Reciprocal effects in true potato seed breeding in short-day length environments

Reciprocal crosses may have significantly distinct performances regarding tuber characteristics in potato, suggesting the importance of cytoplasmic effects in this crop. The selection of parents for true potato seed breeding therefore needs to consider this potential effect when determining the direction of a cross. The aim of this research was to determine whether a broad‐based true potato seed breeding population, developed at the Centro Internacional de la Papa, could be affected by cytoplasmic effects in the short‐day length environments of the tropics. Two random sets of reciprocal biparental crosses were included in the first set of experiments, which were grown in two contrasting Peruvian locations. Only one out of 14 reciprocal crosses showed significantly distinct performance for tuber yield and tuber set. In the second set, only one of each of the 12 reciprocal crosses had distinct performance for vine earliness, days to flowering and flowering intensity, but four of the reciprocal crosses in the second set showed distinct pollen production. The results suggest that cytoplasmic effects in this breeding population are more important for reproductive characteristics such as pollen production than for tuber yield. This finding is not surprising because male sterility in potato results from the interaction between sensitive cytoplasm and dominant nuclear genes.     

Combining ability and correlations for yield components in early generations of potato breeding

A partial diallel set of crosses was made between 14 potato cultivars chosen for their fertility, from those included in a potato breeding programme at the NEIKER – Basque Institute for Agricultural Research. The progeny were grown in completely randomized trials from 1997 to 1999. Performance for yield, tuber number and average tuber weight was analysed in seedling and two clonal generations. Variance estimates due to both general combining ability (GCA) and specific combining ability (SCA) were significant in all generations for all traits under study. However, SCA was more important than GCA in almost all cases. Correlation coefficients among characters, generations, GCA and SCA effects were examined. For tuber yield no relation was obtained between generations; however, average tuber weight and yield were positively associated in all generations. The results indicate that appropriate selection criteria depend strongly on the particular cross. The implication for a breeding strategy are discussed.     

Breeding for impact resistance in potato: use of a pendulum in progeny testing

Breeding for impact resistance in potato is common at the end of a breeding scheme, where there is no more variability for it. The purpose of this study was to propose an objective progeny test using a pendulum and single-year data. Data were recorded over a 5-year period using 275 progenies, organized in small factorial and hierarchical designs. The variability of response of the additive variance made it impossible to generalize parental choice. However, once the crosses were made, the high degree of variability among the progenies and the high level of broad-sense heritability in every case suggest that selection can be carried out efficiently using single-year data.     

Effectiveness of in vitro selection for agronomic characters in potato

Effectiveness of in vitro selection for agronomic characters was studied by finding correlation coefficients between in vitro and in vivo performance of 22 potato genotypes. Evaluation was performed under eight in vitro and two in vivo conditions. Genotypic differences were highly significant for various characters under all in vitro and in vivo conditions. Error mean squares were much lower in in vitro experiments than in in vivo experiments. In vitro selection was found to be highly effective for tuber colour, stem pigment and number of eyes, and moderately effective for average tuber weight, plant vigour and foliage senescence under specific conditions. The results also indicated the possibility of in vitro selection for heat tolerance. For tuber yield and number of tubers, effectiveness of in vitro selection was low to very low, because differences in phenotypic expressions of genotypes were much enlarged under in vitro conditions compared to under in vivo conditions. Correlation coefficients for certain characters were better when in vitro conditions were closer to in vivo conditions. To improve the efficiency of in vitro selection, there is perhaps a need to simulate and identify the in vitro conditions under which genotypic differences are reflected in phenotypes more realistically as they are expressed under in vivo conditions. This revised version was published online in August 2006 with corrections to the Cover Date.     

Behaviour of genetically modified amylose free potato clones as progenitors in a breeding program

Three amylose-free genetically modified potato clones were used both as male and female parents in a breeding program with non-GMO potato clones. Segregation data on the expression of the inserted antisense gene construct in tubers of progeny plants were in agreement with previous molecular analysis of the transgenic clones. The inheritance of the inserted genes was according to Mendelian segregation. Therefore, these clones can be very useful in a breeding program for large scale introduction of amylose free potato cultivars into agriculture. Because of varying number and expression levels of inserts in the GMO-clones, but also because of the varying strength of the endogenous GBSS-alleles of the non-GMO-clones, a segregation into a range of amylose contents occurred. The segregation of the starch colour after iodine staining of pollen of transgenic clones did not follow the obtained segregation in the progeny and was, therefore, not useful in predicting the breeding result. This revised version was published online in July 2006 with corrections to the Cover Date.     

