Effects of Different Soil Treatments on the Development of <Emphasis Type="Italic">Spongospora subterranea</Emphasis> f. sp. <Emphasis Type="Italic">subterranea</Emphasis> in Potato Roots and Tubers in the Greenhouse

Powdery scab caused by Spongospora subterranea f. sp. subterranea (Sss) causes extensive losses in potato production systems globally. Two pot experiments were established in the greenhouse in summer 2013 and winter 2014 to evaluate the effectiveness of different soil chemicals, fumigant, amendments and biological control agents (BCAs) against Sss in the rhizospheric soil, potato roots and tubers. The study used visual assessment methods to assess the effect of treatments on root galling and zoosporangia production, and qPCR to measure Sss concentration in the soil and in the potato roots and tubers. All six soil treatments, namely metam sodium, fluazinam, ZincMax, calcium cyanamide, Biocult and a combination of Bacillus subtilis and Trichoderma asperellum recorded significantly (P < 0.05) lower numbers of zoosporangia in the roots compared to the untreated control. The same effect was observed on the concentration of Sss DNA in the roots at tuber initiation. A more diverse picture was obtained when root gall scores at tuber initiation and Sss DNA in the rhizospheric soil at tuber initiation and harvesting were compared. Significant differences (P < 0.05) were also noted in disease severity, disease incidence, and tuber yield between metam sodium, fluazinam, ZincMax, calcium cyanamide and the untreated control. Calcium cyanamide gave the highest tuber yield. The study demonstrated the potential of soil treatments such as metam sodium, fluazinam, ZincMax and calcium cyanamide in managing Sss in potatoes by reducing the pathogen both in the rhizospheric soil and the roots of the potato plant.     

Chloropicrin Soil Fumigation Reduces <Emphasis Type="Italic">Spongospora subterranea</Emphasis> Soil Inoculum Levels but Does Not Control Powdery Scab Disease on Roots and Tubers of Potato

The effect of chloropicrin fumigation on the soil populations of Spongospora subterranea and the development of powdery scab, formation of root galls and tuber yield was investigated in seven field trials conducted in Minnesota and North Dakota. Sixteen potato cultivars, with different levels of susceptibility to disease on roots and tubers, were planted in plots treated with chloropicrin at rates ranging from zero to 201.8 kg a.i. ha^?1. The amount of S. subterranea DNA in soil was determined using qPCR. Bioassays were conducted to further assess the effect of chloropicrin fumigation on root colonization by S. subterranea in two potato cultivars with contrasting disease susceptibility. In the field, chloropicrin applied at rates between 70.1 to 201.8 kg a.i. ha^?1 significantly decreased S. subterranea initial inoculum in soil but increased the amount of disease observed on roots and tubers of susceptible cultivars. The effect of increasing disease was confirmed in controlled conditions experiments. Although the amount of S. subterranea DNA in roots of bioassay plants increased with increasing chloropicrin rates, it remained similar among potato cultivars. Chloropicrin fumigation significantly increased tuber yield which in cultivars such as Shepody and Umatilla Russet were associated with the amount root galls (r = 0.30; P < 0.03). Results of these studies contradict earlier reports on the use of chloropicrin fumigation for the control of powdery scab. Factors other than inoculum level, such as environmental conditions that affect inoculum efficiency and host susceptibility, may be significant contributors to the development of powdery scab and root gall formation.     

Yield Response of Potato (<Emphasis Type="Italic">Solanum tuberosum L</Emphasis>.) Genotypes to Late Blight Caused by <Emphasis Type="Italic">Phytophthora infestans</Emphasis> in Uganda

