Heritability of late blight resistance in tomato conferred by Solanum pimpinellifolium accession PI 224710

Late blight (LB), caused by the oomycete Phytophthora infestans, is one of the most destructive diseases of tomato and potato worldwide. Identifying and characterizing new sources of resistance is essential given the emergence of new aggressive and fungicide‐resistant P. infestans isolates. Recently, we reported identification of several new sources of LB resistance within the tomato wild species, Solanum pimpinellifolium. In this study, we examined heritability (h²) of LB resistance conferred by the S. pimpinellifolium accession PI 224710 using a parent–offspring regression (correlation) analysis. F₂ and F₃ progeny populations, derived from crosses between PI 224710 and a LB‐susceptible tomato breeding line, were evaluated for response to LB infection. To obtain a better estimate of h², the F₃ progeny were evaluated for LB resistance in two separate replicated experiments. The h² estimates were similar in the two experiments and averaged ~0.87, suggesting that this resistance was highly heritable. Two different methods estimated involvement of one resistance locus. Breeding and mapping efforts are underway to further assess the viability of this newly reported LB resistance.     

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.     

Genetic analysis of resistance to tomato late blight in Solanum pimpinellifolium accession PI 163245

Late blight (LB), caused by Phytophthora infestans, is one of the most devastating diseases of tomato (Solanum lycopersicum) worldwide. Due to the emergence of new and aggressive P. infestans isolates, identifying new genetic resistance to LB is a priority in tomato breeding. Recently, we reported the identification of several Solanum pimpinellifolium accessions with strong LB resistance. In this study, we investigated the utility of resistant accession PI 163245 for tomato breeding by examining heritability (h²) of resistance and the response to selection for resistance. Estimates of h² based on F₂ : F₃ and F₃ : F₄ parent : offspring correlation analyses averaged 0.79 and 0.94, respectively, suggesting the heritable nature of LB resistance in PI 163245. Analysis of response to selection for resistance from F₂ to F₄ generations indicated a realized h² of 0.63, confirming the utility of this resistance in tomato breeding. Two methods of estimating the minimum number of loci involved indicated the presence of one major resistance locus. Currently, genetic mapping and breeding efforts are underway to further confirm the viability of this accession for improving tomato LB resistance.     

Mapping of a novel,major late blight resistance locus in the diploid (1EBN) Mexican Solanum pinnatisectum Dunal on chromosome VII

Late blight caused by the oomycete Phytophthora infestans (Mont.) de Bary (Pi) is the most important foliar disease of potato worldwide. An intraspecific hybrid between individuals of a resistant and a susceptible S. pinnatisectum accession was backcrossed to the susceptible parent to generate a segregating population for late blight resistance consisting of 84 plants. In detached‐leaflet assays, reaction to late blight segregated in a 1r:1s manner in BC₁ progeny indicating the presence of a single dominant resistance gene. A genetic map was constructed based on 1,583 DArT/SSR markers which were allocated to 12 linkage groups, covering 1,793.5 cM with an average marker distance of 1.1 cM. The late blight resistance locus derived from S. pinnatisectum was mapped on chromosome VII. In comparison with the previously reported resistance genes Rpi1 and Rpi2, the new target resistance locus most likely is located on the opposite arm of chromosome VII. Results of this study will serve as a basis for future fine mapping of the late blight resistance locus and the development of locus‐specific markers for marker‐assisted selection.     

The potato R10 resistance specificity to late blight is conferred by both a single dominant R gene and quantitative trait loci

The R10 late blight differential of potato, 3681ad1, exhibits good field resistance. Progeny from the cross between 3681ad1 and the susceptible cultivar ‘Katahdin’ were assessed for late blight resistance to three Phytophthora infestans isolates, using a detached leaf assay. Progeny differed in response to the three isolates. Resistance to isolates IPO‐0 and 99018 was controlled by quantitative trait loci (QTL), whereas resistance to isolate 89148‐9 was inherited as a dominant R gene, designated as R10 in this study. Statistical analysis revealed that one of the resistance QTLs to isolates IPO‐0 and 99018 is linked to the R10 gene, which maps to chromosome 11 in a region where a complex late blight resistance locus has been reported previously. A high‐resolution map of R10 was constructed using a large segregating population, and the gene was delimited to a genetic interval of 0.26 cM. The clustering of the qualitative gene R10 with resistance QTLs could explain the field resistance observed with 3681ad1.     

