Breeding behavior of yield components and hollow heart in tetraploid-diploid vs. conventionally derived potato hybrids

Summary A factorial mating design with six 4x cultivars as stylar parents mated to three groups of pollen parents (4x cultivars, 4x high protein selections, and 2x S. phureja) was used to study breeding behavior of yield, tuber set, marketable yield and hollow heart in potatoes. Mean yield for the 4x-2x hybrids was equivalent to that of their cultivar parents although yield components, tuber set and average tuber size, were greater and less, respectively, than the cultivar parents. High estimates of general combining ability were found in both pollen and stylar parents for yield, tuber set, marketable yield, number of marketable tubers and hollow heart in all three hybrid populations. Specific combining ability estimates were significant only for yield and hollow heart. Frequency of white-fleshed 4x-2x progeny of five heterozygous yellow-fleshed 2x phureja genotypes was used to confirm cytological identification of a first division restitution mechanism operative in the 2x parents. Although three of these phureja genotypes generated high-yielding progeny, two others produced 4x-2x progeny with mean yield similar to that of hybrids among 4x cultivars.Scientific journal series article 11,601 of the Minnesota Agricultural Experiment Station.     

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.     

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.     

Solanum phureja genes are expressed in the leaves and tubers of aneusomatic potato dihaploids

Summary The expression of leaf isozymes and tuber patatin in dihaploids derived from the Solanum tuberosum cv. Pentland Crown was investigated. Seven of the dihaploids were aneusomatic containing additional chromosomes from the S. phureja dihaploid inducer. Of these, four genotypes expressed leaf isozymes characteristic of the S. phureja dihaploid inducer, and the tubers of three aneusomatic dihaploids contained a S. phureja form of patatin. Aneusomatic dihaploids in which the proportion of cells containing additional S. phureja chromosomes was relatively small (i.e. 1–15%) did not express leaf isozyme markers or patatin bands characteristic of the dihaploid inducer or showed only faint expression of one or two markers. However, those with a high proportion of cells containing additional chromosomes (50–55%) had a range of strongly expressed leaf isozymes that were characteristic of the dihaploid inducer and also expressed the S. phureja tuber patatin.One dihaploid genotype was exclusively euploid (2n<24), yet is expressed a S. phureja leaf isozyme marker and S. phureja tuber patatin, suggesting recombination or chromosome substitution between the genome of the S. phureja dihaploid inducer and the cultivar Pentland Crown.     

Tuber yield and quality of 4x-2x (FDR) potato progenies derived from the wild diploid species Solanum berthaultii and Solanum tarijense

Field experiments were carried out in order to evaluate 4x‐2x families derived from crosses between elite 4x potato cultivars and 2x Tuberosum‐Solanum tarijense and 2x Tuberosum‐Solanum berthaultii clones. Three traits were assessed: total tuber yield (TTY), haulm maturity (HM) and general tuber appearance (GTA). The degree of heterosis of these hybrid families was evaluated by comparison with the respective 2x and 4x parents as well as with seven 4x cultivars. The parental haploid species hybrids derived from S. berthaultii and S. tarijense combined two or more positive horticultural characteristics. Expressed as yield percentage of the 4x parents, the TTY of the families ranged from 53% to 246%. For TTY, the best 4x‐2x hybrid family ranked better than seven out of nine elite 4x cultivars. Some families had GTA scores in the range of the highly selected 4x cultivars. The families, however, were generally later maturing than the 4x parent group. Specific combining ability for TTY and GTA were the only two significant sources of variation observed in this genetic material. Parent‐offspring correlation coefficients were low for all traits, and indicated that parental performance would not be informative at either ploidy level. These results parallel previous investigations with distinct haploid species hybrids where a 4x‐2x breeding scheme was found to be an effective strategy for increasing progeny TTY over the 4x parents. However, the high degree of heterosis for TTY along with good GTA scores observed in certain cross combinations derived from these unadapted 2x species was a somewhat surprising result. Thus far, the importance of these two South American wild potato species, from the potato‐breeding standpoint, has been limited to the fact that they are natural reservoirs of major genes controlling resistance against important diseases and insects. However, the level of performance of some 4x‐2x families in comparative assays with elite cultivars suggests the unanticipated possibility of introgressing genetic factors from S. berthaultii and S. tarijense with positive effects on quantitative traits of horticultural importance along with these major resistance genes.     

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 biosystematic study of the origin of the cultivated diploid potato,Solanum x Ajanhuiri Juz. et Buk.

