Genetic linkage of QTLs for late blight resistance and foliage maturity type in six related potato progenies

A set of test crosses of diploid potatoes was used to identify QTLs for foliage resistance against Phytophthora infestans and QTLs for foliage maturity type, and to assess their genetic relationship. The most important locus for both traits was found on chromosome 5 near marker GP21: the allele of marker GP21 that is associated with resistance to late blight is also associated with late foliage maturity. An additional QTL with a small effect on foliage maturity type was identified on chromosome 3, and additional QTLs for late blight resistance were found on chromosomes 3 and 10. Another QTL was detected on chromosome 7 when resistance was adjusted for the effect of foliage maturity type. The additional QTLs for resistance against P. infestans on chromosomes 3 and 10 seem to be independent of foliage maturity type and are not affected by epistatic effects of the locus on chromosome 5. The effects of the additional QTLs for resistance are small, but early maturing genotypes that necessarily have the allele for susceptibility for late blight on chromosome 5 may benefit from the resistance that is provided by these QTLs on chromosomes 3 and 10.     

The effect of seed tuber weight on characters in the first and the second clonal generation of potato populations

Summary Fifty-five populations from crosses of three Solanum tuberosum ssp. tuberosum cultivars and seven selected S. tuberosum ssp. andigena clones adapted to the northern temperate zone have been studied in the seedling and the first and second clonal generation. This paper only deals with the effect of seed tuber weight on various important agronomic characters which are selected for in early clonal generations, independently of the type of crosses.The results showed that a distinction has to be made between the effects of differences in seed tuber weight within and between seedlings, i.e. genotypes. The effects on the first clonal generation of differences in weight of the tubers within seedlings and those of the first heaviest tubers between seedlings were similar and comparable to seed tuber weight effects within cultivars. In contrast to the within seedlings effects the between seedlings effects proved to be heritable. This is concluded from the fact that, on average, independent of seed tuber weight seedlings with larger first heaviest tubers performed better in the second clonal generation than those with smaller ones. Notably plant height, general impression, date of maturity and particularly tuber yield and mean tuber weight in the second clonal generation were related to the weight of the first heaviest tubers of the seedlings. The potentialities of these findings for an early generation selection are discussed.     

Effects of plot size and selection intensity on efficiency of selection in the first clonal generation of potato

To improve efficiency of selection in first year clones, investigations were carried out on the use of two-plant plots instead of one-plant ones. Extended plot size did reduce the environmental variance of characters connected with tuber yield, but had only a slightly positive effect on the selection result, as was concluded from a comparison between the first and the second clonal generation. Only lowering selection pressure in the first year clones seems to be a solution for optimizing selection efficiency, as could be concluded by comparing simulated selection percentages of about 5%, 20% and 50%.Owing to genotype × harvest time-interaction for the majority of traits important to either seed or ware potatoes, assessments at both harvest times are necessary for proper evaluation.     

Correlations within and between characters between and within generations as a measure for the early generation selection in potato breeding

Summary The main objective was to determine the reliability of selection in early generations of a potato breeding programme. To that end data were recorded in the seedling generation, an early and a late harvested experiment of the first clonal generation and the late harvested second clonal generation on up to 30 genotypes of 55 populations from crosses of three Solanum tuberosum ssp. tuberosum cultivars and seven selected S. tuberosum ssp. andigena clones, adapted to the northern temperate zone. Phenotypic correlations were computed per population in almost all possible combinations within and between characters between and within generations. The mean across 55 comparable correlation coefficients was considered to be the best estimate of the relationship concerned.In all four experiments data were collected on number of tubers, tuber yield and mean tuber weight; in the three experiments of the two clonal generations in addition on date of emergence, number of stems, plant height, haulm type, and general impression of the underground plant parts and in the two late harvested clonal generations on date of maturity and under water weight.The results confirmed that general impression which is largely decisive for the selection in early generations, is a little reliable measure. However, from the magnitude of the mean correlations between experiments within characters it must be concluded that selection for most of these separate characters is hardly more reliable. An exception was date of maturity for which a mean correlation was determined of r=0.78 between the two clonal generations. The second strongest mean correlations, perhaps surprisingly, were for tuber yield, viz. r=0.52 between the seedling generation and the late harvested first as well as the second clonal generation and r=0.63 between the late harvested clonal generations. From the correlations between characters it appeared that for a high tuber yield, besides its components, in particular an early emergence, a large plant height, an attractive haulm type and a late maturity are of importance.The main conclusion is that, as long as not enough seed tubers are available for the layout of statistically sound experiments, a negative instead of a positive selection should be applied to avoid the rejection of most of the valuable genotypes in an early stage. On the basis of the results directives for the application of such a negative selection are given.     