Mapping polygenes for tuber resistance to late blight in a diploid Solanum phureja × S. stenotomum hybrid population

Potato tuber blight is a disease caused by the oomycete Phytophthora infestans (Mont.) de Bary. Due to the significant economic impact of this disease, introgression of durable resistance into the cultivated potato is one of the top priorities of breeding programmes worldwide. Though numerous resistance loci against this devastating disease have already been mapped, most of the detected loci are contributing towards foliar resistance while specific information on tuber resistance is limited. To identify the genetic components of tuber resistance and its relationship to foliar resistance and plant maturity we have investigated the host‐pathogen interaction in a segregating diploid hybrid Solanum phureja × S. stenotomum family. Mature tubers from this mapping family were inoculated with a sporangial suspension of P. infestans (US‐8 clonal lineage) and evaluated for lesion expansion. No significant correlation was detected between late blight resistance in foliage and tubers, and between plant maturity and tuber resistance. Four chromosomal regions were significantly associated with tuber resistance to the disease. The largest effect was detected near the marker locus PSC (LOD 10.7) located on chromosome 10. This locus explained about 63% of the total phenotypic variation of the trait. The other three resistance‐related loci were mapped on chromosomes 8 (GP1282, LOD 4.4), 6 (CP18, LOD 4.0) and 2 (CP157, LOD 3.8). None of the four tuber resistance loci coincides with the foliage resistance loci detected in this same family. Tuber blight resistance quantitative trait loci (QTL) on chromosomes 2, 8 and 10 are distinct from the maturity QTLs and have an additive effect on tuber resistance. These results indicate that different genes are involved in foliar and tuber resistance to P. infestans in the present family and that some of the resistance genes might be associated with late maturity.     

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.     

Two distinct potato late blight resistance genes from <Emphasis Type="Italic">Solanum berthaultii</Emphasis> are located on chromosome 10

For breeding potato varieties resistant to late blight, identification of resistance genes to Phytophthora infestans (Rpi genes) is essential. Introduction of Rpi genes from wild Solanum species into cultivated potato is likely to be a good method to achieve durable resistance to P. infestans. In this study, we identified two Rpi genes (Rpi-ber1 and Rpi-ber2) derived from two different accessions of Solanum berthaultii. These two genes are closely linked on the long arm of chromosome 10. There are similarities between the predicted genetic locations of the previously identified Rpi-ber and Rpi-ber1, which given the common origin of these genes, may indicate that they are the same. However, the genetic positions of Rpi-ber1 and Rpi-ber2 are different. Rpi-ber1 is positioned between CT214 and TG63, whereas Rpi-ber2 is located below both of these two markers. In addition, the sequences of four linked markers to both R genes showed different polymorphisms indicating the two Rpi genes could be transmitted from different haplotypes (chromosomes).     

On the influence of extra R-genes on the resistance of the potato to the corresponding P-races1) ofPhytophthora infestans

Summary A study was made on the possible influence of extraR ₃-genes on the resistance to the physiologic racep ₃ (=(3)) ofPhythophthora infestans. The data allowed of the conclusion that such an influence could be small only, not exceeding small differences in the average degree of field resistance existing between the groups of simplex, duplex and triplex plants.     

An evaluation of Solanum verrucosum Schlechtd. For its possible use in potato breeding

Summary Solanum verrucosum is a diploid (2n=24) wild potato species from Mexico. It is self-compatible, but in greenhouse artificial selfing is mostly necessary for seed setting. The species has a good male and female fertility and possesses resistance to different races and isolates of Phytophthora infestans. It is easily crossable with different species from South America. The good crossability, the genomic constitution and resistance to diseases favour its use in potato breeding at the diploid level.     

