Factors affecting the temporal progress of stem canker (Rhizoctonia solani ) on potatoes (Solanum tuberosum)

The effects of different densities of tuber-borne inoculum, selected agronomic treatments (date of planting, irrigation and size of seed tubers) and their interactions on the temporal progress of stem canker ( Rhizoctonia solani ) on potato plants ( Solanum tuberosum ) were investigated in a multifactorial experiment. Data comprising stem number and the incidence and severity of stem canker from planting until tuber initiation in two consecutive growing seasons were analysed using linear contrasts, quadratic contrasts and the area under the disease progress or host growth curve. Differences in the incidence and severity of stem canker were dominated by the effect of different densities of tuber-borne inoculum. The majority of disease progress curves were nonmonotonic for the incidence and severity of stem canker with a rapid rise in disease up to stem emergence and a decline thereafter. Most treatments affected the area under the curve and to a lesser extent the average rate of increase in disease. Of the agronomic treatments, later dates of planting and pre-emergence irrigation reduced the levels of stem canker whereas size of seed tubers did not affect the progress of disease. Little additional information was revealed by scoring for the severity rather than the incidence of stem canker.     

Relationships between stem canker, stolon canker, black scurf (Rhizoctonia solani) and yield of potato (Solanum tuberosum) under different agronomic conditions

Relationships between diseases caused by Rhizoctonia solani on different parts of potato plants ( Solanum tuberosum ) at different stages of crop growth were investigated under selected agronomic conditions. The effects of different densities of tuber-borne inoculum, date of planting, irrigation, size of seed tubers and their interactions on the incidence of stem and stolon canker during crop growth, the incidence and severity of black scurf and the yield of progeny tubers at harvest were quantified in a multifactorial experiment. Differences in the incidence of stem canker, stolon canker and black scurf were dominated by the effect of density of inoculum on seed tubers at planting. Highly positive correlations between the disease variables indicated a close relationship between the incidence of disease at each stage of crop growth although the degree of association between variables measured at an early growth stage and those measured at progressively later stages of crop growth weakened as the time interval increased. Total yield of progeny tubers was not affected by the density of tuber-borne inoculum although there was a shift in the size distribution, with a decrease in the yield of main-sized tubers and an increase in the yield of baker- and oversized tubers at the higher density of inoculum. Of the remaining factors, the effect of season tended to be more pronounced than any of the agronomic treatments although the use of irrigation and later dates of planting did influence the incidence of infection to a limited extent.     

The occurrence of thiabendazole-resistant isolates of Polyscytalum pustulans and Helminthosporium solani on seed potato tubers in relation to fungicide treatment and storage

Tubers infected with thiabendazole-sensitive isolates of Polyscytalum pustulans and Helminthosporium solani were treated annually for 4 years with benomyl, thiabendazole or a formulated mixture of thiabendazole and imazalil, and grown at three farms in Scotland. The proportion of thiabendazole-resistant isolates of H. solani increased, and isolates producing black colonies became more common with successive annual applications of thiabendazole or benomyl. Silver scurf was not reduced after three annual applications. When these fungicides were applied once to untreated seed the incidence of resistant isolates of H. solani was much less in 1988 than in 1991 when tubers had been grown on farms for 3 years from untreated seed.
The proportion of resistant isolates of P. pustulans increased with the number of successive applications of thiabendazole or benomyl but at differing rates on each farm. At one farm, skin spot was not reduced by three annual applications of these fungicides whereas at the other farms it was reduced by 90–100% by four annual applications.
A smaller proportion of resistant isolates of P. pustulans and H. solani was obtained after applying the mixture of thiabendazole and imazalil than after benomyl or thiabendazole alone. Their occurrence was not related to the number of fungicide applications. The mixture also reduced both diseases by more than 75% over the 4 years.
At one farm where resistant isolates of H. solani were present, tubers were infected when stored on trays but not when stored in bags.     

