Testing Taxonomic Predictivity of Foliar and Tuber Resistance to Phytophthora infestans in Wild Relatives of Potato.

Potato late blight, caused by the oomycete phytopathogen Phytophthora infestans, is a devastating disease found in potato-growing regions worldwide. Long-term management strategies to control late blight include the incorporation of host resistance to predominant strains. However, due to rapid genetic changes within pathogen populations, rapid and recurring identification and integration of novel host resistance traits is necessary. Wild relatives of potato offer a rich source of desirable traits, including late blight resistance, but screening methods can be time intensive. We tested the ability of taxonomy, ploidy, crossing group, breeding system, and geography to predict the presence of foliar and tuber late blight resistance in wild Solanum spp. Significant variation for resistance to both tuber and foliar late blight was found within and among species but there was no discernable predictive power based on taxonomic series, clade, ploidy, breeding system, elevation, or geographic location. We observed a moderate but significant correlation between tuber and foliar resistance within species. Although previously uncharacterized sources of both foliar and tuber resistance were identified, our study does not support an assumption that taxonomic or geographic data can be used to predict sources of late blight resistance in wild Solanum spp.

A test of taxonomic and biogeographic predictivity: resistance to Potato virus Y in wild relatives of the cultivated potato.

A major justification for taxonomic research is its assumed ability to predict the presence of traits in a group for which the trait has been observed in a representative subset of the group. Similarly, populations in similar environments are expected to be more alike than populations in divergent environments. Consequently, it is logical to assume that taxonomic relationships and biogeographical data have the power to predict the distribution of disease resistance phenotypes among plant species. The objective of this study was to test predictivity in a group of widely distributed wild potato species, based on hypotheses that closely related organisms (taxonomy) or organisms from similar environments (biogeography) share resistance to a simply inherited trait (Potato virus Y [PVY]). We found that wild potato species with an endosperm balance number (EBN) of 1 (a measure of cross compatibility) shared resistances to PVY more than species with different EBN values. However, a large amount of variation was found for resistance to PVY among and within species. We also found that populations from low elevations were more resistant than those from high elevations. Because PVY is vectored by aphids, we speculate that the distribution of aphids may determine the level of selection pressure for PVY resistance.

A test of taxonomic predictivity: resistance to the Colorado potato beetle in wild relatives of cultivated potato.

Wild relatives of potato offer a tremendous germplasm resource for breeders. Because the germplasm base of potato is so broad and diverse, we have undertaken a series of studies to determine whether we can predict the distribution of valuable genes in wild Solanum species based on taxonomic or biogeographic data. This is the third study in the series. Resistance to defoliation by Colorado potato beetle, Leptinotarsa decemlineata Say, larvae was evaluated in 156 accessions of 41 wild Solanum species. The highest frequencies of resistant accessions were found in diploid species with an endosperm balance number of 1. In contrast to previous studies on resistance to foliar fungal pathogens, there was little variability in defoliation scores among plants within an accession and among accessions within a species, at least for the most resistant species. There was no strong association of Colorado potato beetle resistance in wild potato species to biogeographic data. Resistance was confirmed in species previously characterized by high levels of glycoalkaloids or dense glandular trichomes. However, we have identified additional species with resistance to the Colorado potato beetle. Mechanisms of resistance are being studied in these species and attempts will be made to introgress them into the cultivated potato.

A test of taxonomic predictivity: resistance to early blight in wild relatives of cultivated potato.

Host plant resistance offers an attractive method of control for early blight (caused by the foliar fungus Alternaria solani), a widespread disease that appears annually in potato crops worldwide. We tested the assumed ability of taxonomy to predict the presence of early blight resistance genes in wild Solanum species for which resistance was observed in related species. We also tested associations to ploidy, crossing group, breeding system, and geography. As in a prior study of Sclerotinia sclerotiorum (white mold) resistance, tremendous variation for resistance to early blight was found to occur within and among species. There was no discernable relationship between the distribution of resistant phenotypes and taxonomic series (based on an intuitive interpretation of morphological data), clade (based on a cladistic analysis of plastid DNA data), ploidy, breeding system, geographic distance, or climate parameters. Species and individual accessions with high proportions of early blight resistant plants were identified, but high levels of inter- and intra-accession variability were observed. Consequently, the designation of species or accessions as resistant or susceptible must take this variation into account. This study calls into question the assumption that taxonomic or geographic data can be used to predict sources of early blight resistance in wild Solanum species.

Genetic analysis of the cultivated potato Solanum tuberosum L. Phureja Group using RAPDs and nuclear SSRs.