Inheritance of resistance to the peach root‐knot nematode (Meloidogyne floridensis) in interspecific crosses between peach (Prunus persica) and its wild relative (Prunus kansuensis)

The peach root‐knot nematode, Meloidogyne floridensis (MF), infects majority of available nematode‐resistant peach rootstocks which are mostly derived from peach (Prunus persica) and Chinese wild peach (P. davidiana). Interspecific hybridization of peach with its wild relative, Kansu peach (P. kansuensis), offers potential for broadening the resistance spectrum in standard peach rootstocks. We investigated the inheritance of resistance to MF in segregating populations of peach (‘Okinawa’ or ‘Flordaguard’) × P. kansuensis. A total of 379 individuals from 13 F₂ and BC₁F₁ families were challenged with a pathogenic MF isolate “MFGnv14” and were classified as resistant (R) or susceptible (S) based on root galling intensity. Segregation analyses in F₂ progeny revealed the involvement of a major locus with a dominant or recessive allele determining resistance in progeny segregating 3R:1S and 1R:3S, respectively. Testcrosses with a homozygous‐susceptible peach genotype (‘Flordaguard’ or ‘UFSharp’) confirmed P. kansuensis as a source of new resistance and the heterozygous allelic status of P. kansuensis at the locus conferring resistance to MF. We propose a single‐locus dominant/recessive model for the inheritance of resistance.     

Response of accessions within tomato wild species,Solanum pimpinellifolium to late blight

Late blight (LB), caused by the oomycete Phytophthora infestans, is one of the most devastating diseases of the cultivated tomato (Solanum lycopersicum) worldwide. Most commercial cultivars of tomato are susceptible to LB. Previously, three major LB resistance genes (Ph‐1, Ph‐2, Ph‐3) were identified and incorporated into a few commercial cultivars of tomato. Reduced effectiveness and potential breakdown of the resistance genes has necessitated identification, characterization and utilization of new sources of resistance. We evaluated the response of 67 accessions of the wild tomato species, S. pimpinellifolium to LB, under multiple field and greenhouse (GH) conditions and compared them with six control genotypes. Sixteen accessions were identified with strong LB resistance in both field and GH experiments. However, 12 accessions exhibited resistance similar to a control line which was homozygous for Ph‐2 + Ph‐3. Genotyping accessions with molecular markers for Ph‐2 and Ph‐3 were not conclusive, indicating that resistance in these accessions could be due to these or other resistance genes. Strong correlations were observed between field and GH disease response and between foliar and stem infection.     

Multiple resistance to diseases and pests in potatoes

Summary The potato has more characters of economic importance that need to be considered by the breeder than any other temperate crop. In Europe these include resistance to at least twelve major diseases and pests. Highest priority has been given to resistance to late blight (Phytophthora infestans), virus diseases (particularly those caused by potato leafroll virus and potato virus Y) and potato cyst nematode (Globodera rostochiensis andG. pallida). Useful sources of resistance are available and early generation screening techniques have been developed to allow positive selection for multiple resistance and the breeding value of clones used as parents to be determined. Progress in restriction fragment length polymorphism technology should result in more efficient selection in the future.     

Identification of somatic hybrids of dihaploid Solanum tuberosum lines and S. brevidens by species specific RAPD patterns and assessment of disease resistance of the hybrids

Summary Symmetric somatic hybrids were produced by electrofusion of protoplasts of two dihaploid tuber-bearing potato (Solanum tuberosum L.) lines and Solanum brevidens Phil., a diploid non-tuber-bearing wild potato species. A total of 985 plants was obtained. Verification of nuclear hybridity of putative hybrids was based on additive RAPD patterns, general morphological characteristics and chromosome counts. 53 (90%) calli regenerated into plants which were identified as somatic hybrids. Most of the hybrids were aneuploids at the tetraploid (4×) or hexaploid (6×) level. The 20 hybrids tested expressed a high level of resistance to potato virus Y (PVY ^N ) characteristic of the S. brevidens parent. Resistance to late blight (Phytophthora infestans (Mont.) de Bary) varied between hybrids, but was on average better than that of the fusion parents. Resistance of hybrids to bacterial stem rot (Erwinia carotovora subsp. atroseptica (van Hall) Dye) was not superior to that of commercial potato cultivars.     

Inheritance and stability of the virus-resistant gene in the progeny of transgenic sweet potato

Viral diseases of sweet potato are very prevalent and often seriously damaging to the plants. In particular, the severe strain of the sweet potato feathery mottle virus (SPFMV‐S) causes ‘obizyo‐sohi’ disease in Japan. In order to confer viral resistance against SPFMV using current biotechnology, a transgenic sweet potato has been produced, introducing hygromycin‐resistant (hpt) and SPFMV‐S coat protein (CP) genes, which have shown a significant resistance to SPFMV‐S. In the breeding programme, it is important to confirm that the viral resistance conferred in T₀ plants can be inherited by their progeny. In the present study, progeny were obtained from crosses between the transgenic T₀ and a non‐transgenic variety of sweet potato. The results showed that the CP gene was inherited by the next generation and that the stability of viral resistance was also confirmed. Thus, this production system for the virus‐resistant transgenic sweet potato is useful in practical breeding.     