Identifying superior parents in a potato breeding program using cross prediction techniques

Cross prediction techniques were applied to data collected from over 600 hybrid combinations of potato (Solanum tuberosum) and analyzed to determine the potential of using early generation cross prediction techniques to identify superior parental clones. Performance of parental lines based on parameters collected in early generations were compared with the observed frequencies of desirable recombinants with a common parent in the latter stages of a breeding program. Results showed that value of parents, in their ability to produce desirable recombinants in breeding programs, can be predicted using univariate cross prediction techniques. This type of information can be available from early generation progeny trials and could easily be incorporated into a practical potato breeding scheme. This revised version was published online in August 2006 with corrections to the Cover Date.     

Evaluation of FDR diploid and tetraploid parents in potato under two different day-length environments

Five tetraploid clones of potato were crossed to each of eight diploid first division restitution (FDR) 2n pollen producers and four tetraploid males using a line × tester mating design. A total of 59 families were obtained and evaluated under long days at Rhinelander (USA). A subset of 49 families were grown under short days at four Peruvian locations. Tuber yield (kg per plant) of 4x-2x families was similar to, or greater than, that of 4x-4x families in short day environments. The diploid clones had breeding values greater than or equal to those of the tetraploid clones. Only families derived from FDR 2n pollinators combined high tuber yield with stability and high dry matter content. Reciprocal recurrent selection would be the best breeding scheme, in which the diploids are testers of the tetraploids and vice-versa. Intra-population improvement could be achieved through phenotypic recurrent selection. The best materials from both breeding pools could then be crossed to produce tetraploid hybrids through unilateral sexual polyploidization (4x-2x crosses).     

Selection for verticillium wilt resistance in potato breeding populations derived from four cross types

Verticillium wilt (VW) is one of the important yield-limiting diseases for potato production. To develop resistant clones, the potential for early generation selection was studied using three basic selection methods, individual, family, and within family selection, for two clonal generations. A total of 152 clones were derived from four cross types (2x × 2x, 2x × 4x, 4x × 2x and 4x × 4x). Clones were evaluated for maturity, symptom expression, yield and stem colonization in replicated trials. Heritability and selection response for the traits were estimated for each selection method. Direct selection in the second clonal generation and individual selection showed more gain than that from other methods. Both 2x × 2x and 4x × 2x families were higher yielding and had lower stem colonization scores than 2x × 4x and 4x × 4x crosses. Therefore, 2x × 2x or 4x × 2x crosses between carefully chosen parents with high yield and VW resistance may produce offspring with superior performance.     

Top‐cross vs. poly‐cross as alternative to test‐cross for estimating the general combining ability in potato

Test‐crosses for estimating combining ability in potato although useful are difficult to make being labour intensive and time consuming, and due to the problem of sterility. The objective of the present study was to test methods for estimating general combining ability (GCA) of parents based on fewer matings. The GCA of 12 females estimated based on matings with specific single testers (top‐cross) and three types of bulk pollen (poly‐cross) were compared with those estimated by 72 (12 × 6) test‐crosses, using common testers in three types of matings. The study was conducted for 2 years at each of the two locations representing short‐days subtropical conditions in plains and long‐days temperate conditions in hills of India. Analysis of variance of test‐crosses showed that both general and specific combining ability effects were important for tuber yield, late blight resistance as well as general impression, the characters studied. Correlation coefficients among three types of matings showed that bulk pollen matings had poor effectiveness in predicting the GCA of the females, whereas matings with selected individual testers resulted in GCA estimates similar to those based on six testers. Thus use of top‐cross involving selected tester could substantially reduce the number of matings required for estimating the GCA. Among the six testers used, the tester CFK69.1 could estimate the GCA of the females with reasonably high accuracy both for tuber yield and general impression, and ‘Tobique’ for reaction to late blight.     

A comparison between single plant plots and five plant plots for the initial selection stage of a potato breeding programme

Summary The efficiency of single plant selection in the initial selection stage of a potato breeding programme was examined. A random sample of potato clones was grown in single plant plots and also in tworeplicates of five plant plots. After harvest, each plot was visually assessed by three potato breeders; total tuber weight and number of tubers per plant were also recorded, from which mean tuber weight was calculated. The error variances of the single plant plots were found to be significantly greater than from the five plant plots for total tuber weight, mean tuber weight and number of tubers per plant, but not significantly different for breeders' preference. Coefficients of correlation between single and five plants plots were significantly greater than zero for all traits examined, but they were lower than the corresponding correlations between the two replicates of five plant plots. Although a random sample of clones was examined, each clone had been assessed for breeders' preference the previous year. When the clones were grouped according to the previous years preference scores, it was found that the correlation coefficients between single and five plant plots for breeders' preference were inversely related, in magnitude, to the mean preference score of each group. It was concluded that single plant selection was generally ineffective, particularly when only the better clones, from the previous years assessment, are examined.     