Effect of temperature on the transmission of Potato mop-top virus from seed tuber and by its vector, Spongospora subterranea

Potato mop-top virus (PMTV) causes disease in both the growing plant and tubers (spraing) of potato and is transmitted by the plasmodiophorid Spongospora subterranea , the cause of powdery scab. The effect of temperature during plant growth on the transmission of PMTV from infected seed tubers and from infested growing media was investigated in a series of glasshouse experiments. Symptoms developed on foliage of plants derived from infected seed tubers but none developed when PMTV was transmitted by S. subterranea in soil. The incidence of foliar symptoms was greatest on plants grown at 12°C, less at 16°C, few at 20°C and absent at 24°C. The transmission of PMTV from infected seed tubers was not significantly affected by temperatures between 12 and 24°C, but when the virus was transmitted by S. subterranea , minimal tuber infection occurred at 24°C and no differences were recorded at temperatures between 12 and 20°C. The incidence of powdery scab on tubers was greatest at 12 and 16°C and very low at 20 and 24°C. However, the incidence and severity of root galling caused by S. subterranea , was greatest at 20 and very low at 24°C. The incidence of powdery scab was greater on tubers of plants derived from infected seed tubers grown in a fluctuating temperature regime of 12 h at 20°C followed by 24 h at 12°C than on those grown at a constant 20°C, whereas the incidence of tuber infection by PMTV and spraing was similar for both regimes. This demonstrates that infection of roots can occur at a higher temperature than that for powdery scab on tubers and that this root infection can enable the transmission of PMTV into the potato plant.     

The importance of the infected seed tuber and soil inoculum in transmitting Potato mop‐top virus to potato plants

Potato mop‐top virus (PMTV), the cause of spraing in potato tubers, is transmitted by Spongospora subterranea, the cause of powdery scab, and by planting infected seed tubers. This study was undertaken to determine the relative importance of these sources of infection in seed potato production in Scotland. The transmission of PMTV from tested seed tubers to daughter plants was examined over 2 years and six cultivars. The development of foliar symptoms varied with year and cultivar. Infection of daughter tubers derived from PMTV‐infected seed tubers was more prevalent on plants affected by foliar symptoms than those without symptoms. The rate of transmission of PMTV from infected seed tubers to daughter tubers ranged from 18 to 54%. Transmission was affected by cultivar and by origin of seed tubers used for a cultivar, but not by a cultivar's sensitivity to PMTV infection. The incidence of PMTV in daughter tubers of cv. Cara grown from seed potatoes from one source (common origin) by more than 25 seed producers was examined over two successive generations. The incidence of PMTV in daughter tubers was not correlated with that in the seed tubers but appeared to be strongly associated with soil inoculum. The incidence of PMTV was correlated with powdery scab in those crops in which both were present. There was some evidence from soil tests conducted in 2006 using a tomato bait plant and real‐time RT‐PCR that planting PMTV‐infected seed potatoes could increase the risk of introducing the virus into land not infested by PMTV.     

Pathogen resistance in aerial tubers of potato cultivars

Increases in resistance of aerial tubers (cultivars Désirée, Epicure and King Edward) against nine pathogenic potato fungi were determined in vitro using freeze-dried powder derived from subterranean and aerial tubers as a culture medium, and in vivo by inoculating both tuber types with the storage pathogens Fusarium sulphureum, F . solani var. coeruleum, Phoma exigua var. foveata and Polyscytalum pustulans . Significantly reduced growth rates and spore and conidiophore numbers were recorded when pathogens were cultured on freeze-dried powder derived from aerial tubers. No effects on spore size of F . solani var. coeruleum were recorded but spore and conidiophore lengths of remaining pathogens were reduced and width increased, except for Colletotrichum coccodes , where the diameter of acervuli increased, and P . exigua var. foveata , where spore length and width were reduced. Reduced disease incidence, severity and spore numbers were recorded on aerial compared with subterranean tubers inoculated in vivo with the four storage pathogens. Spore length of P . pustulans was reduced and spore width increased, while spore length and width of P . exigua var. foveata were reduced on aerial compared with subterranean tubers inoculated with these pathogens. No effects on spore size of Fusarium spp. were recorded. Concentrations of glycoalkaloids, chlorogenic acid, calcium and magnesium were higher in aerial than in subterranean tuber tissue, which may have contributed to increased resistance and effects on size and morphology of fungal spores. The implications of using aerial tubers as an alternative technique for the production of seed potatoes and their importance to the potato industry are discussed.     

