Potato Breeding at the Scottish Plant Breeding Station and the Scottish Crop Research Institute: 1920–2008

Seventy-two potato cultivars have been bred at the Scottish Plant Breeding Station and the Scottish Crop Research Institute since 1920. The original genetic base contained resistance to wart disease and to viruses, but not comprehensive resistance to all strains. Introgression of resistance genes from the wild and cultivated potato species of Latin America started for late blight in 1932, for viruses in 1941 and for potato cyst nematodes in 1952. Just seven of the 219 wild tuber-bearing species recognized by Hawkes in 1990 feature in the pedigrees of our cultivars, with Solanum demissum for blight resistance in 58, S. vernei for nematode resistance in 19 and S. microdontum for Potato virus Y resistance in 15, the other four species being S. multidissectum, S. commersonii, S. maglia and S. acaule. Resistance to other fungal and bacterial diseases has been mainly due to chance rather than deliberate breeding. From 1970, selection for yield and quality included processing quality, and despite lack of commercial success, prospects remain good for cultivars resistant to sweetening during cold storage. Since 1990 prebreeding has combined desirable traits through efficient recurrent selection based on progeny testing and provided parents for the commercially funded breeding of finished cultivars. Only one cultivar is a Neotuberosum–Tuberosum hybrid, whereas 15 cultivars have the H1 gene for resistance to Globodera rostochiensis introgressed from group Andigena. Long-day Phureja cultivars are finding a market niche for their flavour attributes. Breeding strategies and methods are critically reviewed from a genetic viewpoint.     

Potato viruses in China

Potato (Solanum tuberosum L.) is globally the fourth largest staple crop after rice, wheat and maize. China is the largest potato producer worldwide, accounting for 26.3% and 22.2% of the global total area and yield, respectively. Viral diseases have been a limiting factor for sustainable production of potato in China. This article provides update and comprehensive information on the most important viruses, their effect on yield and their control in China. Meristem culture is currently the most widely used technique for production of virus-free potato plants and these plants are used as nuclear stock for the production of certified seed tubers. Although a significant increase in tuber yield has been achieved using certified seed potatoes, lack of application of a national seed potato production scheme has hindered progress. Conventional breeding using virus-resistant cultivars introduced from other countries as parents has already led to the production of Chinese cultivars. Use of virus-resistant wild Solanum species and genetic transformation techniques will provide new virus-resistant varieties in the future to further enhance control of potato viruses in China.     

Cisgenesis,a New Tool for Traditional Plant Breeding,Should be Exempted from the Regulation on Genetically Modified Organisms in a Step by Step Approach

Modern potato breeding requires over 100,000 seedlings per new variety. Main reasons are (1) the increasing number of traits that have to be combined in this tetraploid vegetatively propagated crop, and (2) an increasing number of traits (e.g., resistance to biotic stress) originates from wild species. Pre-breeding by introgression or induced translocation is an expensive way of transferring single traits (such as R-genes, coding for resistance to biotic stress) to the cultivated plant. The most important obstacle is simultaneous transfer of undesired neighbouring alien alleles as linkage drag. Stacking several genes from different wild sources is increasing this linkage drag problem tremendously. Biotechnology has enabled transformation of alien genes into the plant. Initially, transgenes were originating mainly from microorganisms, viruses or non-crossable plant species, or they were chimeric. Moreover, selection markers coding for antibiotic resistance or herbicide resistance were needed. Transgenes are a new gene source for plant breeding and, therefore, additional regulations like the EU Directive 2001/18/EC were developed. Because of a strong opposition against genetic modification of plants in Europe, the application of this Directive is strict, very expensive, hampering the introduction of genetically modified (GM) crops and the use of this technology by small and medium-sized enterprises (SMEs). Currently, GM crops are almost the exclusive domain of multinationals. Meanwhile, not only transgenes but also natural genes from the plant species itself or from crossable plant species, called cisgenes, are available and the alien selection genes can be avoided in the end product. This opens the way for cisgenic crops without alien genes. The existing EU directive for GM organisms is not designed for this new development. The cisgenes belong to the existing breeders’ gene pool. The use of this classical gene pool has been regulated already in agreements regarding breeders’ rights. We are proposing a step by step approach starting with a crop and gene specific derogation and monitoring towards a general exemption of cisgenic plants from the Directive. Two examples, i.e. development of cisgenic potato for resistance to Phytophthora infestans and cisgenic apple for resistance to Venturia inaequalis are discussed shortly for illustration of the importance of cisgenesis as a new tool for traditional plant breeding. Cisgenesis is simplifying introgression and induced translocation breeding tremendously and is highly recommended for SMEs and developing countries.     