Potato (Solanum tuberosum L.) is a major food and cash crop, mainly grown by small-scale farmers in the highland regions of Uganda. Potato late blight is one of the major diseases limiting production with potential yield losses over 70%, making host resistance a strong element in integrated disease management. This study was carried out to screen and select high yielding potato genotypes with resistance to late blight in Uganda. Forty-eight genotypes, including advanced clones from the population B3C2 of the International Potato Centre, commercial and farmers’ varieties, were evaluated under two environments for two seasons. Trials were laid out in an 8?×?6 alpha lattice design with three replications. Genotypes showed significant differences in yield and resistance to blight. A higher disease severity was observed in Karengyere (56%). The average RAUDPC (= 100 max) across locations indicated that genotypes 395,077.12 and 392,657.8, with disease severity of 12% and 14%, respectively, were the most resistant. Genotypes Victoria (53%) and NKRN59.124 (48%) were the most susceptible. Mean tuber yield under late blight infection was19.8 t ha^?1. The best yielding genotype across sites was 395,112.32 (35.6 t ha^?1) while 394,905.8 (10.3 t ha^?1), yielded the lowest. The mean marketable tuber weight was 8.9 kg with genotypes 395,112.32 and 395,109.34 having the highest marketable weight of 16.5 kg and 15.6 kg respectively. Correlations between yield and yield related parameters were positive (p ≤?0.001), while those between RAUDPC were negative. The following genotypes, 395,112.32, 391,919.3, 393,220.54. 393,077.54, 396,038.107. 392,657.8, Kinigi, 395,014.17, NKRN59.58, NKRK19.17 and 395,011.2, were identified as promising parents for a late blight resistance breeding program. These exhibited high to medium resistance to late blight disease and high yields.     

Maturity-Adjusted Resistance of Potato (<Emphasis Type="Italic">Solanum tuberosum</Emphasis> L.) Cultivars to Verticillium Wilt Caused by <Emphasis Type="Italic">Verticillium dahliae</Emphasis>

Verticillium wilt is a fungal disease of potato caused by two species of Verticillium, V. dahliae and V. albo atrum. The pathogen infects the vascular tissue of potato plants through roots, interfering with the transport of water and nutrition, and reducing both the yield and quality of tubers. We have evaluated the reaction of 283 potato clones (274 cultivars and nine breeding selections) to inoculation with V. dahliae under greenhouse conditions. A significant linear correlation (r = 0.4, p < 0.0001) was detected between plant maturity and partial resistance to the pathogen, with late maturing clones being generally more resistant. Maturity-adjusted resistance, that takes into consideration both plant maturity and resistance, was calculated from residuals of the linear regression between the two traits. Even after adjusting for maturity, the difference in the resistance of clones was still highly significant, indicating that a substantial part of resistance cannot be explained by the effect of maturity. The highest maturity-adjusted resistance was found in the cv. Navajo, while the most susceptible clone was the cv. Pungo. We hope that the present abundance of data about the resistance and maturity of 283 clones will help potato breeders to develop cultivars with improved resistance to V. dahliae.     

Characterization of <Emphasis Type="Italic">Phytophthora infestans</Emphasis> Isolates from Jersey,Channel Islands

Potato production on the island of Jersey, in the English Channel, is dominated by Jersey Royal, a selection of the early cultivar International Kidney. Jersey Royal is very susceptible to Phytophthora infestans, the cause of potato late blight, and Jersey’s climate is frequently conducive to infection. During 2004–2006, isolates of P. infestans were obtained from Jersey Royal plant samples from 41 different sites (24 crops, one infected tuber, 16 volunteers) and from five outdoor tomato crops and characterised by mating type, mtDNA haplotype, Gpi and Pep allozyme genotype, metalaxyl resistance and RG57 fingerprint. A subset of 17 isolates was additionally characterised by simple sequence repeat (SSR). All but one isolate from potato belonged to a single A1 multilocus genotype or a variant. The five isolates from tomato represented three distinct genotypes, none of which was found on potato and included one which was A2 mating type. The populations of P. infestans on potato and tomato in Jersey appear distinct, with that on potato being highly clonal.     

An <Emphasis Type="Italic">In Vitro</Emphasis> Assay Method for Resistance to Bacterial Wilt (<Emphasis Type="Italic">Ralstonia solanacearum</Emphasis>) in Potato

To develop an in vitro assay method for bacterial wilt resistance in potato, resistant and susceptible standard genotypes were grown in vitro, and different conditions of inoculation with Ralstonia solanacearum phylotype I/biovar 4 were examined. The optimal condition was the inoculation of 6–8 leaf stage plants with a bacterial concentration of 10² CFU ml^?1 and an incubation temperature of 28 °C. Evaluation of stem wilting was more reliable than that of leaf wilting. Using this method, nine genotypes with different resistance levels in the field were evaluated. Lower disease indices were obtained for genotypes with high resistance levels in the field, suggesting that this assay is useful for evaluating bacterial wilt resistance in a controlled environment.     

