Resistance to late blight in potato. A putative gene that suppresses R genes and is elicited by specific isolates

Eleven progenies, ten bi-parental and one selfed, of tetraploid potato were assessed for phenotypic segregation ratios of resistance to susceptible individuals against two isolates of P. infestans, one avirulent to all 11 known potato R genes (avirulent pathotype) and one virulent to eight R genes (virulent pathotype). Assuming Mendelian inheritance of independent R genes, the numbers of individuals segregating for resistance to both races could not be explained sufficiently for all progenies. However, the assumed presence of dominant, specific suppressors of the R gene function that were elicited by specific isolates of the fungus and that segregated in the host independently from R genes explained all frequencies of resistant and susceptible individuals in each of the progenies studied. This revised version was published online in July 2006 with corrections to the Cover Date.     

马铃薯资源晚疫病抗性的全基因组关联分析

广谱抗性基因的挖掘是马铃薯高抗晚疫病品种选育的基础。本研究以288份国际马铃薯中心筛选的晚疫病抗性群体为试验材料,经过连续2年田间调查,计算AUDPC和sAUDPC值,评估群体晚疫病抗性;利用SLAF-seq方法进行群体简化基因组测序,通过对晚疫病抗性表型数据的全基因组关联分析,挖掘晚疫病抗性相关的遗传位点和候选基因,为晚疫病抗性品种选育和抗病机理研究提供一定的理论和材料基础。结果表明,晚疫病抗性在288份材料间存在着广泛的遗传差异;基于5种分析模型,共鉴定到82个与晚疫病抗性显著关联的位点;在关联区间关联到54个已知或可能与晚疫病抗性相关的基因。其中,23个基因为抗性基因,包括晚疫病抗性基因R1同源基因、Sw-5同源基因(R8)和Rpi-vnt1以及编码多效性耐药蛋白基因;5个基因编码MAPK蛋白和WRKY转录因子;1个基因参与茉莉酸途径;3个基因与水杨酸途径相关;6个基因是病程相关的基因;3个基因参与苯基丙酸类合成途径;其他与晚疫病抗性相关的基因,如HMGR基因(2个)、细胞色素P450(21个)。     

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.     

Linkage and quantitative trait locus mapping of foliage late blight resistance in the wild species Solanum vernei

The global cultivation of potato (Solanum tuberosum) is threatened by epidemics caused by new variants of the late blight pathogen, Phytophthora infestans. New sources of durable late blight resistance are urgently needed and these may be found in wild Solanum species. The diploid wild species, S. vernei, has not previously been subjected to mapping of quantitative trait loci (QTLs) for late blight resistance. Two populations designated HGIHJS and HGG, originating from a cross between a clone of S. vernei and two different S. tuberosum clones were evaluated in field trials for late blight infestation. The relative area under the disease progress curve (RAUDPC) was estimated and used for QTL mapping. A linkage map of S. vernei, comprising 11 linkage groups, nine of which could be assigned to chromosomes, was constructed. Results indicated that the resistance in S. vernei was quantitatively inherited. Significant QTLs for late blight resistance were identified on chromosomes VIII (HGG), VI and IX (HGIHJS). In addition, potential QTLs were detected on chromosomes VII (HGIHJS) and IX (HGG). A putative and a significant QTL for tuber yield were found on chromosomes VI and VII in HGG, but no linkage between yield and resistance was indicated. The QTL for late blight resistance, which mapped to chromosome IX, could be useful for late blight resistance breeding as it was located close to the microsatellite marker STM1051 in both populations.     

Vertical resistance to late blight in wild potato species from Argentina

The possible presence of vertical resistance to late blight conferred by R genes in Argentine wild Solanum species, which presumably have not evolved under the pressure of the fungus, was investigated. Solanum microdontum, S. commersonii and S. chacoense clones were tested in the greenhouse and with detached leaves in the laboratory after inoculation with a complex race and a non-virulent race of Phytophthora infestans. Two progeny tests were carried out to assess the performance of contrasting parents in regards to their resistance. A varying frequency of incompatible clones was detected among the species, with S. microdontum having the higher proportion of clones yielding incompatible reactions both in the laboratory and in the greenhouse. S. chacoense and S. commersonii showed a small but still considerable frequency of incompatible clones. The progeny tests confirmed the presence of R genes in a S. commersonii cross and their absence in a S. chacoense cross. The unexpected presence of R genes, its evolutionary significance and the consequences on the use of these species in breeding is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Molecular markers for late blight resistance breeding of potato: an update

Late blight is the most devastating disease of the potato crop that can be effectively managed by growing resistant cultivars. Introgression of resistance (R) genes/quantitative trait loci (QTLs) from the Solanum germplasm into common potato is one of the plausible approaches to breed resistant cultivars. Although the conventional method of breeding will continue to play a primary role in potato improvement, molecular marker technology is becoming one of its integral components. To achieve rapid success, from the past to recent years, several R genes/QTLs that originated from wild/cultivated Solanum species were mapped on the potato genome and a few genes were cloned using molecular approaches. As a result, molecular markers closely linked to resistance genes or QTLs offer a quicker potato breeding option through marker‐assisted selection (MAS). However, limited progress has been achieved so far through MAS in potato breeding. In near future, new resistance genes/QTLs are expected to be discovered from wild Solanum gene pools and linked molecular markers would be available for MAS. This article presents an update on the development of molecular markers linked to late blight resistance genes or QTLs by utilization of Solanum species for MAS in potato.     