The relations of broad nematode resistance to quality characteristics as a consequence of marker-assisted selection in potato breeding programs

Cultivating resistant varieties of potato is the most effective and environmentally sound method of protecting potato crops against pests and diseases. Potato cyst nematodes (PCN) are major nematode pests causing severe constraints in potato production worldwide. There are five pathotypes of Globodrea rostochiensis (Ro1–Ro5) and three of G. pallida Pa1–Pa3. Cultivation of potato varieties with broad nematode resistance may influence the growth of the wide spectrum of PCN pathotypes, but there is limited availability of such varieties on the market. The use of molecular markers allows for the effective selection of resistant genotypes at early stages of breeding. However, the impact of early selection for nematode resistance on the agronomic value of the final selected clones is a cause of concern for potato breeders. This study investigates the relationships between the presence of the combined resistance genes H1, Gro1-4 and GpaV_vrn, which confer resistance to the nematodes, and certain agricultural traits. Clones with broad nematode resistance conferred by the genes H1, Gro1-4 and GpaV_vrn presented yields and tuber morphology traits similar to those of the clones without identified resistance genes.     

Early selection for extreme resistance to potato virus Y and tobacco etch virus in potato using a β-glucuronidase-tagged virus

The Ry_sto gene from Solanum stoloniferum introduced into potato cultivars (Solanum tuberosum L. ssp. tuberosum) confers resistance to potato virus A, potato virus V and potato virus Y (PVY). In addition to PVY, tobacco etch virus (TEV) and a TEV construct that encodes β‐glucuronidase (TEV‐GUS) were inoculated to determine the inheritance of resistance to these viruses in progenies obtained from potato cultivars containing the Ry_sto gene. While cultivars ‘Karlena’ and ‘Delikat’ were susceptible, ‘Bettina’ and clone 927eY were resistant to PVY, TEV and TEV‐GUS, as determined by enzyme linked immunosorbent assay, biotest and GUS assay, respectively. The segregation ratios obtained from the progenies of ‘Bettina’בDelikat’ and 816eY בKarlena’ indicate that resistances to PVY and TEV are governed by one dominant gene or two genes tightly linked in coupling phase. Evidently, Ry_sto confers broad spectrum resistance to potyviruses. TEV resistance could be reliably detected 4 days after inoculation with the TEV‐GUS construct by GUS assay. Therefore, the GUS‐tagged TEV construct can be used for early selection for resistances based on the gene Ry_sto or closely linked genes.     

Analysis of dihaploid induction and production ability and seed parent x pollinator interaction in potato

Summary The production of dihaploids is the first step in a potato breeding program at the diploid level. Dihaploid induction ability, dihaploid production ability and seed parent x pollinator interaction were analyzed for 28 × 3, seed parent x pollinator combinations. This is the first report on significant interaction between seed parents and pollinators. Despite this interaction, IVP 101 had a significantly higher dihaploid induction ability than the widely used pollinators IVP 35 and IVP 48. Previous findings on significant differences in dihaploid production ability between seed parents were clearly confirmed.     

Combining ability of seed vigor and seed yield in soybean

Studies have shown no consensus in relationships between seed yield and vigor in soybean [Glycine max (L.) Merrill]. The lack of information regarding the inheritance of seed vigor prompted this study to determine the types of gene action and combining ability estimates for seed vigor and its related traits. Five high and six low seed vigor soybean genotypes were crossed in a diallel, and selfed to produce 55 F₂ progenies, which were examined, along with the parents, for seed vigor, yield, and seed weight. Significant genotype and environment effects were found for seed vigor and yield. General combining ability (GCA) effects for seed vigor and seed yield were significant (p≤ 0.01) and larger than specific combining ability (SCA) effects. Significant GCA and SCA effects were found for seed weight, indicating that both additive and non additive genetic effects were involved in conditioning seed weight. The ratios of mean square, 2GCA / (2GCA+SCA), were 0.96 for seed vigor and 0.93 for seed yield. These ratios indicated that additive gene effects were more important than non additive gene effects for seed vigor and seed yield in these crosses. Mean seed vigor(83.8%), as determined by accelerated aging germination, and mean seed yield (2,155 kg ha^-1)in high vigor × high vigor crosses were higher than the high vigor × low vigor and low vigor × low vigor crosses. Mean percent accelerated aging germination rates in F₂ populations from diallel crosses were significantly related to mid-parent seed vigor(r² = 0.52^**) and midparent seed size (r² = 0.31^**). These results indicated that levels of seed vigor can be improved through breeding, while maintaining high yields because of the predominance of GCA effects in both seed vigor and seed yield. This revised version was published online in July 2006 with corrections to the Cover Date.     