Optimization of Hairy Root Induction in <Emphasis Type="Italic">Solanum tuberosum</Emphasis>

An efficient protocol for hairy root induction in Solanum tuberosum was established using Agrobacterium rhizogenes. Transformation of Desiree was more efficient than with Shepody or Purple Majesty. Transformation efficiency of Desiree tuber discs (97% transformation efficiency) was higher in comparison to root (67% transformation efficiency), stem (61% transformation efficiency) and leaf (14% transformation efficiency) explants. Bacterial density, culture parameters, explant type, and genotype influenced the transformation efficiency, as well as the growth of hairy roots. Hairy root transformation efficiency in explants treated with acetosyringone was enhanced from 72 to 89% in Purple Majesty tubers and 56 to 80% in roots during co-cultivation. The growth rate of hairy roots from tuber discs was about 5 times greater than that of hairy roots from leaves. Tuber hairy roots were highly resistant to kanamycin in the absence of NPT II. This study describes the efficient generation of hairy roots in different potato tissues and cultivars that can provide a fast method to generate transformed tissue or be used for production of secondary metabolites.     

Impact of Different US Genotypes of <Emphasis Type="Italic">Phytophthora infestans</Emphasis> on Potato Seed Tuber Rot and Plant Emergence in a Range of Cultivars and Advanced Breeding Lines

Seed pieces of different potato cultivars and advanced breeding lines (ABLs) from north central US breeding programmes were inoculated with different genotypes of Phytophthora infestans (US-1, US-1.7, US-8, US-11 and US-14). The effect of these genotypes of P. infestans on seed piece rot severity after re-storage was assessed using an image analysis technique. P. infestans genotypes demonstrated variable ability to cause seed piece rot and to reduce plant emergence measured as final plant stand (%) and the relative area under the plant emergence curve (RAUEPC). The US-8 genotype of P. infestans was the most aggressive genotype, as indicated by tuber rot severity across all cultivars/ABLs tested, followed by US-14 in both years. The US-1, US-1.7 and US-11 genotypes were the least aggressive, causing only moderate seed piece rotting across cultivars/ABLs tested. Similar trends were observed in two field experiments, where the US-8 and US-14 genotypes delayed or reduced emergence. Values of final plant stand (%) and RAUEPC demonstrated that the cultivars/ABLs Atlantic, MSJ453-4Y and Torridon were the least susceptible across all P. infestans genotypes. In both experiments cv. Pike was the most susceptible. Other cultivars/ABLs demonstrated variable responses to different genotypes of P. infestans. No symptoms of P. infestans were observed on emerged plants up to 60 days after planting. The variability of susceptibility of tubers to different genotypes of P. infestans has implications for plant breeding efforts in that the major emphasis in the past has been to breed for foliar resistance, with limited emphasis on the reaction of the tuber. Results from this study suggest that highly aggressive genotypes of P. infestans such as US-8 may lead to severe tuber rotting and deterioration of tubers before emergence, but despite this observation the US-8 genotype is still predominant in North America.     

Amplification of the <Emphasis Type="Italic">Phytophthora infestans</Emphasis> RG57 loci Facilitates <Emphasis Type="Italic">in planta</Emphasis> T-RFLP Identification of Late Blight Genotypes

Late blight, caused by the oomycete Phytophthora infestans (Mont.) de Bary, is a devastating disease in potato and tomato and causes yield and quality losses worldwide. The disease first emerged in central America and has since spread in North America including the United States and Canada. Several new genotypes of P. infestans have recently emerged, including US-22, US-23 and US-24. Due to significant economic and environmental impacts, there has been an increasing interest in the rapid identification of P. infestans genotypes. In addition to providing details regarding the various phenotypic characteristics such as fungicide resistance, host preference, and pathogenicity associated with various P. infestans genotypes, information related to pathogen movement and potential recombination may also be determined from the genetic analyses. Restriction fragment length polymorphism (RFLP) analysis with the RG57 loci is one of the most reliable procedures used to genotype P. infestans. However, the RFLP procedure requires propagation and isolation of the pathogen and relatively large amounts of DNA. Isolation of the late blight pathogen is sometimes impossible due to the poor condition of the infected tissues or the presence of fungicide residues. In this study, we describe a procedure to identify P. infestans at the molecular level in planta using terminal restriction fragment length polymorphism (T-RFLP) of the RG57 loci. This T-RFLP assay is sufficiently sensitive to detect and differentiate P. infestans genotypes directly in planta without propagation and isolation of the pathogen, to facilitate the timely implementation of best management practices.     