Inoculum density and spatial pattern of Rhizoctonia solani in field plots of Solanum tuberosum: effects of cropping frequency

Bait plants, comprising micropropagated and commercial seed tubers, were used to estimate the effects of rotation on the density and spatial pattern of inoculum of Rhizoctonia solani in large field plots of potatoes. Soilborne inoculum of R. solani produced economically significant levels of stem canker and its incidence and severity varied with rotation, with most disease in 2-year and less in 4- and 6-year rotations. The rates of loss of inoculum during intercrop periods differed amongst rotations with a rapid fall to low levels occurring after 1 year in a 6-year rotation and after 2 years in a 4-year rotation. Replenishment of inoculum to soil was rapid following the growth of a susceptible crop, with comparatively high levels of infection and disease, even in long rotations. Disease occurred in patches and the size of patches and the density of R. solani within patches differed with cropping frequency. The degree of spatial autocorrelation also differed amongst rotations but there was no evidence for any significant differences in the rate of change of spatial autocorrelation during intercrop periods in the three rotations.     

Development of symptoms caused by Erwinia carotovora ssp. atroseptica under field conditions and their effects on the yield of individual potato plants

Infection of seed tubers by pectinolytic Erwinia species can lead to the development of various symptoms during vegetative growth of potato crops, including non-emergence of plants, chlorosis, wilting, haulm desiccation and typical blackleg. The relationships between types of symptoms and yield are poorly documented, and are investigated by following the development of symptoms in potato plants grown under field conditions from seed tubers artificially inoculated with E. carotovora ssp. atroseptica ( Eca ) , and measuring the yield of each plant. Symptoms were classified into five main types (non-emergence, wilting/chlorosis, blackleg, haulm desiccation and plant death). Each plant was scored for types of symptom on four successive dates; plants without visible symptoms were scored as healthy. The method of inoculation and inoculum concentration proved major factors for the subsequent development of symptoms. Disease development was more severe after vacuum infiltration of bacteria into seed tubers than after shaking tubers in contaminated sand. Disease usually progressed from chlorosis and/or wilting to partial or total desiccation on a given plant. Yield losses varied according to symptom type, but the relationship between symptoms recorded and yield also depended on scoring dates. The data suggest that the beginning of tuber growth might be the most suitable stage for predicting yield losses from symptom observations. In both cultivars studied (Bintje, highly susceptible, and Désirée, moderately resistant), the yield of symptomless plants growing from inoculated seed tubers was significantly less than that of control plants, indicating that the presence of bacteria on the seed tuber was detrimental, even in the absence of visible symptoms. Differences in symptom expression in the field between cultivars matched the level of visible infection of tubers at harvest, as Bintje tubers showed a higher incidence of rot than Désirée tubers.     

Effect of soil inoculum level and environmental factors on potato powdery scab caused by Spongospora subterranea

The effects of soil inoculum level and three environmental factors (soil type, soil moisture regime and temperature) on the incidence and severity of powdery scab caused by Spongospora subterranea were investigated in potato plants grown under controlled environmental conditions. Symptoms of powdery scab on tubers were assessed visually, after which DNA was extracted from tuber peelings and quantified in a real-time polymerase chain reaction assay using primers and a TaqMan® probe specific to S. subterranea to establish tuber infection levels. Soil inoculum concentration of S. subterranea did not significantly affect the incidence and severity of either tuber infection or powdery scab symptoms at maturity. No significant differences in disease incidence and severity were found between sandy, loamy and clay soils, although the two lighter soils yielded more powdery scab than clay soil. Constant dampness of the soil resulted in significantly more disease than a fluctuating moisture regime. Infection and disease levels were high at all three temperatures tested (9, 12 and 17°C), but symptoms were most severe at 12°C. The percentage of plants with infected tubers did not increase after tuber initiation, although the amount of S. subterranea DNA detected in tubers and the severity of powdery scab symptoms increased in mature plants. Latent tuber infections were found to be common, especially under conditions suboptimal for disease development. This new information may be important for the prevention of powdery scab in potato-growing areas around the world.     