Molecular breeding for resistance to Phytophthora infestans (Mont.) de Bary in potato (Solanum tuberosum L.): a perspective of cisgenesis

Late blight caused by Phytophthora infestans is one of the most devastating diseases in potato cultivation and is mostly controlled by the application of chemicals. However, introduction of combinations of resistance ( R ) genes conferring broad-spectrum resistance from wild Solanum species into cultivated potatoes is considered the most practical and promising approach to achieve durable resistance. This can be realized via classical breeding or genetic modification (GM). Because classical breeding is very time-consuming and is often hampered by linkage drag, a GM approach seems logic in this heterozygous and vegetatively propagated crop. During the last decades, many R genes have been identified in several wild Solanum species. Some have been cloned and more will follow. When these genes are derived from species crossable with cultivated potato (so-called cisgenes), application in resistance breeding using a GM approach is similar to an introgression breeding approach, in that the exploited genes are indigenous to the crop. Pending deregulation or derogation of cisgenesis, the use of cisgenic R genes would be an ideal strategy to accomplish durable resistance in potato.     

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.     

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.     

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.     

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.     

Early generation selection for potato tuber quality in progenies of late blight resistant parents

Developing disease resistant cultivars is one of the major objectives for a potato (Solanum tuberosum L.) breeding program, but many resistant clones have not achieved commercial acceptance because of late maturity and non-marketable tuber characteristics. Selection for tuber quality should have greater emphasis inbreeding disease resistant cultivars. The objectives of this study were 1) to evaluate the ability of late blight (Phytophthora infestans (Mont.) de Bary) resistant parents to transmit chip-processing (tuber appearance, specificgravity, and chip-color) or tablestock (tuber appearance) quality to the offspring and 2) to compare selecting for tuber quality in single-hill versus eight-hill generations. We made crosses among eight unadapted potato cultivars (B0718-3,Bertita, Bzura, Greta, Libertas, Stobrawa, Tollocan, and Zarevo) with reported late blight resistance with adapted susceptible cultivars/breeding clones to generate 95populations (4,750 seedlings). Approximately 10% of the progeny from each cross were selected from single-hill plots based on tuber appearance, number, shape, and internal defects. These selected clones (408) were evaluated for tuber appearance, specific gravity, and chip-color. The same evaluations in the following year were made on tuber samples from eight-hill plots. Libertas and Tollocan were the best parents for transmitting chip-color; B0718-3, Zarevo, and Tollocan for transmitting tuber appearance; and Bzura, Libertas, and Zarevo for transmitting high specific gravity to the highest percentage of the offspring. Overall, 50% and 56% of the clones based on single- and eight-hill generation, respectively, were considered to possess chip-processing quality; over 90% of the clones had acceptable tablestock quality. A total of 71% of the clones possessing acceptable chip-processing and 95% of the clones possessing acceptable tablestock quality selected in both generations were identified in single-hill plots. The evaluation of tuber quality characteristics in single-hill generation not only permitted the identification of clones with acceptable chip-processing and tablestock, but also increased the amount of clonal information for the following generation of selection. In crosses between late blight resistant and susceptible clones, selection for tuber quality traits can be initiated in single-hill generation using a moderate selection intensity and precede late blight testing. This revised version was published online in August 2006 with corrections to the Cover Date.     

中国东北部与河北省致病疫霉SSR基因型的组成与分析

为了解中国东北部致病疫霉基因型分布及变化规律。利用2对SSR引物Pi4B和Pi4G对2008-2010年期间采自于河北、黑龙江、辽宁和吉林的373株致病疫霉的SSR基因型进行分析,4个省份共鉴定出16种SSR基因型,分别为A-03、A-09、B-01、D-03、F-01、F-04、F-05、F-06、F-07、F-08、G-02、G-03、I-01、J-01、K-01和L-01,其中主导基因型为F-01,占所测菌株的67.83%,其次为F-06基因型(14.75%);河北、黑龙江、辽宁和吉林4个省份SSR基因型的种类分别为11种、8种、4种和3种;16种基因型中,A-09、F-07、F-08、I-01、J-01、K-01和L-01等7种基因型在东北及河北地区属首次报道;2008-2010年期间,河北省和黑龙江省致病疫霉SSR基因型数量均有所增加,辽宁和吉林基因型种类变化不明显,但总体上东北地区和河北省致病疫霉优势基因型较单一,这为上述地区制定马铃薯晚疫病综合防治策略、尤其是品种的合理布局提供了一定的理论依据。     

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.     