Summary The origin and nature of the diploid cultivated potato species S. ajanhuiri Juz. et Buk. was studied. Several lines of evidence indicate that S. ajanhuiri might be derived from natural crosses between primitive cultivars of the diploid species S. stenotomum and the wild species S. megistacrolobum. Morphological comparisons were made between S. stenotomum x S. megistacrolobum F₁ hybrids and naturally occurring S. ajanhuiri to investigate this hypothesis. Comparisons were also made between S. ajanhuiri x S. stenotomum crosses and the F₂ generation of the first-mentioned cross.Crosses between the two major groups of S. ajanhuiri cultivars, Ajawiri and Yari, showed not only genetic breakdown but also a wide range of phenotypic variation similar to those of artificial F₂ families of S. stenotomum x S. megistacrolobum. Furthermore, there was strong evidence showing that the Yari group of S. ajanhuiri could almost certainly be an F₁ S. stenotomum x S. megistacrolobum hybrid, whereas the Ajawiri group could be a backcross of an F₁ hybrid to S. stenotomum. These results added further support to the hypothesis of a hybrid origin of S. ajanhuiri, as well as indicating its putative parents. It is suggested that this hybridogenic taxon be retained at the species level under the name Solanum x ajanhuiri.     

The effect of chromosome doubling on fertility,meiotic behaviour and crossability of Solanum etuberosum × S. pinnatisectum

Summary Doubling the chromosome number of sterile F₁ hybrids of S. etuberosum × S. pinnatisectum resulted in fertile allotetraploid hybrids (code 4x-EP) with nearly normal chromosome pairing at meiosis. Many seeds were obtained from selfing and sibbing, but seed set varied considerably. The 4x-EP hybrids were successfully crosses as females with S. stoloniferum and S. polytrichon and as males with S. verrucosum. The crosses with S. stoloniferum and S. polytrichon were carried out reciprocally and unilateral incompatibility was detected. Crosses of 4x-EP hybrids as males onto S. phureja and S. tuberosum cv. Gineke produced parthenocarpic berries only. Nearly 1700 flowers of colchicine-doubled S. pinnatisectum × S. bulbocastanum and the trispecific hybrid (S. acaule × S. bulbocastanum) × S. phureja did not yield a single berry after pollination with a pollen mixture from 20 4x-EP plants.It is concluded that the use of the bridging species S. pinnatisectum and either S. stoloniferum, S. polytrichon or S. verrucosum may enable gene transfer from the non-tuberous Etuberosa species to potato cultivars.     

Introgression of the low browning trait from the wild Mexican species Solanum hjertingii into cultivated potato (S. tuberosum L.)

Internal discoloration of tubers resulting from impact damage (blackspot bruise) is a serious quality problem in potato production and utilization, reducing profits to growers and increasing costs for processors. Resistance to blackspot bruise has been identified in the wild species Solanum hjertingii and is therefore a potential germplasm resource for genetic resistance to this problem. A bridging cross between S. hjertingii and a cultivated diploid clone was used to produce a triploid hybrid population that exhibited very low tuber browning potential, indicating a dominant pattern of inheritance for this trait. The triploid progeny were subjected to in vitrochromosome doubling and the resulting hexaploid clones were screened for browning potential. A hexaploid clone selected for low browning was reciprocally crossed with cultivated S. tuberosum cultivars exhibiting high susceptibility to blackspot bruise. Tubers obtained from the seed progeny of these 4x-6x crosses (hereafter referred to as the BC₁ populations) were evaluated for browning potential and polyphenol oxidase (PPO) activity. Tubers from the BC₁ populations displayed a very low potential for melanin production, while PPO activity was quite variable. The low Pearson correlation coefficient (r² = 0.45), between browning potential and PPO activity suggests that the mechanism of blackspot bruise resistance derived from S. hjertingii cannot be explained simply as a reduction in the initial PPO activity. The expression of substantial resistance to browning and dominant expression pattern in these BC₁ progeny indicate that utilizing genetic elements derived from S. hjertingii provides a robust approach for developing blackspot bruise resistant potato varieties. This revised version was published online in August 2006 with corrections to the Cover Date.     

Reciprocal differences between hybrids of Solanum tuberosum Groups Tuberosum (haploid) and Phureja