Identification and Quantification of Steroidal Alkaloids from Wild Tuber-Bearing <Emphasis Type="Italic">Solanum</Emphasis> Species by HPLC and LC-ESI-MS

Wild Solanum species are characterized by several types of glycoalkaloids (GAs), which are usually not found in commercial potato cultivars. These alkaloids serve as defence compounds against herbivores and microorganisms, especially fungi. In this study, the GA composition of tuber and leaf material of 17 wild Solanum species was analysed qualitatively by liquid chromatography-electrospray ionization-mass spectrometry (LC-ESI-MS) and quantitatively by high-performance liquid chromatography. The GAs identified were α-solanine, α-chaconine, β-chaconine, solasonine, solamargine, demissine, dehydrodemissine, α-tomatine, dehydrotomatine, commersonine, dehydrocommersonine, leptine I and leptine II. Most species accumulated the common potato GAs α-solanine and α-chaconine in tubers and solasonine and solamargine additionally in leaves. In some species, such as S. acaule ssp. acaule, S. demissum and S. polyadenium, substantial amounts of unusual alkaloids were detected. By using LC-ESI-MS, we detected several minor alkaloids such as dehydrogenated forms of α-tomatine, demissine and commersonine for the first time. Total GA content, expressed as the sum of the four main alkaloids α-solanine, α-chaconine, solamargine and solasonine differed from species to species. In general, GA contents in leaf tissue were higher and GA patterns were more complex than those of tubers.     

Rooting Characteristics of <Emphasis Type="Italic">Solanum chacoense</Emphasis> and <Emphasis Type="Italic">Solanum tuberosum in Vitro</Emphasis>

The precise level of environmental control in vitro may aid in identifying genetically superior plant germplasm for rooting characteristics (RC) linked to increased foraging for plant nitrogen (N). The objectives of this research were to determine the phenotypic variation in root morphological responses of 49 Solanum chacoense (chc), 30 Solanum tuberosum Group Phureja – Solanum tuberosum Group Stenotomum (phu-stn), and three Solanum tuberosum (tbr) genotypes to 1.0 and 0.5 N rate in vitro for 28 d, and identify genotypes with superior RC. The 0.5 N significantly increased density of root length, surface area, and tips. All RC were significantly greater in chc than in either phu-stn or tbr. Based upon clustering on root length, surface area, and volume, the cluster with the greatest rooting values consisted of eight chc genotypes that may be utilized to initiate a breeding program to improve RC in potato.     

Impact of Agronomic Strategies (Seed Tuber Pre-sprouting,Cultivar Choice) to Control Late Blight (<Emphasis Type="Italic">Phytophthora infestans</Emphasis>) on Tuber Growth and Yield in Organic Potato (<Emphasis Type="Italic">Solanum tuberosum</Emphasis> L.) Crops

