not as well developed or were shrunken. The histological analysis revealed that inviable seeds had less developed or collapsed endosperms and thicker endotheliums than viable ones. A gradation of crossabilities was observed among species. The self-compatible species Solanum acaule had good performance as a female but not as a male parent. Among the self-incompatible species, Solanum gourlayi was the best male parent but had a poor performance as a female, whereas Solanum commersonii had an intermediate behavior. Differences in crossabilities among genotypes within species were also observed. These results can not be solely explained by the EBN hypothesis. It is, therefore, suggested that the EBN may be part of a more complex system of interspecific barriers acting at the pre- and post-zygotic levels. This revised version was published online in July 2006 with corrections to the Cover Date.  相似文献   

3.
Endosperm balance number (EBN) was used to predict the outcome of crosses among a series of complex polyploid interspecific hybrids between Trifolium repens and T. nigrescens. Normal endosperm development, and hence seed‐set, requires a 2 : 1 maternal : paternal EBN ratio in endosperm. Based on their crossability, T. repens (2n = 4x = 32) and T. nigrescens (2n = 2x = 16) were assigned the same EBN of 4. This number was used to predict the relative success of different backcrosses and intercrosses. The method was further developed to successfully predict which gametes would effect fertilization when complex polyploid hybrids with uneven genomic constitutions (5x, 7x) produced mixtures of euploid and aneuploid gametes. This was achieved by allocating EBNs to all potential gametes of complex polyploids. Allocation of gamete EBNs enabled not only prediction of the success or failure of a cross, but also which gametes would be filtered to effect fertilization in complex crosses. This enhanced ability to understand the potential outcomes of complex crosses will increase the efficiency of plant breeding programmes that integrate secondary gene‐pools into economic species by interspecific hybridization.  相似文献   

4.
Overcoming hybridization barriers in potato   总被引：3，自引：0，他引：3  
The cultivated potato is a major crop worldwide. It is a high input crop with complex quality requirements at harvest and during storage. Potato breeders are fortunate to have access to a very diverse and accessible germplasm resource. Wild Solanum relatives provide genetic diversity as well as genes for valuable production and quality traits. In most cases, crossing success can be predicted based on endosperm balance number (EBN), or effective ploidy, of the parents. Crossing barriers between most wild species and the cultivated potato are the consequence of differences in EBN and can be easily overcome using ploidy manipulations and bridge crosses. The most common ploidy manipulations include haploid extraction to reduce EBN and 2n gamete production to increase EBN. Additional methods to produce fertile interspecific hybrids include mentor pollination, embryo rescue, hormone treatments, reciprocal crosses, selection of cross‐compatible genotypes and somatic fusion. Knowledge of crossing barriers and mechanisms to overcome them allows potato breeders access to the rich gene pool in the genus Solanum.  相似文献   

5.
Summary The extent and pattern of polyploidy in the tuber-bearing Solanums varies among the many taxanomic series that have been identified in this subsection of Solanum. While several series appear to be entirely diploid, others exhibit a range of ploidy levels from 2x to 6x, and some contain only polyploid species.In many diploid, triploid and tetraploid species 2n gametes (gametes or gametophytes with the sporophytic chromosome number) have been detected. Both 2n eggs and 2n pollen occur. 2n gametes provide the opportunity for unilateral and bilateral sexual polyploidization. The genetic determination and consequences of sexual polyploidization strongly suggest that 2n gametes have been the major instrument in the polyploid evolution of the tuber-bearing Solanums. Somatic doubling of species and interspecific hybrids appears to be of very limited importance.New evidence for the occurrence of 2n eggs and 2n pollen in many species is reported, and data from the literature are added to illustrate the widespread distribution of 2n gametes throughout the subsection. A very high correlation is found between polyploidy and 2n gametes, and its significance is discussed. Proof is presented for the occurrence of alleles governing 2n pollen production in the cultivated tetraploids, providing additional evidence for the hypothesis that 2n gametes have been involved in their origin.Multiple unilateral and bilateral sexual polyploidizations are proposed for the origin of the cultivated tetraploids: this accounts for the large variability encountered in this group, which closely resembles that of the related diploids. Similar evolutionary pathways are hypothesized for the other polyploid complexes in the subsection.A scheme is proposed in which participation of both 2n and n gametes link together all ploidy levels in the tuber-bearing Solanums, thus overcoming the ploidy barriers and providing for gene flow throughout the sympatric species of the subsection.Paper No. 2093 from the Laboratory of Genetics. Research supported by the College of Agricultural and Life Sciences, a gift from Frito-Lay, Inc., and grants from NIH (GM 15422) and The International Potato Center. Plant materials, unpublished chromosome counts and technical help of IR-1 are gratefully acknowledged.  相似文献   