The Solanum tuberosum L. Phureja Group consists of potato landraces widely grown in the Andes from western Venezuela to central Bolivia, and forms an important breeding stock due to their excellent culinary properties and other traits for developing modern varieties. They have been distinguished by short-day adaptation, diploid ploidy (2n = 2x = 24), and lack of tuber dormancy. This nuclear simple sequence repeat (nSSR or microsatellite) study complements a prior random amplified polymorphic DNA (RAPD) study to explore the use of these markers to form a core collection of cultivar groups of potatoes. Like this prior RAPD study, we analyzed 128 accessions of the Phureja Group using nuclear microsatellites (nSSR). Twenty-six of the 128 accessions were invariant for 22 nSSR markers assayed. The nSSR data uncovered 25 unexpected triploid and tetraploid accessions. Chromosome counts of the 102 accessions confirmed these nSSR results and highlighted seven more triploids or tetraploids. Thus, these nSSR markers (except 1) are good indicators of ploidy for diploid potatoes in 92% of the cases. The nSSR and RAPD results: (1) were highly discordant for the remaining 70 accessions that were diploid and variable in nSSR, (2) show the utility of nSSRs to effectively uncover many ploidy variants in cultivated potato, (3) support the use of a cultivar-group (rather than a species) classification of cultivated potato, (4) fail to support a relationship between genetic distance and geographic distance, (5) question the use of any single type of molecular marker to construct core collections.

Late blight resistance linkages in a novel cross of the wild potato species Solanum paucissectum (series Piurana).

The cultivated potato, Solanum tuberosum, is affected by a variety of diseases with late blight, caused by Phytophthora infestans, being the most severe. Wild potato species have proven to be a continuing source of resistance, sometimes of an extreme type, to this disease. The present study constructs the first late blight linkage map of a member of series Piurana, S. paucissectum, a tuber-bearing relative of potato, using probes for conserved sequences from potato and tomato. Eight probes mapped to unexpected linkage groups, but syntenic differences with prior maps of potato were not supported by any blocks of rearranged chromosome segments. All 12 linkage groups were resolved and significant associations with late blight resistance were found on chromosomes 10, 11 and 12. A major quantitative trait locus (QTL) on chromosome 11 accounts for more than 25% of the phenotypic variance measured in a field trial. Crossing of S. paucissectum with cultivated potato resulted in very few seeds indicating partial reproductive barriers. Differential reactions of accessions of this potential donor species with simple and complex isolates of P. infestans suggest that it carries major resistance genes that are not those previously described from the Mexican species, S. demissum. However, the additivity of the QTL effects argues for the quantitative nature of resistance in this cross.

Nuclear and chloroplast DNA reassessment of the origin of Indian potato varieties and its implications for the origin of the early European potato.

The modern cultivated potato was first recorded in Europe in 1562, but its area(s) of exportation has long been in dispute. Two competing hypotheses have proposed an "Andean" area (somewhere from upland Venezuela to northern Argentina) or a lowland south central "Chilean" area. Potato landraces from these two areas can be distinguished, although sometimes with difficulty, by (1) cytoplasmic sterility factors, (2) morphological traits, (3) daylength adaptation, (4) microsatellite markers, and (5) co-evolved chloroplast (cp) and mitochondria (mt) DNA. The Chilean introduction hypothesis originally was proposed because of similarities among Chilean landraces and modern "European" cultivars with respect to traits 2 and 3. Alternatively, the Andean introduction hypothesis suggests that (1) traits 2 and 3 of European potato evolved rapidly, in parallel, from Andean landraces to a Chilean type through selection following import to Europe, and (2) the worldwide late blight epidemics beginning in 1845 in the United Kingdom displaced most existing European cultivars and the potato was subsequently improved by importations of Chilean landraces. We reassess these two competing hypotheses with nuclear microsatellite and cpDNA analyses of (1) 32 Indian cultivars, some of which are thought to preserve putatively remnant populations of Andean landraces, (2) 12 Andean landraces, and (3) five Chilean landraces. Our microsatellite results cluster all Indian cultivars, including putatively remnant Andean landrace populations, with the Chilean landraces, and none with the "old Andigenum" landraces. Some of these Indian landraces, however, lack the cpDNA typical of Chilean landraces and advanced cultivars, indicating they likely are hybrids of Andean landraces with Chilean clones or more advanced cultivars. These results lead us to reexamine the hypothesis that early introductions of potato to Europe were solely from the Andes.

Selection of highly informative and user-friendly microsatellites (SSRs) for genotyping of cultivated potato.

Characterization of nearly 1,000 cultivated potato accessions with simple sequence repeats (SSRs; also referred to as microsatellites) has allowed the identification of a reference set of SSR markers for accurate and efficient genotyping. In addition, 31 SSRs are reported here for a potato genetic map, including new map locations for 24 of them. A first criterion for this proposed reference set was ubiquity of the SSRs in the eight landrace cultivar groups of the potato, Solanum tuberosum. All SSRs tested in the present study displayed the same allele phenotypes and allele size range in the diverse germplasm set as in the advanced potato cultivar germplasm in which they were originally discovered. Thirteen of 13 SSR products from all cultivar groups are shown to cross-hybridize with the corresponding SSR product of the source cultivar to ascertain sequence homology. Other important SSR selection criteria are quality of amplification products, locus complexity, polymorphic index content, and well-dispersed location on a potato genetic map. Screening of 156 SSRs allowed the identification of a highly informative and user-friendly set comprising 18 SSR markers for use in characterization of potato genetic resources. In addition, we have identified true- and pseudo-multiplexing SSRs for even greater efficiency.