In order to optimize potato crop management in organic farming systems, knowledge of crop growth processes in relation to limitations and reductions by nitrogen (N) and disease is necessary. This paper examines the effect of different prevention measures (seed tuber pre-sprouting, choice of cultivars: resistance to Phytophthora infestans; earliness of tuber initiation) against disease-related constraints on yields, depending on the N-mediated growing conditions of organic potato crops. Under conditions of a relatively high N supply, accelerating the early development and tuber initiation of potato crops by seed tuber pre-sprouting (yield increase of c. 18–23%) or the selective choice of cultivars with an earlier tuber initiation (yield increase of c. 0–21%) are the most effective strategies in combating late blight. They bring forward the crop development c. 7–10 days, escaping the negative impact of the fungus on tuber growth. Under conditions of a relatively low N supply, these strategies do not affect final tuber yield. The reason for the differences of the effect depending on N supply of the crops is, that the lower the N supply, the shorter the period of time over which tuber growth takes place, independent of whether P. infestans becomes severe or not. Tuber growth of organic crops low in N has mostly ceased by the time late blight becomes potentially important for limiting yield, with the consequence that preventative measures against P. infestans are meaningless. Surprisingly, the level of cultivar haulm resistance to late blight did not affect tuber yields in years with an early and strong late blight development. Probably, the positive effect of a longer resistance to the fungus (c. 1 week) was counterbalanced by a later tuber initiation (also c. 1 week). In years with a moderate late blight incidence, less susceptible cultivars were able to produce higher tuber yields on fields with a lower N availability (c. 17–20%), probably due to higher N use efficiency and a better match of N mineralization and N uptake. Under conditions of a relatively high N supply, the tested cultivars with a higher haulm resistance to late blight were not able to improve tuber yields.     

Resistance of Potato Tubers to a Highly Aggressive Isolate of Phytophthora infestans in Relation to Tuber Age

The goal of these studies was to evaluate the influence of tuber age on the expression of resistance to Phytophthora infestans in five selected potato genotypes (three cultivars and two clones) varying in tuber resistance and maturity type, and also to select the best time for testing potato tubers for resistance to blight. Resistance was evaluated by inoculating whole tubers at 13 different times as they progressively aged during growth and storage. Tuber age was expressed as weeks after planting. Tubers were rated from week 13 (late July) until week 43 (late February) for tubers planted on 27–28 April, over three consecutive seasons (2001–2003). Analysis of variance performed on tuber resistance data showed significant effects of genotype, year and tuber age. Significant contributions of genotype × year, genotype × tuber age, year × tuber age and year × genotype × tuber age interactions were also detected. A slight increase in tuber resistance with tuber age was observed for cvs Bzura, Sokół and Irga, while the opposite trend was observed for the clones 97-A-63 and DG 92-227. Changes in tuber resistance with age of tested genotypes were not related to their maturity type. The period of most stable expression of tuber resistance was observed when tubers were tested between the 16th and 28th week after planting.     

Predictive Markers for Cold-Induced Sweetening Resistance in Cold Stored Potatoes (<Emphasis Type="Italic">Solanum tuberosum</Emphasis> L.)

An approach has been developed to screen a large number of potato clones for cold induced sweetening (CIS) resistance in breeding programs. Two key enzymes responsible for reducing sugar accumulation during cold storage were identified. Clones with the A-II isozymes of UDP-glucose pyrophosphorylase coupled with low activity of vacuolar acid invertase enzyme had increased resistance to CIS by forming less suc, which is subsequently hydrolyzed to the undesirable reducing sugars, glc and fru. Six named cultivars and 192 genetically diverse clones from various breeding programs in USA were analyzed over two years for the two key enzymes and sugar concentration in cold stored tubers. The predictability for CIS resistance during cold storage was 94% both years. Clones classified as class A accumulated low concentration of reducing sugar glc during cold storage. It is suggested that these two predictor enzymes can be used for screening parents and selections in potato breeding program.     

Relationship Between Tuber Storage Proteins and Tuber Powdery Scab Resistance in Potato

The potato genotypes (Solanum tuberosum L.) with russet tuber skin are generally resistant to powdery scab (Spongospora subterranea f.sp. subterranea or Sss). Lipoxygenase (LOX; EC 1.13.11.12) and patatin are two key storage proteins that are known to offer resistance to several diseases and insects. The objective of this study was to find out the relationship of these proteins in stored tubers with potato tuber powdery scab resistance, especially in russet skinned potatoes. An evaluation of potato germplasm with different tuber characteristics in a greenhouse environment over several years (2006–’11) suggests that russet skinned tuber genotypes (Mesa Russet, Centennial Russet and Russet Nugget) with negligible tuber disease severity index (DSI) and 100 % marketability were resistant to powdery scab. Higher physiological levels of LOX protein (on a dry weight basis) were negatively correlated with tuber DSI and positively correlated with tuber russet skin. Tuber total protein and patatin-lipase levels did not have a significant relationship with tuber powdery scab resistance. The proposed role of LOX protein in suberin- and/or non-suberin-mediated mechanisms of powdery scab resistance in russet skinned tubers are discussed here. The physiological levels of LOX protein can be considered as a useful marker for powdery scab resistance in potato breeding programs.     