6.
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.  相似文献   

7.
Genetic resistance to potato cyst nematode is considered as one of the most effective means of increasing yield and reducing nematode infestation levels in potato fields. In this study, resistance to this nematode was successfully transferred from diploid tuber-bearing Solanums to the tetraploid gene pool using a 4x-2x breeding approach. More specifically, resistance from Solanum vernei, S. sparsipilum and haploids of S. tuberosum group Andigena was introgressed into conventional tetraploid clones, using first division restitution (FDR) 2n gametes. Furthermore, some of the FDR diploid parents had similar breeding values as advanced resistant tetraploid clones which were developed only after several cycles of selection against the potato cyst nematode. This revised version was published online in July 2006 with corrections to the Cover Date.  相似文献   

8.
The cultivated potato has over 200 extant wild relatives many of which contain genes valuable for disease resistance, hardiness, processing and agronomic traits. Crossability of these wild species directly with the cultivated potato is complicated by several reproductive phenomena such as stylar and ploidy barriers and Endosperm Balance Numbers (EBN). However, a systematic analysis of crossability with many of these wild relatives has never fully been examined. Reciprocal crosses were made between cultivated potato and over 400 wild potato accessions; stylar barriers and 2n gamete production were examined as was the fertility of many of the putative hybrids. Generally, the seed/fruit ratio increased the more closely related the species were to the cultivated potato. However, a few crosses were successful in spite of predicted failure due to ploidy or EBN differences. This revised version was published online in August 2006 with corrections to the Cover Date.  相似文献   

9.
Summary Bacterial ring rot of potato (Solanum tuberosum) has not been successfully controlled through management of certified seed. Therefore, the identification of immunity to this pathogen and its introgression into the cultivated potato is vitally important to the potato industry. Immunity was detected in the disomic tetraploid 2EBN (Endosperm Balance Number) species S. acaule. Immune and nonimmune parents were crossed in a 4 × 4 mating scheme that consisted of four immune × immune crosses, four immune × nonimmune, four nonimmune × immune, and four nonimmune × nonimmune crosses. Analysis was performed on the 16 F1 populations and 54% of the progeny was found to exhibit an immune response to inoculation with Clavibacter michiganensis subsp. sepedonicus (Spieck & Kotth.) Davis et al. Immunity appears to be conferred by two dominant alleles, one at each locus, and may be associated with minor or modifying genes. Phenotypic expression of nonimmune progeny ranged from susceptible to resistant, probably due to minor or modifying genes. These results indicate that it may not be difficult to transfer immunity into the cultivated potato.  相似文献   

10.
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.  相似文献   

11.
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.  相似文献   

12.
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.  相似文献   

13.
Summary Pollen grain germination, pollen tube growth and micropylar penetration were investigated in intraspecific and intergeneric crosses involving Brassica napus L. ssp. oleifera (Metzg.) Sinsk. (oil-seed rape or fodder rape), xBrassicoraphanus Sageret (Raparadish) and diploid (2x) and tetraploid (4x) accessions of Sinapis alba L.(white mustard). For the reciprocal intergeneric crosses between B. napus and xBrassicoraphanus no effective barriers to pollen tube growth on stigmata or in styles were observed. The resulting low frequency of hybrid plants was mainly associated with a low rate of ovules with micropylar penetration per siliqua or with embryo abortion. Hybrid plants could be obtained without use of embryo rescue. In reciprocal crosses between B. napus and S. alba 2x or 4x incongruity barriers were observed on the stigma, in the style, and in the ovary resulting in a low frequency of ovules with micropylar penetration per siliqua. Open flower-pollination compared to bud-pollination generally was the more favourable procedure for pollen grain germination and pollen tube growth in crosses involving S. alba, but for micropylar penetration and seed set no differences were observed. Crosses between S. alba 2x () and B.napus () were found to result in a higher frequency of ovules with micropylar penetration as compared to reciprocal crosses or crosses with S. alba 4x. All reciprocal crosses between B. napus and S. alba 2x or 4x were unsuccessful when no embryo rescue was applied. Embryo rescue shortly after pollination, i.e. 2 to 5 days, however, resulted in hybrid seeds and plants, but only when applied to crosses between S. alba 2x () and B. napus (). The possible effects of the genome constitution, taxonomic distance and the parthenogenetic and parthenogenesis inducing ability of the parental genotypes on the observed malfunctions at the pre-and/or post-zygotic stage of the pollen grain-pistil interactions are discussed.Abbreviations DAP Days After Pollination - IAA Indole-3-Acetic Acid - kin Kinetin  相似文献   