Influence of light-enhanced glycoalkaloids on resistance of potato tubers to Fusarium sulphureum and Fusarium solani var. coeruleum

Increases in resistance to Fusarium sulphureum and Fusartium solani var. coeruleum were determined in seed tubers of cultivars Desiree, Pentland Hawk and Kerr's Pink with light-enhanced glycoalkaloids. Glycoalkaloid concentration in the tubers ranged from 78.0 to 885.9 mg kg⁻¹ fresh weight. Exposure of tubers to light prior to inoculation decreased fungal invasion, the greater reductions generally reflecting longer light exposure periods. In most cases, no direct relationship between resistance and glycoalkaloid concentrations was recorded for F . sulphureum. Surface infection indices and reduced depth of rot caused by F. solani var. coeruleum were closely correlated with tuber glycoalkaloid content. No significant effects on spore length and width were recorded. Exposure to light of seed tubers that are not destined for human or animal consumption could provide a pretreatment to reduce fungal invasion by F. sulphureum and F. solani var. coeruleum .     

Spread of Phoma exigua var. foveata to healthy potato plants

Contamination of stem-cutting bait plants exposed at various locations in and around a potato crop was greatest within the crop, less outside it and least at a site c. 20 km from commercial crops. Within the crop about half the stem infection and tuber contamination detected had occurred before haulm destruction. The incidence of contamination on tubers of bait plants placed within the crop was similar to that on tubers from the surrounding plants. Phoma exigua var. foveata was recovered from green leaves sampled at intervals between June and early September. In one year contamination by P. exigua var. foveata of tubers sampled from healthy plants surrounding a diseased plot was greatest in samples adjacent to the plot and decreased progressively away from it. This pattern was less apparent in two other years. These results are discussed in relation to the spread of airborne inoculum.     

Direct infection of potato tubers by the root parasite Orobanche aegyptiaca

Parasitic plants of the Orobanchaceae are known as obligate root parasites that develop haustoria that connect to roots of various host plants. This article describes, for the first time, a case where the root parasite successfully connected to potato tubers, i.e. to the swollen portion of an underground stem rather than to a root. The rhizosphere of Orobanche aegyptiaca and of its host Solanum tuberosum (potato) was carefully examined. In anatomical studies, the adventitious roots were directly connected to potato tubers. Numerous secondary haustoria, which developed along the adventitious roots in close vicinity to the potato tuber, penetrated the tuber epidermis and the perimedullary tuber parenchyma and developed xylem strands that are presumably connected to the minor xylem strands within the tuber cortex. These findings indicate that parasites of the Orobanchaceae that normally attack host roots may also parasitise underground stem tubers.     

Development of resistance to thiabendazole in Helminthosporium solani (silver scurf) as a result of potato seed tuber treatment

Applying thiabendazole to potato seed tubers affected with silver scurf caused by Helminthosporium solani sensitive to thiabendazole decreased the severity of disease on progeny tubers at harvest, but about 50% of the isolates from these were resistant to the fungicide. The disease was not decreased when samples of the progeny tubers were treated with thiabendazole and planted in the following year, and the incidence of resistant isolates increased. Resistant isolates continued to be present when tubers were planted in the next 2 years without fungicide treatment. Treatment with a mixture of thiabendazole and imazalil also decreased the disease and fewer isolates were resistant than when treated with thiabendazole alone, although the proportion increased after treatment with the mixture in the following year. When seed tubers were infected with thiabendazole-resistant H. solani , silver scurf on progeny tubers was not affected by thiabendazole applied to the seed tubers but was decreased by the mixture of thiabendazole and imazalil. Imazalil was equally effective against H. solani sensitive or resistant to thiabendazole.
Some isolates of H. solani had grey aerial mycelium and of 516 of these isolates obtained in 4 years 29% were resistant to the fungicide. Other isolates produced small, black colonies and their frequency increased with thiabendazole treatment of seed tubers. Of 244 of these isolates, 62% were resistant.     