Granule-bound starch synthase (GBSSI) gene phylogeny of wild tomatoes (Solanum L. section Lycopersicon [Mill.] Wettst. subsection Lycopersicon).

Eight wild tomato species are native to western South America and one to the Galapagos Islands. Different classifications of tomatoes have been based on morphological or biological criteria. Our primary goal was to examine the phylogenetic relationships of all nine wild tomato species and closely related outgroups, with a concentration on the most widespread and variable tomato species Solanum peruvianum, using DNA sequences of the structural gene granule-bound starch synthase (GBSSI, or waxy). Results show some concordance with previous morphology-based classifications and new relationships. The ingroup comprised a basal polytomy composed of the self-incompatible green-fruited species S. chilense and the central to southern Peruvian populations of S. peruvianum, S. habrochaites, and S. pennellii. A derived clade contains the northern Peruvian populations of S. peruvianum (also self-incompatible, green-fruited), S. chmielewskii, and S. neorickii (self-compatible, green-fruited), and the self-compatible and red- to orange- to yellow-fruited species S. cheesmaniae, S. lycopersicum, and S. pimpinellifolium. Outgroup relationships are largely concordant with prior chloroplast DNA restriction site phylogenies, support S. juglandifolium and S. ochranthum as the closest outgroup to tomatoes with S. lycopersicoides and S. sitiens as basal to these, and support allogamy, self-incompatibility, and green fruits as primitive in the tomato clade.

Geographic distribution of wild potato species.

The geographic distribution of wild potatoes (Solanaceae sect. Petota) was analyzed using a database of 6073 georeferenced observations. Wild potatoes occur in 16 countries, but 88% of the observations are from Argentina, Bolivia, Mexico, and Peru. Most species are rare and narrowly endemic: for 77 species the largest distance between two observations of the same species is <100 km. Peru has the highest number of species (93), followed by Bolivia (39). A grid of 50 × 50 km cells and a circular neighborhood with a radius of 50 km to assign points to grid cells was used to map species richness. High species richness occurs in northern Argentina, central Bolivia, central Ecuador, central Mexico, and south and north-central Peru. The highest number of species in a grid cell (22) occurs in southern Peru. To include all species at least once, 59 grid cells need to be selected (out of 1317 cells with observations). Wild potatoes occur between 38° N and 41° S, with more species in the southern hemisphere. Species richness is highest between 8° and 20° S and around 20° N. Wild potatoes typically occur between 2000 and 4000 m altitude.

Comparison of four molecular markers in measuring relationships among the wild potato relatives Solanum section Etuberosum (subgenus Potatoe).

We evaluated chloroplast DNA (cpDNA), isozymes, single to low-copy nuclear DNA (RFLPs), and random amplified polymorphic DNAs (RAPDs) in terms of concordance for genetic distance of 15 accessions each of Solanum etuberosum and S. palustre, and 4 accessions of S. fernandezianum. These self-compatible, diploid (2n=24), and morphologically very similar taxa constitute all species in Solanum sect. Etuberosum, a group of non-tuber-bearing species closely related to Solanum sect. Petota (the potato and its wild relatives). Genetic distance and multidimentional scaling results show general concordance of isozymes, RFLPs and RAPDs between all three taxa; cpDNA shows S. etuberosum and S. palustre to be more similar to each other than to S. fernandezianum. Interspecific sampling variance shows a gradation of resolution from allozyme (low) to RAPD to RFLP (high); while intraspecific comparisons graded from RFLPs (low) to RAPDs (high; lack of sufficient allozyme variability within species precluded comparisons for allozymes). Experimental error was low in RFLPs and RAPDs.

RFLP analysis of the wild potato species, Solanum acaule Bitter (Solanum sect. Petota).

Intraspecific variation of a wild potato species, Solanum acaule Bitt., was analyzed by RFLPs of genomic DNA. One hundred and five accessions were selected throughout the distribution area, including all subspecies, i.e., ssp. albicans (hexaploid), ssp. punae (tetraploid), ssp. acaule (tetraploid) and ssp. aemulans (tetraploid). Twenty-seven low-copy DNA clones (probes) were Southern hybridized with EcoRI, EcoRV, HindIII, and XbaI digests of total DNA of all accessions. In total, 238 RFLPs were detected from 94 enzyme x probe combinations. Among them, 49 RFLPs were specific to ssp. albicans, suggesting that the additional third genome is distinct from its two other genomes. RFLPs between and within subspecies were analyzed by principal component analysis. DNA similarities between subspecies coincided with a former taxonomic treatment in the sense that ssp. albicans is the most distantly related to ssp. acaule and ssp. aemulans is distantly related. Subspecies acaule and ssp. punae were indistinguishable. In addition, RFLPs could be used to distinguish groups within subspecies. Subspecies aemulans, confined to Argentina, was divided into two populations, one from the province of La Rioja and the other from the province of Jujuy. In ssp. acaule, some accessions from the southernmost distribution area were clearly distinguishable, while the others varied continuously, showing a geographical cline from Peru to Argentina.