Correlation between late blight resistance and foliage maturity type in potato

The genetics of race-non-specific foliage resistance against Phytophthora infestans, of foliage maturity type, and of their association in potato (Solanum tuberosum) were studied. Six progenies were derived from a half-diallel set of crosses between diploid potato clones that represented a broad pool within the genus Solanum and were free of any of the 11 known R genes for late blight resistance. The progenies were evaluated for resistance to late blight and for foliage maturity type, and five of them showed a significant correlation between the two traits. The correlation did not account for all variation that was present for both traits, as reflected in the analysis in which the relative AUDPC values were adjusted for foliage maturity type. The present study adds to previous results: resistance against P. infestans always coincides with late foliage maturity. However, the results also indicate that some selection for late blight resistance without affecting the foliage maturity type should be possible.     

Half-sib progeny evaluation and selection of potatoes resistant to the US8 genotype of Phytophthora infestans from crosses between resistant and susceptible parents

The objectives of this study were to evaluate the use of potato (Solanum tuberosum L.) late blight (Phytophthora infestans (Mont.) de Bary) resistant parents in cultivar development and identify superior clones possessing moderate to high late blight resistance combined with acceptable maturity and tuber quality. Ninety-five crosses were made between eight unadapted parents with reported late blight resistance (B0718-3, Bertita, Bzura, Greta, Libertas, Stobrawa, Tollocan and Zarevo) and susceptible parents (cultivars or advanced breeding clones) adapted to North American growing conditions. A total of 408 field selected clones were assessed for late blight resistance in the greenhouse and in the field using a mixture of US8 P. infestans isolates (A2 mating type, metalaxyl resistant) that overcame all known R-genes except R8 and R9. Clones with ≤ 10% infected foliar area in the greenhouse test or ≤ 0.30 RAUDPC (relative area under the disease progress curve) value in the field in 1998 were re-tested in 1999. A total of 118 (29% of 408) putative late blight resistant clones were selected. The eight late blight resistant parents differed in both the ability to transmit late blight resistance and in the level of resistance transmitted to the progeny. The Tollocan and B0718-3 families (half-sib progeny) had the greatest degree of resistance and frequency of resistant clones. Scott-Knott cluster analysis ranked 79 clones (67% of 118) in the high and moderate late blight resistant groups. Among these 79 clones, 19 clones had vine maturity equal to or earlier than mid-season combined with acceptable tuber quality. Further selection in 2000 resulted in eight advanced selected clones (six from Tollocan and two from B0718-3 families) with the same level of resistance as the parent combined with vine maturity and tuber quality equivalent to Atlantic, a standard cultivar for chip processing in North America. The results indicate that this breeding approach can be used to select parents for late blight resistance breeding and to identify superior clones with high levels of late blight resistance and marketable vine maturity and tuber quality. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of resistance to late blight (Phytophthora infestans) in potato

Under epiphytotic conditions for late blight in spring seasons, data were recorded on its intensity four times at 4 days intervals from the start of the disease in the field, in 114 (19 females × 6 males) progenies and their parents planted in randomized complete block design in the years 2005 and 2006. Regression and stepwise regression analysis showed that observations during the rapidly increasing phase of disease between initial and last phase of disease are more important than the observations at initial and last phase of the disease. Combining ability analysis on a sub-set of 68 progenies showed that the additive component of genetic variance was more important than the non-additive component of variance in inheritance of quantitative resistance to late blight. The per se performance of the parents does give an idea about their general combining ability (GCA). However, selection of parents based on their GCA will be very useful for breeding for quantitative resistance to late blight. Parents JX 90, JF 4841, CP 3356, CP 1358, CP 3290, JN 1197 and CP 3125 were found to have good GCA for quantitative resistance for late blight and the best six crosses for late bilght resistance based on mean performance involved parents with good combining ability only.     