Relative importance of seed‐tuber and soilborne inoculum in causing black dot disease of potato

Controlled‐environment and field experiments were done to quantify the individual contribution of seed‐tuber and soilborne inoculum of Colletotrichum coccodes in causing black dot disease of potato tubers. Seed‐tuber and soilborne inocula of C. coccodes were quantified using an existing real‐time PCR assay and related to subsequent incidence and severity of disease. In four field trials, a controlled‐environment experiment and through the monitoring of 122 commercial crops, seed‐tuber inoculum was found to be relatively less important than soilborne inoculum in causing black dot, and the level of seed‐tuber inoculum did not significantly affect either the incidence or severity of disease or the percentage of progeny tubers deemed unmarketable. By contrast, soilborne inoculum had the potential to result in high levels of disease and the level of C. coccodes soil infestation (pg DNA g^?1 soil) was found to have a significant effect. At soil infestation levels below 100 pg DNA C. coccodes g^?1 soil, 7% of commercial crops had an incidence of black dot greater than 20%, increasing to 40% and 57% of crops at levels of 100–1000 pg g^?1 and >1000 pg g^?1 soil, respectively. These arbitrary threshold levels for soilborne inoculum related to disease risk are discussed. Interpretation of disease risk based on inoculum levels must, in the future, be informed by agronomic variables and potential control strategies.     

Resistance to thiabendazole in isolates of Polyscytalum pustulans (skin spot) and Fusarium solani var. coeruleum (dry rot) in Scotland

Between 1988 and 1991, isolates of Polyscytalum pustulans were obtained from potato tubers grown in Scotland and tested for sensitivity to thiabendazole (TBZ) on agar. Sensitive isolates did not grow at 1 mg TBZ/1, whereas growth of resistant isolates was unaffected at 1 mg and reduced by 10–30% at 10 mg/1. Resistant isolates were present on all classes of seed potatoes, and were also recovered from the air of stores during grading. Isolates from a stock of seed tubers from Northern Ireland were resistant. All 44 isolates of Fusarium solani var. coeruleum collected in 1991 were sensitive to TBZ.     

Factors affecting the incidence and severity of Spongospora subterranea infection and galling in potato roots

The incidence and severity of root infection and root galling caused by Spongospora subterranea were assessed in potato plants (cv. Estima) grown under controlled environmental conditions. The effects of temperature, soil type, soil moisture regime and soil inoculum level on infection and root gall development were determined by molecular and visual methods at two plant growth stages. Root gall severity was scored at harvest, after which DNA was extracted from the roots and quantified in a real-time polymerase chain reaction (PCR) assay specific for S. subterranea . Root galling was severe at 17°C, with a disease score of 3·1 on a 0–4 scale, low (0·6) at 12°C, and did not occur at 9°C. The level of inoculum in soil, in the form of artificially added sporosori, had no effect on the incidence and severity of visual symptoms, with 21%, 41% and 33% incidence observed at 5, 15 and 50 sporosori g⁻¹ soil, respectively. Incidence of infection, as detected by the real-time PCR assay, was greater with increasing soil inoculum concentrations, ranging from 48% at 5 sporosori g⁻¹ to 59% (15 sporosori g⁻¹) and 73% (50 sporosori g⁻¹) of plants infected at maturity, but this effect was not statistically significant. No correlation was found between the occurrence of galls on roots and powdery scab on tubers of the same plants.     

Anastomosis group and pathogenicity of isolates of Rhizoctonia solani from potato crops in South Australia

Isolates of Rhizoctonia collected from the stems, roots, tuber sclerotia and soil of potato crops in Virginia and Lenswood, South Australia, were identified to anastomosis groups (AG). Of the 301 multinucleate isolates of Rhizoctonia solani tested, 90% were AG-3, 7% were AG-4 and 2% were AG-5; 12 isolates were binucleate Rhizoctonia spp. This is the first report of isolates of AG-4 and AG-5 causing disease in potato crops in South Australia. All AG-3, AG-4 and AG-5 isolates tested caused rhizoctonia disease symptoms on the potato cultivar Coliban in pathogenicity trials conducted under glasshotise conditions. Both AG-3 and AG-5 isolates caused black scurf and stem cankers, although symptoms of black scurf were less severe with AG-5. AG-4 isolates produced the most severe stem and stolon cankers of all isolates tested. The pathogenicity of tuber-borne inoculum was confirmed by growing plants from sclerotia-infested tubers. AG-8 isolates from diseased barley and wheat produced severe root cankers and caused loss of feeder roots on inoculated potato plants. Results suggest that rhizoctonia disease in potato fields in South Australia is caused by a combination of different anastomosis groups and this has important implications for crop rotations.     