Combination of resistance to potato late blight in foliage and tubers by intraspecific dihaploid protoplast fusion

Five dihaploid Solanum tuberosum genotypes, encompassing different levels of polygenically inherited resistance to potato late blight disease (Phytophthora infestans (Mont.) de Bary) in foliage and tubers, were used in four intraspecific protoplast fusion combinations. Vigorous growing putative hybrid calli were selected four weeks after electrofusion. Intact plants were regenerated from 6% of the selected calli. Verification of hybridity was accomplished by use of RAPD analysis which revealed that 53% of the regenerated plants were true somatic hybrids. The score of true somatic hybrids in the different fusion combinations ranged within 21% – 100%. The hybrid plants were analysed for resistance to foliage blight in a field trial and assessed for tuber blight resistance by use of a laboratory test. Resistance to late blight in foliage and tubers varied between the hybrids. Very high levels of resistance to both foliage and tuber blight were obtained in some hybrids. However, loss of resistance in some hybrids as compared to the parental plants were also observed. Possible reasons for the phenotypic disappearance of resistance to either foliage or tuber blight are suggested. This revised version was published online in July 2006 with corrections to the Cover Date.     

Heritability of resistance to foliar late blight in a diploid hybrid potato population of Solanum phureja×Solanum stenotomum

The emergence of new races of Phytophthora infestans has necessitated the search for additional sources of potato germplasm with resistance to late blight. This study examined 281 clones, derived from 72 families of a diploid random-mated hybrid population of Solanum phureja×Solanum stenotomum. The clones were evaluated in a replicated field trial for 2 years with the control cultivar ‘Atlantic’ in Pennsylvania, USA. The P. infestans US-8 A2 mating type culture was used to inoculate spreader rows of susceptible S. tuberosum cv.‘Russet Burbank’. Percent defoliation caused by the late blight fungus was estimated visually in each plot three times near the end of the growing season. Area under the disease progress curve (AUDPC) was estimated and showed that late blight was more severe in 1997 than in 1996. Genetic differences among clones and significant, but small, clone–environment interaction were detected for AUDPC. Broad-sense and narrow-sense heritability estimates, over years, were 0.79 ± 0.05 (P = 0.05) and 0.78 ± 0.29, respectively. Seventy-five percent of the diploid clones had a significantly lower mean AUDPC than Atlantic. These results support the idea that this diploid population is worthy of use in breeding for late blight resistance in tetraploid potato cultivars.     

Genetics of resistance to P3 isolate of Phytophthora blight in pigeonpea

Seedlings from six crosses between susceptible × resistant and two between susceptible parents were screened against P₃ isolate of Phytophthora drechsleri f. sp. cajani in the glasshouses. Disease reaction was scored on a rating scale of 1–5. Resistance was found to be controlled by one dominant gene and some minor genes and is affected by ontogeny. The resistance gene appeared to be different from the one reported earlier (Pd₁) and is designated here as Pd₃. This revised version was published online in July 2006 with corrections to the Cover Date.     

Production of somatic hybrids between S. tuberosum L. and late blight resistant Mexican wild potato species

Interspecific somatic hybrids between dihaploid breeding clones of potato, S. tuberosum and two accessions of wild Mexican species S. pinnatisectum and the hybrid line S. pinnatisectum × S. bulbocastanum were regenerated following electrofusion of mesophyll protoplasts to combine important agricultural traits of S. tuberosum and a high level of late blight resistance from selected wild accessions. In two fusion combinations 239 calli were regenerated; 162 from 195 calli analysed were identified as hybrids by means of isozyme analysis of peroxidases and, for some hybrid clones, by RAPD analysis. Depending on the fusion combination, 47–89 percent of the somatic hybrids had the expected ploidy level and 7–16 percent were mixoploids. Somatic hybrids were phenotypically intermediate as compared to their parents and some of them were able to be backcrossed sexually with potato. Fertility and crossability depended on combination and ploidy level of the somatic hybrids. In tests with detached leaves the wild partner clones had a high late blight resistance score of 8,6 and 8,9; the susceptible tuberosum-partners of 2,8 and 3,5, respectively. Nearly 25 percent of somatic hybrids had a resistance level of 6 or higher in the first year of assessment. The average resistance value of most somatic hybrids was lower than the average parental level. The reasons for variation in resistance values are discussed in connection with the practical application of fusion hybrids. This revised version was published online in July 2006 with corrections to the Cover Date.