Somaclonal variation in explant-derived potato clones over three tuber generations

Summary Tuber characteristics of potato clones (Solanum tuberosum L.) regenerated from cultured leaf, rachis or stem pieces of cv. Desiree were assessed in field trails over three years. In the first two seasons, when the crop was grown for seed potato production, tuber numbers and the tuber yields were normally distributed and the population means of the clones were about the same as those of the controls. When 78 of the clones were grown as a ware crop in a replicated field trial there were statistically significant differences between most clones and the controls in total yield (generally lower) and ware yield (generally higher), brought about by alterations of the size distributions of tubers. In all years the population of clones had fewer scab (Streptomyces scabies) lesions than the controls. Clones with consistent differences in flesh colour and eye depth were also observed. Two out of 78 clones, in 1984, had significantly higher dry matter content. No stable decrease in susceptibility to the cyst nematode Globodera pallida was observed amongst 167 clones tested. These potato plants regenerated from explants clearly show that somaclonal variation can include agronomically relevant changes.     

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.     

Resistance to potato late blight (Phytophthora infestans (Mont.) de Bary) in Solanum nigrum,S. villosum and their sexual hybrids with S. tuberosum and S. demissum

Summary Solanum nigrum and S. villosum, and their sexual hybrids with S. tuberosum and S. demissum respectively, were inoculated with a complex race of Phytophthora infestans. No visible reaction was seen on S. villosum and one genotype of S. nigrum. Another genotype of S. nigrum and the hybrids showed a hypersensitive response on most inoculated leaves. In one experiment, some sporulation was observed on detached leaves of a hybrid derived from S. nigrum. Microscopical examination of infections in S. nigrum and in a hybrid from S. nigrum, showed that penetration of epidermal cells and subsequent intercellular growth of the pathogen into the spongy mesophyll occurred, but without the formation of haustoria, and that invaded and neighbouring cells became necrotic. Callose appositions were found in epidermis and mesophyll cells of all inoculated genotypes, and also in epidermal cells of the unrelated nonhost species Brassica campestris.     

Search for sources of early blight (Alternaria solani) field resistance not associated with vegetative late maturity in tetraploid potato germplasm

Summary A total of 934 potato cultivars and clones from different breeding programmes across the world were tested for their reaction to early blight (Alternaria solani) in Brasilia-DF, Brazil. This field trial was set up in order to identify tetraploid potato genotypes with useful levels of resistance to early blight (EB) that were not correlated with late maturity or lateness. Artificial inoculation (spread ofA. solani infected leaves over plant canopy) was employed even though EB occurs naturally at epidemic levels in the test area. Four disease assessments were made every 10 days, beginning 45 days after planting. Based on the evaluation made 65 days after planting, only 27 clones were identified that had less lesioned leaf area than the resistant standard (cv. Aracy). Area under disease progress curve (AUDPC) was also used as a criterion of resistance. Using AUDPC as a parameter, 22 genotypes were identified presenting values lower than or identical to cv. Aracy. Differences in the commercial yield were recorded in genotypes with identical response to EB, suggesting that distinct levels of tolerance to EB also exist. In spite of the strong correlation between EB resistance and late maturity, some genotypes derived from the parental materials NDD 277-2, Kufri Jyoti, CIP 377888-7, Maine-28, and the clones CFS 69.1 and I-853 were identified as having good levels of EB field resistance not linked with the negative phenotypic trait of lateness. These materials presented medium to medium-early maturity with a vegetative cycle of around 90–95 days. Some selected genotypes presented more acceptable agronomic characteristics including resistance toPhytophthora infestans, potato leaf roll luteovirus (PLRV) and potato virus Y (PVY). The EB resistance sources identified in this work can be recommended as parental materials for widening the genetic base for EB disease resistance in breeding programmes for highland tropical and subtropical areas.     

Field resistance to late blight (Phytophthora infestans (Mont.) De Bary) in potato (Solanum tuberosum L.) somaclones associated with instability and pleiotropic effects

Summary Adventitious regenerants (somaclones) of Bintje and their vegetative progeny were screened for field resistance to Phytophthora infestans as follows: the area under the disease progress curve was computed and correlated with resistance rating in Bintje and reference varieties. The resistance rating of the somaclones was determined from this relationship.Clones with stable improved field resistance in successive years' trials were detected, however, most of such clones were also maturation mutants. Variation in resistance rating in clone replicates and between years was detected in most clones.The possible basis of the field resistance and reasons for its instability are discussed.Abbreviations AUDPC area under the disease progress curve     

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.     

Transgenic potato plants resistant to viruses

Summary Traditional potato breeding is a laborious process in which, by intercrossing, valuable traits from different parental clones are combined in a progeny genotype. Depending on the availability of genes, molecular techniques can be used to add specific genes to existing cultivars that, although otherwise satisfactory, lack a lew commercially important traits. For virus resistance the gene for the coat protein of a given virus transplanted into the genome of the plant renders the plant resistant to that virus. In conferring such resistance to potato varieties it proved to be possible to preserve their intrinsic properties.     