Germplasm Release: Three Tetraploid Potato Clones with Resistance to Common Scab

Common scab caused by the soil-borne bacterium Streptomyces scabies is a serious disease for the potato industry. We have identified a strong source of resistance in the diploid wild relative Solanum chacoense. This resistance has been introgressed into tetraploid cultivated potato via unilateral sexual polyploidization. This paper describes three hybrid clones (M8, M16, M17) for use by potato breeding programs to enhance resistance to common scab. They were created by crossing a diploid clone (50% S. tuberosum, 50% S. chacoense) to tetraploid cultivars, producing hybrids with 75% cultivated and 25% wild germplasm. The clones are male and female fertile, and are adapted to temperate zone production environments.     

Screening for Chip-Processing Potato Line from Introgression of Wild Species’ Germplasms with Post-Harvest Storage and Chip Qualities

The processing of potatoes into chips is expanding in China. There is a need for new processing cultivars that are well-suited for the local ecological environment. Eleven potato varieties were bred from hybridization and backcrossing between the local cultivated potato varieties (Solanum tuberosum L.) and wild potato species. Lines from different wild species’ germplasms showed that the ability to accumulate reducing sugars was significantly different during low temperature storage (4 °C). A correlation analysis was conducted to determine the correlation coefficient among reducing sugars, acid invertase, free amino acids, chip colors and the content of acrylamide after storage at room and low temperatures. The lines 0706-116, 0737-6 and 0726-205 had low levels of reducing sugars, acrylamide content and acceptable chip colors for potato chip processing. The results indicated that the transfer of wild species’ processing traits into local cultivars by hybridization and continuous backcrossing is an effective potato breeding method and that the wild germplasm resources S. phureja and S. chacoense are suitable for improving the processing traits of local varieties.     

The Effect of Photoperiod on Tuberisation in Cultivated × Wild Potato Species Hybrids

Wild Solanum species offer a valuable source of genetic diversity for potato improvement. Most of these species are found in equatorial South and Central America and they do not tuberise under long-day photoperiods typical of those in the major potato production areas of North America, Europe and Asia. Crosses between two haploids of Solanum tuberosum Chilotanum Group as females and two wild Solanum species clones generated four hybrid families. The parents and progeny were subjected to two greenhouse trials to evaluate tuberisation at 20-, 14-, and 8-h photoperiods. Parents and offspring tuberised most readily at the 8-h photoperiod, and poorly or not at all at the 20-h photoperiod. Segregation for tuberisation in hybrids was apparent at the 14-h photoperiod and depended on the cultivated parent, but not the wild species parent. The data support previous studies and best fit a model in which tuber production under long photoperiods is controlled by two dominant genes in cultivated × wild species hybrids.     

Salt tolerance inSolanum kurzianum andS. tuberosum cvs Alpha and Russet Burbank

Summary The cultivated potatoSolanum tuberosum cvs Alpha and Russet Burbank and the wild speciesS. kurzianum were compared with respect to salt tolerance. Plants of the wild species were found to be more salt tolerant than those of the cultivated species since their growth was less impaired by the salinity, although they accumulated more sodium and less potassium ions in the shoot. Unlike the whole plant, callus derived from the wild plants was not more tolerant than that from the cultivated species. Differences in the responses to salinity between cultivated and wild plants, and between the whole plants and calli derived from them are discussed. Based on these differences and the similarity of the physiological responses to salt stress betweenS. kurzianum and the wild salt-tolerant relatives of tomato, the former is suggested as a potential source of genes for increasing the salt tolerance of potato.     