Phenotypic and Genomic Modifications Associated with <Emphasis Type="Italic">Globodera pallida</Emphasis> Adaptation to Potato Resistances

Studying phenotypic and genomic modifications associated with pathogen adaptation to resistance is a crucial step to better understand and anticipate resistance breakdown. This short review summarizes recent results obtained using experimentally evolved populations of the potato cyst nematode Globodera pallida. In a first step, the variability of resistance durability was explored in four different potato genotypes carrying the resistance quantitative trait loci (QTL) GpaV_vrn originating from Solanum vernei but differing by their genetic background. The consequences of the adaptation to resistance in terms of local adaptation, cross-virulence and virulence cost were then investigated. Finally, a genome scan approach was performed in order to identify the genomic regions involved in this adaptation. Results showed that nematode populations were able to adapt to the QTL GpaV_vrn, and that the plant genetic background has a strong impact on resistance durability. A trade-off between the adaptations to different resistant potato genotypes was detected, and we also showed that adaptation to the resistance QTL GpaV_vrn from S. vernei did not allow adaptation to the colinear locus from S. sparsipilum (GpaV_spl). Unexpectedly, the adaptation to resistance led to an increase of virulent individual’s fitness on a susceptible host. Moreover, the genome scan approach allowed the highlighting of candidate genomic regions involved in adaptation to host plant resistance. This review shows that experimental evolution is an interesting tool to anticipate the adaptation of pathogen populations and could be very useful for identifying durable strategies for resistance deployment.     

Progress in Mapping and Cloning Qualitative and Quantitative Resistance Against <Emphasis Type="Italic">Phytophthora infestans</Emphasis> in Potato and Its Wild Relatives

Cultivated potato is susceptible to many pests and pathogens, none of which is more of a threat to potato agriculture than the late blight disease, caused by the oomycete Phytophthora infestans (Mont.) de Bary. To date all efforts to thwart this most adaptive of pathogens have failed, and early attempts to deploy ‘R genes’ introgressed from the wild Mexican hexaploid Solanum demissum ended in abject failure. With the advent of facile gene mapping and cloning, allied to knowledge of plant resistance gene structure, renewed efforts are leading to mapping and isolation of new sources of late blight resistance in potato wild species, many of which are being performed under the auspices of the BIOEXPLOIT project (Sub-project 2). We document recent advances in late blight resistance gene mapping and isolation, and postulate how these genes, allied to knowledge of pathogen effectors and their recognition specificity, may greatly enhance our chances of halting the progress of late blight disease in potato crops worldwide.     

Analysis of <Emphasis Type="Italic">Potato Mop-Top Virus</Emphasis> Survival Probability in Post-Harvest Storage

Potato mop top virus (PMTV) induced necrosis can cause tuber quality loss at harvest and storage. Stored potatoes help maintain a constant supply of potatoes to the market and processing industry. PMTV-induced necrosis continues to develop during storage and appropriate timelines are needed for growers to make marketing decisions of their potatoes before incurring any significant quality losses. Survival analysis was used to estimate the time to event occurrence (PMTV-induced necrosis) in four (red-, russet-, white-, and yellow-skinned) potato market types across six post-harvest assessments conducted over two years. At each assessment the presence and absence of PMTV-induced necrosis was recorded and probability of tuber survival was estimated. Survival curves were significantly different among the four market type potato cultivars (Log-Rank test, P < 0.0005). Red- and russet-market type cultivars showed low and high survival probability, respectively, demonstrating that red cultivars need priority marketing. The survival probabilities decreased with increased storage time during both years, indicating that PMTV-induced necrosis development is dependent on potato cultivar and post-harvest storage. The median (50% of tubers with symptoms) survival times were estimated as 167 and 214 days for red- and other market type potato cultivars. The information from this study could potentially help growers regulate storage times for their cultivars to minimize tuber quality loss due to PMTV-induced tuber necrosis.