Spongospora subterranea root infection assessed in two potato cultivars differing in susceptibility to tuber powdery scab

Infection by Spongospora subterranea of roots of two potato (Solanum tuberosum) cultivars, either very resistant or very susceptible to powdery scab on their tubers, was studied in a glasshouse experiment. Plants grown in sand/nutrient solution culture were inoculated with S. subterranea sporosori 2 weeks after planting. Plant parameters, the intensity of zoosporangium infection in roots, numbers of Spongospora root galls and amounts of Spongospora DNA in roots, measured using quantitative PCR (qPCR), were assessed at sequential harvests. Inoculation with S. subterranea reduced water use (56 days after planting) by 26% in the tuber resistant cultivar compared with uninoculated plants, and by 60% in the susceptible cultivar. Inoculation did not affect growth of the resistant cultivar, nor shoot mass of the susceptible cultivar, but caused a 38% reduction in root mass of the susceptible cultivar. The intensities of zoosporangium development in both cultivars were similar. The susceptible cultivar had approximately four times more Spongospora root galls g^?1 root mass than the resistant cultivar. Quantitative PCR detected S. subterranea DNA in roots 1 week after inoculation, and indicated a twofold greater amount of pathogen DNA in roots of the susceptible than the resistant cultivar. This study suggests that the S. subterranea zoosporangium stage in host roots is affected differently by host resistance factors than the sporosorus (root gall and tuber scab) stages. The study has also demonstrated the usefulness of qPCR for sensitive and consistent detection of S. subterranea across the duration of potato root infection.     

Detection of <Emphasis Type="Italic">Potato virus Y</Emphasis> in industrial quantities of seed potatoes by TaqMan Real Time PCR

Potato (Solanum tuberosum) is the largest crop in Israel. Production is based on the import of seed tubers from Europe for the spring crop. Imported tubers are generally free from virus infection. The most important virus infecting potato is Potato virus Y (PVY), which may cause severe damage to marketable yields. In Israel, tubers from the spring harvest are stored over the summer for planting in the autumn. It is important to be able to determine the infection rate of seed tuber lots from the spring harvest prior to storage. Commonly, infection is measured by sprouting tubers and measuring virus titre in the leaves using ELISA (the “Growing-On test”), which takes at least 6 weeks to give results. There is a need for a faster method to produce results, such as Taqman Real Time PCR (qPCR), for direct analysis of viral infection in tubers at harvest. To use qPCR as a diagnostic tool, it is necessary to demonstrate that both techniques give comparable results on batches of field-grown tubers. Such a comparison was performed on potential seed tuber lots of 14 different cultivars over three Israeli spring harvests (2013–2015). The agreement between the results of the two techniques was not of high statistical significance. However, the qPCR technique can distinguish well, by binary classification, between tuber lots with a low PVY infection rate (<5% by Growing On test; suitable for seed) and those unsuitable for seed (≥5% by Growing On test). Therefore, qPCR is an appropriate technique for determination of the PVY infection rate of seed tuber lots in Israel.     

Uptake of iprodione and control of diseases on potato stems

When applied to the roots of potato plants growing in different soils iprodione became systemic in the plants. Although iprodione was adsorbed on the soils tested, sufficient fungicide was present in potato stems to prevent infection by Phoma exigua var. foveata on all plants except those grown in peat soils. The amount of fungicide present in the aerial parts of the plant was related to the soil/water distribution coefficients. Application of iprodione to the sprouts of seed potato tubers prior to planting decreased the incidence and severity of infection by Rhizoctonia solani (stem canker) and Polyscytalum pustulans on stem bases.