Breeding Potential of Potato Somatic Hybrids: Evaluations for Adaptability,Tuber Traits,Late Blight Resistance,Keeping Quality and Backcross (BC<Subscript>1</Subscript>) Progenies

Ten tetraploid interspecific potato somatic hybrids developed earlier through protoplast fusion between the dihaploid Solanum tuberosum L. ‘C-13’ and the diploid wild species Solanum pinnatisectum Dun. were used in this study. Somatic hybrids and standard control cultivars were evaluated for adaptability, tuber traits, late blight resistance and keeping quality attributes (dormancy, sprouting, weight loss and appearance after 75 days of storage) during two successive winter crop seasons (short-days) in the sub-tropical plains of India, where nearly 90% of India’s potatoes are grown. Somatic hybrids showed medium to good plant vigour and late to very late plant maturity. Though none of the somatic hybrids produced equal or higher tuber yield than the control cultivars, except clone P7, all other somatic hybrids produced significantly higher marketable and total tuber yield after a 90-day growth than the parent C-13. Most of the somatic hybrids possessed higher tuber dry matter concentration, resistance to late blight and better keeping quality attributes than the control cultivars. Notably, the tubers of somatic hybrids showed a tendency for colour change from white to purple on exposure to sunlight. Breeding potential of somatic hybrids was further assessed by crossing them with commonly grown local potato cultivars for the development of backcross (BC₁) progenies. In the hybridization programme, berries were formed while somatic hybrids were used either as male or female parents with common cultivars. Our results revealed that the use of bulk pollen of somatic hybrids not only resulted in higher pollination success but also helped to achieve higher numbers of true potato seeds (TPS) per berry. Despite the formation of berries, TPS was absent in 22 crosses using somatic hybrids as a female or male parent which may be due to ovule or embryo abortions. Segregating BC₁ generations were evaluated under the same sub-tropical conditions for further exploitation in potato breeding. Based on our study, the somatic hybrids P4, P8 and P10 can be utilized for the introgression of important characters such as high tuber dry matter concontration, resistance to late blight and excellent keeping quality attributes into the cultivated potato via conventional breeding methods for cultivar development in the sub-tropical plains of India.     

Seed protein variation within accessions of wild and cultivated potato species and inbredSolanum chacoense

Summary Soluble proteins in seed potato were analyzed by electrophoresis (PAGE). The Index of Variation (IV) was calculated in order to estimate variation within each accession of wild and cultivated potato species andS. chacoense selfed lines. Seeds of 34 accessions representing 14 species from six taxonomic series were analyzed. The IV ofS. chacoense selfed lines decreased from 47.7 in the S₂ to 12.9 in the S₇ generation. The average IV for the self-incompatible diploid species was 41.2, whereas that of the selfcompatible diploid species,S. polyadenium, was 21.3. TetraploidS. tuberosum ssp.andigena andS. sucrense had an average IV of 62.2, which is much higher than that observed in any other diploid species. The naturally selfing speciesS. acaule (4x), and its colchicine-doubled derivative (8x), showed very low variation (IV=7.5). The IV appears to reflect heterozygosity and is affected by the breeding mode and the genetic constitution of diploid parents for derived polyploids.     