The breeding values of potato parents for field resistance to late blight measured by whole seedlings

Summary Parents used in a potato breeding programme were intercrossed in a multiple mating scheme. The resulting progenies were assessed for resistance to late blight, using as criteria the size and frequencies of lesions on leaves, on petioles and on stems, and an overall score. Analysis of variance showed that all the statistically significant genetic variation was attributable to general combining ability (GCA) differences. The seven variates were highly correlated.     

Field resistance to late blight and potato root eelworm in group tuberosum dihaploids

Summary An examination of the feasibility of obtaining dihaploids with high levels of field resistance to important potato pathogens, showed that dihaploids may be produced which are more resistant to late blight or to potato root eelworm pathotype E than are their tetraploid parents. The significance of this is discussed in relation to the possible types of gene action involved in conferring field resistance.     

Study on the inheritance of potato tuber yield by means of harvest index components and its consequences for choice of parental material

Summary The inheritance of potato tuber yield was studied considering the seasonal course of harvest index. Harvests of several progenies and their parents were performed at intervals and fresh weight of foliage, underground stems, tubers and stolons was assessed.Tuber yield depends on the time of tuber initiation and the proportion of the assimilates allocated to the tubers, as reflected in the increase rate of harvest index. The time of tuber initiation of the progenies was associated with the time of tuber initiation of the earliest tuberizing parent. The mean increase rate of the harvest index of the progenies was in most cases equal to the mean values of parents, however in crosses combining early with late maturing parents the rate equaled the value of the maternal parent. Using these relations, differences in harvest index between progenies can be predicted with a high degree of precision by using the harvest index parameters of the parents. This estimated harvest index showed a fairly good correlation to tuber yield at mid-September and can be used in the prediction of the yield of the progeny. The use of the mid-parent value for this purpose is not suitable.     

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.     

Evaluation of seedling selection for salinity tolerance in potato (Solanum tuberosum L.)

Summary Salt-tolerance in potato (Solanum tuberosum L.) was selected on the basis of germination and survival of seed sown in trays of perlite suspended in either 75 or 150 mM NaCl. Salinity reduced the germination of seed. Genetic differences in salt-tolerance were apparent with salt reducing germination more in seed collected from cv. Cara than in that collected from cv. Maris Piper. Progeny from the seedling selection were then grown to maturity to produce tubers.The relative tolerance of the parental cultivars and of unselected and selected progeny to long-term exposure to salinity was examined in a pot experiment in which plants were irrigated with either fresh water or 50 mM NaCl solution from one week after plant emergence. In this experiment, salinity significantly reduced leaf conductance, total dry matter production and partitioning of assimilate to tubers. Salinity reduced dry matter production and assimilate partitioning to tubers to a greater extent in Cara than in M. Piper. Progeny selected for short-term salt-tolerance did not exhibit greater long-term salt-tolerance than unselected progeny, and both were more sensitive than M. Piper.These results demonstrate genetic variation in salt-tolerance in potato. However, although there was a correlation between the performance of the parent to long-term salinity and survival of progeny in the seedling selection, there was no correlation between short- and long-term salt-tolerance. This suggests that characters underlying short-term tolerance may contribute to long-term tolerance but do not of themselves confer long-term tolerance. Future progress in selecting for improved salt-tolerance depends on understanding the effects of salinity on the physiological processes underlying growth and carbon partitioning.     

Breeding behavior for tuber protein in Solanum tuberosum and Tuberosum-phureja hybrids

Summary True protein content among tubers within a plant of Solanum tuberosum cv. Oneida was found to be negatively correlated with tuber size. A similar study of S. phureja genotype 148-17 revealed no correlation. Tuber protein was determined for 250 genotypes in each of four hybrid potato populations in a factorial mating design with four potato cultivars as stylar parents mated to four groups of pollen parents (4x cultivars, 4x high protein selections, 2x S. phureja, 2x high protein S. phureja). The hybrid population derived from the 4x high protein selections was significantly higher in protein content than the other populations. High estimates of general combining ability for tuber protein content were found for both stylar and pollen parents. Poor photoperiodic adaptation to growing conditions in northern latitudes was a possible explanation for the variable protein phenotypes typical of S. phureja and the lack of transmission of the high protein character in phureja to 4x-2x hybrids. Although protein content was negatively correlated with total yield, high protein segregates with good yield potential were identified in all four populations.Scientific Journal Series Article 11, 616 of the Minnesota Agricultural Experiment Station.