Summary Yield components including total tuber yield, total tuber number, vine maturity, vine vigor and average tuber weight as well as seed germination and pollen stainability were studied in 28 exact reciprocal hybrid families, between 12 Solanum tuberosum Group Tuberosum (haploid) (TH) and 12 S. tuberosum Group Phureja (PH) clones in a NC Mating Design II in 4 sets of 3 by 3 reciprocal hybrids at three locations, one in Minnesota (USA) and two in Morocco. TH parents were insensitive to daylength whereas most PH clones required critical photoperiods equal to or less than 14 hours for tuberization. Mean reciprocal differences were significant for most traits at all 3 locations. Hybrids with TH cytoplasm were superior to those with PH cytoplasm by 18% for tuber yield, 21% for tuber number, and 9% for vine maturity, but inferior to hybrids with PH cytoplasm by 11% for vine vigor and 19% for time to reach 80% seed germination. Reciprocal differences were not detected for average tuber weight. Six reciprocal crosses revealed significant differences for pollen stainability but the overall mean difference (one location) was not significant. Possible reasons for the observed reciprocal differences included gamete selection, pseudo-self-compatibility and/or interaction of cytoplasm with both nuclear genes and environment. Differences for critical photoperiod of TH and PH parents were also demonstrated as a probable source of reciprocal differences.Scientific Journal Series Article 14896 of the Minnesota Agricultural Experiment Station supported in part by the U.S. Agency for International Development, contract no. 608-0160, IAV Morocco Project.     

Competition between 2x and x pollen in styles of Solanum tuberosum determined by a quick in vivo method

Summary Pollen competition in vivo was studied by placing two pollen samples on one stigma and comparing the length of the tubes under a U.V. microscope. Comparisons involving pollen of Solanum tuberosum and S. phureja showed that 2x pollen grew faster than x pollen, both in 2x and x styles. Consequences for induction of dihaploids in S. tuberosum are discussed.     

Analysis of pollen tube growth in situ to investigate self-incompatibility in the wild potato Solanum commersonii

Summary Pollen tube growth was investigated in a diallelic crossing design with seven genotypes of the diploid wild potato species Solanum commersonii, accession O/S UR-9, CIP 762459. Pollen tube growth in the style was recorded using a combined quantitative and qualitative evaluation scale. Clear-cut differences in pollen tube growth behavior in compatible and in partially or completely incompatible crosses were detected. Diallelic crossing of the seven randomly chosen genotypes, intercrossing within two progeny families, and backcrossing of two progeny populations to the parents revealed the existence of a one-locus gametophytic system of stylar incompatibility. The S-allele status of all genotypes investigated was determined.     

Integration of transgenes into sexual polyploidization schemes for potato (Solanum tuberosum L.)

Most potato transgenic research has focused on development of resistance to pathogens and modification of potato physiology. Many transgenes, particularly those conferring pathogen resistance, could substantially lower potato production costs in developing countries. However, transgenes have not been reported in sexually propagated 4x-2x potato hybrids commonly grown in developing countries. Two transgenes,the Bacillus thuringiensis cry3Aa endotoxin protein gene and the PVY°coat protein gene, were engineered intodiploid and tetraploid potato using Agrobacterium tumefaciens-mediated transformation. Cry3Aa was produced at high levels in several lines while the PVY° coat protein was not expressed. Diplandroid and tetraploid genotypes were crossed to produce transgenic 4x-2x hybrids. Genetic transformation had no discernable effect on fertility ofthe primary transformants, germination of4x-2x seed derived from the transformants and agronomic performance(tuber set, average tuber weight and total tuber yield) of the 4x-2xhybrids. The transgenic 4x-2xhybrids produced non-viable pollen and could only be crossed as female parents. Results suggest that transgenes, such ascry3Aa, could be expressed in 4x-2x hybrids to lower costs of production with no significant effect on plant phenotype. This revised version was published online in July 2006 with corrections to the Cover Date.     

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)     

Genetic base of Indian potato selections as revealed by pedigree analysis

Pedigree analysis of the 77 advanced Indian potato selections showed that their origin could be traced to only 49 ancestors out of which 29 were exotic, which accounted for 69.52% of the total genomic constitution and maximum contribution (40.65%) was by 10 ancestors from U.K. Breeders tendency was to involve adapted advanced clones as immediate parents. Coefficient of relationship between pair of selections ranged from 0 to 0.68. The success of the parents used was not related to region-specific choice of parents. The findings are discussed in the context of genetic uniformity and the relevance of having separate breeding programmes for the three major areas of potato cultivation in India. Attempt has also been made to give some directions for choosing selections for use as parents and release as cultivars based on their coefficient of relationship so as to broaden the genetic base of the future potato cultivars.     