UseoftheEndospermBalanceNumber(EBN)inPotatoGeneticsandBreeding

在马铃薯初花期喷施植物生长调节剂多效唑,试验结果表明:多效唑显著抑制植株的高度,单株结薯数下降,大薯率上升,当每667m2喷施多效唑75g,浓度在2.5g.L-1时是本试验中提高产量的最佳浓度,干物质和淀粉均有明显的提高。说明喷施多效唑具有控上促下的作用,从而有利于提高产量和品质。     

多效唑对晋薯7号马铃薯的调控作用

在马铃薯初花期喷施植物生长调节剂多效唑,试验结果表明:多效唑显著抑制植株的高度,单株结薯数下降,大薯率上升,当每667m2喷施多效唑75g,浓度在2.5g.L-1时是本试验中提高产量的最佳浓度,干物质和淀粉均有明显的提高。说明喷施多效唑具有控上促下的作用,从而有利于提高产量和品质。     

An analysis of the hybrid nature ofsolanum xedinense Berth, using molecular, cytological and crossability studies

Summary Solanum xedinense Berth., a pentaploid (2n=5x=60) interspecific hybrid between the cultivated potato and the wild speciesSolanum demissum Lindl., represents evidence of genetic exchange between wild and cultivated species. It has been found in Central Mexico at altitudes between 2000 and 3500 meters, growing in and along the edges of cultivated fields, roadside thickets, forest margins and along irrigation ditches. This study examined the hybrid nature of S.xedinense using molecular, cytological and crossability data. RAPD and AFLP markers were used to supportS. xedinense as a hybrid between the cultivated potato andS. demissum. Meiotic analysis and crossability studies were used to examine the potential ofS. xedinense to form offspring. Several accessions ofS. tuberosum L. subsp.tuberosum and subsp.andigena Hawkes,S. demissum and an accession ofS. xedinense were used in the study. Artificial hybrids betweenS. tuberosum subsp.tuberosum andS. demissum were included for comparison. Molecular analysis positioned the natural and artificial hybrids as an intermediate group between the parental populations, supporting their relationship with the parental populations. Meiosis was abnormal in both natural and artificial hybrids, and reduced crossability was noted forS. xedinense. Although gene flow appears to happen between wild and cultivated potato, the potential for the natural hybrids to become established may depend mostly on their ability to clonally propagate themselves.     

Introgression of cold (4 C) chipping from 2x (2 endosperm balance number) potato species into 4x(4EBN) cultivated potato using sexual polyploidization

Cold-chipping cultivars could reduce microbial spoilage and chemical use due to cold storage of chipping potatoes. Sexual polyploidization breeding schemes that introgress cold chipping from diploid potato species may develop improved cultivars at an accelerated rate. The research objectives were (1) to determine the breeding behavior of cold chipping introgressed from 2× 2 Endosperm Balance Number (EBN) potato species into 4×(4EBN) progeny using sexual polyploidization, and (2) to determine if differences exist between 2n gametes and n tetrasomic gametes for transmission of cold chipping. Experimental families of 4×(4EBN) progeny were synthesized using sexual polyploidization (2× × 4× and 4× × 2×) and 4× × 4× matings using cold-chippingS. phureja and cultivated potato -wild species (C -W) hybrid parents. Control families from 4× × 4× matings using as parents current industry leading germplasm were also made. Progeny were field grown and evaluated for chip color (1 light-10 dark, ≤4.0 is industry acceptable) after 3 and 6 months storage at 4 C. A larger percentage of progeny from experimental families had acceptable chip color compared to control families, likely due to introgression of cold-chipping alleles from diploid potato species. Out of experimental families, the best chip color means, highest phenotypic variance, and greatest percentage of acceptable chipping progeny resulted in 2× × 4× and 4× × 2× matings. The ability of 2n gametes to transmit dominant alleles to high frequencies of 4× progeny may contribute to the superiority of 4× -2× crosses. Breeding schemes that use sexual polyploidization in conjunction with early generation selection should rapidly develop 4× cold-chipping germplasm.     