The use of systems analysis and modelling of growth and development in potato ideotyping under conditions affecting yields

Annual crops are usually only grown during part of the year when environmental conditions are favourable. The factors that limit the length of the available growing season for a crop are discussed from the points of view of meteorology, crop physiology and crop management. Crop yields that may be expected depend on the length of the growing season, how well the available season is used by a growing crop and on environmental factors that define (e.g. temperature, daylength and solar radiation), limit (e.g. the availability of water and nitrogen) or reduce (e.g. pests and diseases) yields. Designing genotypes for such environments requires quantitative knowledge of the influence of environmental factors on the length of the season and on dry matter accumulation and partitioning. This approach is discussed and as example the potato crop is used grown under various climatic conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

The reaction of potato differentials to Phytophthora infestans isolates collected in nature

Available data have been evaluated on the reaction of potato differentials to over 5000 Phytophthora infestans isolates, collected in various parts of the world. The differentials were able to identify up to 11 virulence factors in P. infestans. The isolates differed in virulence expression, depending on the isolate and testing conditions. All 11 virulence factors were found in both ‘old’ and ‘new’P. infestans populations. The resistance of individual differentials was not overcome with equal frequency. The resistance of differentials R5, R8 and R9 was overcome least frequently. This may be due to instability of expression of the respective virulence factors in P. infestans and/or the kind of resistance present in the differentials. Whatever the reason, such resistance may possibly be used in breeding potato cultivars with durable resistance to P. infestans.     

Genetic divergence for cross prediction in potato

To study the effectiveness of genetic divergence for cross prediction in potato, progeny means, heterosis and specific combining ability effects were correlated with parents’ genetic distances (D values) estimated under six in vitro and four in vivo conditions, for tuber yield in 72 crosses (18 × 4) of 22 parents under autumn crop conditions for three successive generations. Genetic distances under in vitro conditions had no relationship with the progeny means for tuber yield. Whereas, those under in vivo conditions in the autumn seasons were positively associated with the progeny means. Similarly, heterosis for tuber yield had a stronger relationship with genetic distances based on an in vivo crop than those based on an in vitro crop. All correlation coefficients between genetic distances and specific combining ability effects were non-significant. The magnitudes of the significant correlation coefficients showed that genetic divergence can be used as an indirect parameter of moderate effectiveness in selecting parents to produce heterotic high yielding progenies. Such cross predictions, however, would be effective only if parents are evaluated under the conditions similar to those under which crosses are likely to be evaluated. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effectiveness of selection at microtuber crop level in potato

Correlation coefficients were computed between microtuber crops and normal seed crop in order to study the selection efficiency for agronomic characters in potato at microtuber crop level. Two types of microtuber, green and white, were produced by inducing microtuberization under light and dark conditions, respectively. Genotypic differences were significant for various characters in both microtuber and normal seed crops. Green and white microtubers were equally useful for selection, judging by the crops raised from them. Selection at microtuber crop level was found to be highly effective for plant height, stem habit, tuber colour and general impression, and moderately effective for internode length, leaf length, leaflet width, stem pigment, plant vigour and tuber shape. Selection efficiency was low for tuber yield, tuber number, average tuber weight, number of nodes, number of stems, leaf colour and number of eyes. The implications of these results for potato breeding and germplasm evaluation programmes are discussed and a procedure for selecting superior genotypes at microtuber crop level is outlined.     

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.     

Expression of nonacclimated freezing tolerance and cold acclimation capacity in somatic hybrids between hardy wild Solanum species and cultivated potatoes

The expression of freezing tolerance was characterized in interspecific somatic hybrids between S. tuberosum (tbr) and two cold-hardy wild species, S. brevidens (brd) and S. commersonii (cmm). The nonacclimated freezing tolerance (NA) and acclimation capacity (ACC, increase in freezing tolerance in response to low nonfreezing temperature), two main genetic components of freezing tolerance, were evaluated separately. In contrast to cmm, which exhibited excellent NA and ACC, the freezing tolerance of brd was mainly due to ACC. However, the ACC of brd was only moderately expressed in the somatic hybrids. The NA of cmm was also suppressed in combination with tbr genomes. However, with acclimation, some of the tbr (+) cmm somatic hybrids achieved freezing tolerance comparable to pure hardy species such as brd used in this study. Analysis of chloroplast DNA type by RFLP markers revealed no significant difference in ACC between somatic hybrids carrying chloroplasts from either tbr or cmm. The reasons for the reduced expression of freezing tolerance from either the brd or cmm parent and the utilization of these somatic hybrids in breeding programs are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.