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.     

Resistance to Phytophthora infestans in Potato Genotypes Originating from Wild Species

The reaction to P. infestans in the foliage and tubers of potato was evaluated in resistant genotypes and in their progeny. The genotypes originated from wild species known to carry genes for specific resistance. Evidence was obtained that three types of resistance to P. infestans were presented in the genotypes: (1) Resistance due to major, dominant genes was predominant. The presence of these genes was expressed in both foliage and tubers and did not influence maturity. Some of the genes were R genes, providing specific resistance, but for others this could not be determined as they were effective against the fungus inoculum with the highest virulence available: 1.2.3.4.5.7.10.11. (2) Resistance was present, associated with late maturity. (3) Tuber resistance was present, expressed after inoculation of whole tubers but not after inoculation of tuber slices. Testing of the progeny was essential for the determination of the type of resistance present in the genotypes evaluated.     

Differentiation of the high molecular weight glutenin subunit D t x 2.1 of Aegilops tauschii indicated by partial sequences of its encoding gene and SSR markers

The devastating late blight pathogen Phytophthora infestans infects foliage as well as tubers of potato. To date, resistance breeding has often focused on foliage blight resistance, but tuber blight resistance is becoming more and more important in cultivated potatoes. In this study, a reliable tuber assay for resistance assessment was developed and foliage and tuber blight resistance (R) was compared in four mapping populations. In the RH4X-103 population, tuber blight resistance inherited independently from foliage blight resistance. Three specific R genes against P. infestans were segregating. The Rpi-abpt and R3a genes function as foliage-specific R genes, whereas the R1 gene acts on both foliage and tuber. In the segregating populations SHRH and RH94-076, tuber and foliage blight resistance correlated significantly, which suggests that resistance in foliage and tuber is conferred by the same gene (could be R3b) and quantitative trait loci (QTL), respectively. In the CE population neither tuber nor foliage resistance was observed.     

Some notes on the relations between field resistance to Phytophthora infestans in leaves and tubers and ripening time in Solanum tuberosum subsp. Tuberosum

A survey is made of the correlations between three characters, viz. field resistance of the foliage and of the tubers to Phytophthora infestans and ripening time. The relation between resistance of the foliage and ripening time is so close, that to all appearances it seems to be difficult to combine earliness with a high degree of resistance. This conclusion holds true only for the subspecies tuberosum of Solanum tuberosum and the breeder must draw some optimism from the possibility that this close correlation can be broken by the use of wild and cultivated South American and Mexican material.There is not such a close correlation between ripening time and tuber resistance. From some preliminary experiments the impression is gained that the skin of the tuber is an important barrier for the penetration of the fungus into the tuber in many cases. However, the defence reactions of the flesh are also instrumental in determining of degree of field resistance.     

Late blight resistance in a diploid full-sib potato family

Late blight, caused by Phytophthora infestans (Mont.) de Bary, is the most destructive disease of potato worldwide. As this pathogen can rapidly overcome major race‐specific resistance genes, identifying the basis for enhanced quantitative resistance has become a crucial element for implementing advanced breeding strategies. A population of 230 full‐sib progeny derived from a cross between two diploid hybrid Solanum phureja × S. stenotomum clones was evaluated for foliage resistance against late blight in replicated trials at multiple locations in Pennsylvania between 1999 and 2002. In field experiments, plants were evaluated visually for per cent defoliation, and area under the disease progress curve (AUDPC) was determined. The two parents and three control cultivars (‘Atlantic’, ‘Kennebec’ and ‘Katahdin’) were included in all trials. In all three experiments, the presence of a significant number of clones exhibiting transgressive segregation were observed. There were significant differences among environments as well as among clones, and the clone × environment interaction was also significant. Stability analysis revealed that 37 clones made a significant contribution to the overall environment × clone interaction. Broad‐sense heritability for resistance, measured as AUDPC, was estimated as 0.67. The overall results indicate the presence in this potato family of a high level of field resistance against late blight. This segregating diploid family appears to be a good candidate for quantitative trait loci mapping to identify and characterize the genetic components of partial late blight resistance.