Evaluation of tuber-bearingSolanum species for nitrogen use efficiency and biomass partitioning

Modern potato cultivars (Solanum tuberosum L.) require high rates of fertilizer nitrogen (N). This practice is costly and can pose a serious threat to surface and groundwater. Previous evaluation of wild potato germplasm demonstrated the existence of species capable of producing high total biomass under low N conditions, with the ability to make maximum use of added N. Therefore, a two-year field experiment was conducted in 1994 and 1995 to investigate the response of selected wild potato accessions and their hybrids with the haploid USW551 (USW) to low and high N environments. The haploid USW and cultivars Russet Burbank, Red Norland, and Russet Norkotah were also included in the study. Uniform propagules and seedlings from the variousSolanum species were transplanted to a Hubbard loamy sand (Udic Haploboroll) at Becker, Minn. and were subjected to two N treatments: 0 and 225 kg N ha^-1. At harvest, total dry biomass of wild and hybrid potato germplasm was equal to or higher than that of the cultivars. However, cultivar biomass partitioning was 1% to roots, 15% to shoots, 0% to fruits, and 84% to tubers, whereas wild potato species partitioned 18% to roots plus nontuberized stolons, 52% to shoots, 23% to fruits, and only 7% to tubers. Hybrids were intermediate, allocating 9% of their biomass to roots plus nontuberized stolons, 39% to shoots, 14% to fruits, and 38% to tubers. Nitrogen use efficiencies for many of the species and crosses were comparable to that for Russet Burbank and greater than those for Red Norland and Russet Norkotah. Of the wild species tested,S. chacoense accessions had the highest biomass accumulation and N uptake efficiencies and may be the best source of germplasm for improving NUE in a potato breeding program.     

Germplasm enhancement with wild tuber-bearing species: introgression of PVY resistance and high dry matter content fromSolanum berthaultii, S. gourlayi, S. tarijense andS. vernei

Summary The narrow genetic base of the cultivated potato is a severe constraint for potato breeding. Wild tuber-bearing species constitute an important source of genes for resistance to diseases and pests as well as abiotic factors. Seven interspecific hybrid populations were generated from crosses between diploidTuberosum clones and four wild species:Solanum berthaultii, S. gourlayi, S. tarijense andS. vernei, and evaluated in a field experiment in Burgos (Spain) as a way to broaden the genetic base of the cultivated potato. Good tuberization ability and great variability for yield within the different families were detected. Hybridization with the wild species resulted in high levels of PVY resistance inS. berthaultii andS. vernei hybrids and a large increase of dry matter content in all populations except theS. berthaultii hybrids.     

Tuber and starch quality of wild and cultivated potato species and cultivars

Summary Between 1993 and 1998 205 different potato cultivars and 1220 accessions/genotypes of wild and cultivated potato species from the IPK Genebank Gatersleben were evaluated. Parameters interesting for starch isolation and especially for the use of starch were determined. Altogether, there was a higher variability in wild potato species than in cultivated potatoes for all characteristics investigated: dry matter content, starch content, protein content, amylose content and mean particle diameter of starch granules.     

Susceptibility of potato varieties and advanced breeding lines (Solanum tuberosum L.) toPhytophthora infestans (Mont.) de Bary in greenhouse screenings

Late blight (Phytophthora infestons (Mont.) de Bary) has re-emerged as an important pathogen of the cultivated potato (Solanum tuberosum subsp.tubero-sum L.) in North America. The purpose of this study was to evaluate the relative susceptibility of potato germplasm in the greenhouse in order to initiate a breeding program for resistance to the US-8/A2 mating type which is the more aggressive and prevalent strain of late blight. Whole plants of 147 cultivars and breeding lines were evaluated. Percent plant area infection was visually assessed. Seven days after inoculation, infection ranged from 0 to 100% and the overall mean was greater than 50%. Two-thirds of the cultivars and breeding lines tested were very susceptible to the US-8 genotype. The highest resistance was identified in the somatic hybrids betweenS. tuberosum andS. bulbocastanum and their backcross derivatives. Pike and Snowden were less susceptible than the other North American cultivars. Zarevo was most resistant among the European cultivars. Seven of the advanced breeding lines were equivalent to Zarevo in infection levels. The host plant resistance identified among the material tested in this study can be used by breeding programs to develop improved cultivars with resistance to US-8 genotypes of late blight.     