Early generation selection for potato tuber quality in progenies of late blight resistant parents

Developing disease resistant cultivars is one of the major objectives for a potato (Solanum tuberosum L.) breeding program, but many resistant clones have not achieved commercial acceptance because of late maturity and non-marketable tuber characteristics. Selection for tuber quality should have greater emphasis inbreeding disease resistant cultivars. The objectives of this study were 1) to evaluate the ability of late blight (Phytophthora infestans (Mont.) de Bary) resistant parents to transmit chip-processing (tuber appearance, specificgravity, and chip-color) or tablestock (tuber appearance) quality to the offspring and 2) to compare selecting for tuber quality in single-hill versus eight-hill generations. We made crosses among eight unadapted potato cultivars (B0718-3,Bertita, Bzura, Greta, Libertas, Stobrawa, Tollocan, and Zarevo) with reported late blight resistance with adapted susceptible cultivars/breeding clones to generate 95populations (4,750 seedlings). Approximately 10% of the progeny from each cross were selected from single-hill plots based on tuber appearance, number, shape, and internal defects. These selected clones (408) were evaluated for tuber appearance, specific gravity, and chip-color. The same evaluations in the following year were made on tuber samples from eight-hill plots. Libertas and Tollocan were the best parents for transmitting chip-color; B0718-3, Zarevo, and Tollocan for transmitting tuber appearance; and Bzura, Libertas, and Zarevo for transmitting high specific gravity to the highest percentage of the offspring. Overall, 50% and 56% of the clones based on single- and eight-hill generation, respectively, were considered to possess chip-processing quality; over 90% of the clones had acceptable tablestock quality. A total of 71% of the clones possessing acceptable chip-processing and 95% of the clones possessing acceptable tablestock quality selected in both generations were identified in single-hill plots. The evaluation of tuber quality characteristics in single-hill generation not only permitted the identification of clones with acceptable chip-processing and tablestock, but also increased the amount of clonal information for the following generation of selection. In crosses between late blight resistant and susceptible clones, selection for tuber quality traits can be initiated in single-hill generation using a moderate selection intensity and precede late blight testing. This revised version was published online in August 2006 with corrections to the Cover Date.     

Breeding for resistance to potato tuber moth,Phthorimaea operculella (Zeller), in diploid potatoes

Summary Sixty-two 2x families were generated by intermating 16, 2x clones and evaluated for resistance to potato tuber moth (PTM), Phthorimaea operculella under natural infestation in a storage at San Ramon, Peru and in laboratory tests. The following conclusions could be drawn: (1) relatively simple inheritance was observed for resistance derived from Solanum sparsipilum (spl), (2) the high level of resistance of the original spl has been transferred, undiminished, into an advanced 2x population, (3) simple phenotypic selection was successfully applied to transfer resistance into an improved 2x population, (4) there was a strong indication of reciprocal effects, however spl cytoplasm is not essential for the expression of nuclear resistance genes, (5) antibiosis and antixenosis are the mechanism of PTM resistance in this population, and (6) 4x × 2x crosses could be used to transfer the resistance into commercial cultivars.     

Low temperature tolerance of biomass and pollen germination in potato clones of Andean and European origin

Summary The temperature-related performances of six tetraploid potato clones, two Andean, three European, and one hybrid, were compared by cultivating them in growth chambers under one of two temperature regimes: 10°C day/4°C night or 20°C day/10°C night. Preformance was measured in terms of biomass production and pollen germinability.For dry matter yield at second harvest, the temperature-related performance ratios (performance at 10°C/4°C divided by performance at 20°/10°C) were highest for the Andean clones, intermediate for the Andean/European hybrid clone 201₅×S₁₂, and lowest for the European cultivars. The ability of the European cultivars to maintain their normal rates (i.e. rates at 20/10°C) of biomass production when cultivated at low temperatures varied greatly, with clone S₃ performing most poorly at 10/4°C.For pollen germinability in vitro, performance ratios were highest for the hybrid clone 201₅×S₁₂ and lowest for the European clones.The high tuber yield of hybrid 201₅×S₁₂ at 10/4°C can be attributed to its low-temperature tolerance and it being daylength neutral.     

A breeding method for accelerated development of cold chipping clones in potato

Summary The development of potato cultivars having acceptable chipping quality after cold storage is important because it reduces costs to growers while ensuring continuity of supply to processors throughout the year. Over 5100 single-hill progenies were planted to select for cold chippers. A breeding procedure for selecting and evaluating these progenies for cold chipping ability was used, and 38 desirable genotypes were identified. These selections were increased in the second clonal generation and evaluated for yield, specific gravity, and seven chipping treatments of varying storage duration/temperature/reconditioning duration and were compared to standard chipping cultivars Atlantic, Monona, Norchip, and Snowden. Twenty-two selections yielded Norchip the highest yielding cultivar, while 15 selections had a specific gravity Atlantic, the highest specific gravity cultivar. Snowden was the best chipping cultivar overall and some selections were not significantly different than Snowden. Overall, nine selections combined high yield and specific gravity with the ability to produce attractive chips from 4° C. Single-hill selection for cold chipping could potentially save four years in the breeding process by the identification of good parents, the recycling of good parental cross combinations, and the evaluation of progeny for chipping earlier in the breeding program.     

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.