14.
Ietje W. Boukema 《Euphytica》1982,31(3):981-989
Summary The inheritance of the resistance to Didymella lycopersici was studied on F3- and Bc1-lines from interspecific crosses of L. esculentum with L. hirsutum and with L. hirsutum glabratum. The resistance is not monogenic and is inherited in dominant fashion. The high h2 values based on line means offer possibilities to seleet efficiently for enhanced levels of resistance.  相似文献   

15.
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.  相似文献   

16.
J. Chat    P. Y. Dumoulin    Y. Bastard  R. Monet 《Plant Breeding》1996,115(5):378-384
Interspecific crosses between Actinidia arguta (In = 4x) and A. deliciosa (2n = 6x) were carried out in order to transfer cold resistance from a wild to the cultivated kiwifruit species. Of the 65 progenies obtained and analysed by flow cytometry, 16 were diploid, one was tetraploid and 48 were, as expected, pentaploid. Morphologically, the diploids can be distinguished from the pentaploids by their longer lamina. The existence of 5x progenies confirms the existence of gene flow between A. arguta and A. deliciosa, whereas the existence of 2x and 4x progenies provides evidence for parthenogenesis within the Actinidia genus following an interspecific cross. The study demonstrates the utility of flow cytometry for genomic analysis of progenies derived from interspecific crosses. Possible uses of both sexual and parthenogenetic progenies in the breeding and genetics of kiwifruit are discussed.  相似文献   

17.
Summary The occurrence of natural triploid hybrids between tetraploid Solanum acauleBitter subsp. acaule and diploid S. infundibuliformePhilippi both tuber-bearing Solanums, in the Puna region of Jujuy, is reported. The natural hybrids, which are morphologically intermediate between their putative parents and completely male sterile, have a somatic chromosome number of 36. The artificial hybrids, obtained by crossing both parental species, closely resemble the natural ones thereby confirming their parentage. It is proposed to designate the natural hybrids Solanum x viirsooi hybr. nov.  相似文献   

18.
Summary Interspecific hybrids were generated from crosses between 4 Zinnia angustifolia clones (maternal parents) and an inbred line of Z. elegans Orange King (paternal parent) to investigate the influence of Z. angustifolia genotype on morphology and flowering of hybrids. Leaf length, leaf width, flower diameter, number of ray petals, and days to flowering of interspecific hybrids were significantly influenced by Z. angustifolia clonal genotype. Genetic analysis of hybrid morphology and development was performed using 2 parental Z. angustifolia clones and 18 F1 (9 seedlings from each of 2 populations generated from reciprocal crosses between parental clones) as maternal parents in crosses with Z. elegans Orange King. Genotype of Z. angustifolia significantly affected plant height, fresh weight, number of nodes, and days to flowering of interspecific hybrids. Control of these traits was through the Z. angustifolia nuclear genome. Improvement in interspecific hybrid morphology or development, i.e., shorter stature, earlier flowering, or larger flowers, may be possible by selection of superior-performing Z. angustifolia clones as parents.  相似文献   

19.
The objectives of this study were to evaluate the tolerance to low temperatures and tuber soft rot in hybrids between Solanum commersonii and Solanum tuberosum. The experimental materials consisted of F1 triploid, BC1 pentaploid‐near pentaploid and BC2 tetraploid–near tetraploid hybrids. The F1 triploids had a freezing tolerance and acclimatization capacity closest to S. commersonii. This indicated that the endosperm barriers which prevent the introgression of 1EBN S. commersonii into 4EBN S. tuberosum had been overcome. Indeed, the triploids produced 2n eggs, thus giving a compatible maternal to paternal EBN ratio in the hybrid endosperm generated by the 3x(2EBN) × 4x(3EBN) crosses. The tolerance to low temperatures of BC1 and BC2 hybrids was lower than that of the F1. However, a number of genotypes were identified which were able to withstand temperatures down to ‐5°C. Some BC2 hybrids were also tested for their tolerance to tuber soft rot, and some resistant hybrids were detected. A number of them combined the capacity for cold acclimatization with tolerance to tuber soft rot. These hybrids have an EBN of 4; they are fertile and have been used in backcrosses with 4EBN S. tuberosum.  相似文献   

20.
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.  相似文献   

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