Wild annual Helianthus anomalus and H. deserticola for improving oil content and quality in sunflower

Within the past decade, the desire for alternative sources of fuels, chemicals, and other industrial materials has received increased attention. Sunflower (Helianthus annuus L.) oil has the potential to be improved for nutritional and industrial purposes through selection and breeding. The narrow genetic base of cultivated sunflower has been broadened by the infusion of genes from wild species, resulting in a continuous improvement in agronomic traits. The genus Helianthus is comprised of 51 species and 19 subspecies with 14 annual and 37 perennial species. Interest in using wild species in breeding programs has increased, but concerns about the introduction of low oil concentration and quality from the wild species persist. Two annual desert species, Helianthus anomalus Blake and H. deserticola Heiser, are excellent candidates for increasing oil concentration and enhancing quality based on their adaptation to desert environments. The objective of this study was to collect achenes of H. anomalus and H. deserticola from the desert southwest USA and assess their potential for improving oil concentration and quality in cultivated sunflower. The sunflower collection took place from 16 to 23 September 2000 and covered a distance of 4100 km in three states: Utah, Arizona, and Nevada. The only H. deserticola population collected had an average oil concentration of 330 g/kg, whereas the two populations of H. anomalus had an oil concentration of 430 and 460 g/kg, the highest concentration recorded in any wild sunflower species. The linoleic fatty acid concentration in the oil of H. anomalus populations was uncharacteristically high for a desert environment, approaching 700 g/kg. A linoleic acid concentration of 540 g/kg in H. deserticola was more typical for a desert environment. H. anomalus has the largest achenes and the highest oil concentration of any wild sunflower species, and the same chromosome number (n = 17) as cultivated sunflower. These features will facilitate the introduction of genes from this wild annual progenitor into cultivated sunflower. The lower saturated fatty acid profile in this species is also a desirable trait offering the potential to reduce saturated fatty acids in cultivated sunflower. Further research will be needed to determine the inheritance of the fatty acids and oil concentration. Other agronomic traits will need to be maintained during the introgression of these traits into cultivated sunflower oil.     

T-type chloroplast DNA inSolarium tuberosum L. ssp.tuberosum was conferred from some populations ofS. tarijense Hawkes

The highly heterozygous and tetraploid nature of potato (Solanum tuberosum L. ssp.tuberosum) has hampered discovery of its wild ancestral species. Chloroplast DNA is a very reliable indicator to trace maternal ancestry of crops. Most of the common potato (grown worldwide) has unique, T-type chloroplast DNA derived from Chilean cultivated potato (both areS. tuberosum ssp.tuberosum). Analyzing seven different chloroplast DNA markers, I found all the T-type accessions of cultivated potatoes shared the same chloroplast DNA haplotype only with some accessions ofS. tarijense Hawkes and its derived hybrids. Thus, I conclude that some populations ofS. tarijense acted as the maternal ancestor of potato.     

Molecular marker-assisted selection for potato breeding

The use of molecular markers in potato breeding offers new opportunities for the selection of genotypes. To date, many markers linked to useful traits have been found. Indeed, the potato molecular map is saturated with more than 350 markers uniformly distributed on 12 chromosomes. More than 25 single dominant genes have been localized on the potato map, most of them being pest-resistance genes, together with some quantitative trait loci (QTL) controlling yield and tuber quality traits. Molecular markers linked to these genes are now available for rapid, efficient assisted selection (positive assisted selection). Moreover, in order to transfer useful genes, interspecific crosses between wild and cultivated genotypes are often performed, and in this case the use of species-specific molecular markers would allow the wild genomic content to be reduced in few backcross generations (negative assisted selection). The following review discusses the progress achieved in potato breeding through molecular marker-assisted selection.