The effect of soil- and tuber-borne inoculum on the incidence of potato gangrene

Under optimum growing conditions neither tuber- nor soil-borne Phoma exigua var. foveata inoculum appreciably affected stand or yield of the subsequent potato crop. Seed tubers with gangrene rots caused high levels of stem and tuber symptoms when planted in var. foveata contaminated or uncontaminated land; contaminated seed tubers with no rots also produced progeny with a high gangrene potential. Sufficient soil-borne inoculum was carried over in land that produced a gangrene affected crop in the previous year to override the effect of tuber disinfection. Effective gangrene control was achieved by a combination of tuber disinfection shortly after harvest over successive years with a 1 in 5 yr potato crop rotation. Gangrene rots usually developed through injuries to the tuber periderm, rots in other tubers being associated with pustules of powdery scab (Spon-gospora subterranea).     

The Influence of the Inoculum Source of Rhizoctonia solani on Development of Black Scurf on Potato

Rhizoctonia solani, the causal agent of stem canker and black scurf on potato, survives as sclerotia on tubers, in soil and in plant residues. The objective of the present study was to evaluate the importance of inoculum source on disease development. Disease‐free minitubers and seed tubers contaminated with low levels of R. solani were planted in fumigated or artificially inoculated growth mixture in greenhouse experiments. Black scurf incidence and severity were significantly higher when the inoculum was present in both seed tubers and soil, compared with either of them separately. The severity of disease symptoms on the subterranean parts of the plant also were significantly higher in plots where both seed tubers and soil were contaminated, compared with plots where the inoculum source was either the seed tubers or the soil. Thus, both major sources of inoculum, seed tubers and soil, are important in disease development. However, when both sources are present, black scurf incidence and severity are increased, leading to economical damage to tuber yield and quality. Additional results from field trials support these findings. Disease incidence and severity on daughter tubers were correlated with levels of contamination in seed tubers and soil. When seed tubers and soil were heavily infested, the levels of black scurf incidence and severity on daughter tubers were very high; when seed tuber and soil infestation were very low, black scurf incidence and severity on progeny were also lower. Disease levels were reduced by in‐furrow fungicide treatment, but were less effective when the initial levels of the fungus on the seed tubers and in the soil were high.     

Infection of potato stems by the gangrene pathogen, Phoma exigua var. foveata

Limited transport of pycnidiospores of Phoma exigua var. foveata within transpiring stems was demonstrated by isolating from cut mature stems placed for 24 h in a spore suspension. Pieces of shoot dipped in spore suspension and subsequently rooted contained the pathogen which was redistributed as the plants grew but without causing symptoms until after desiccation. The fungus was only rarely detected within stems growing from rotted seed tubers suggesting that spore transport in the vascular system was relatively unimportant in these experiments. Stems were readily infected from spore suspensions applied to unwounded epidermis especially when the inoculum was held in cotton wool and secured with wax film. The progressive colonisation of growing stems was demonstrated by isolations made at different positions on the stem and at different times after inoculation. In field experiments in two years stems were inoculated with four spore concentrations on four dates from late June to late August. After desiccation the extent of pycnidial lesions was greatest from the earliest inoculation date and the largest spore concentration and these differences were reflected subsequently in the incidence of gangrene on damaged progeny tubers. Younger stems may be more susceptible and infection in early summer probably results in extensive growth within the stem before desiccation and thus in massive pycnidial production.     

Persistence of Phoma exigua xar. foveata and Polyscytalum pustulans in dry soils from potato stores in relation to reinfection of stocks derived from stem cuttings

The incidence of Phoma exigua var. foveata and Polyscytalum pustulans in dry soil and dust from potato stores was assessed at 10 farms in Scotland producing virus-tested stocks of seed potatoes derived from stem cuttings (VTSC). Samples were collected on three occasions during 1976: during the storage period (February-March), soon after the potatoes were planted (May) and just before the new crop was lifted and stored (August-October). Both pathogens were detected at all three sampling times but P. exigua var. foveata was more frequently detected at the last one than P. pustulans. Soil and dust from various sites within the stores, such as floors, ledges, graders and boxes, were contaminated by these pathogens. Propagules of both pathogens remained airborne for at least 12–17 min after the floors were swept. P. exigua var. foveata remained viable for at least 16 months in dry soil from tuber surfaces stored at 4–6°C. When VTSC tubers were dusted before planting with a dry store soil contaminated by P. exigua var. foveata the gangrene potential of the progeny tubers was much greater than that from untreated tubers or from tubers treated with an uncontaminated soil. The role of inoculum surviving in dry soil is discussed in relation to reinfection of VTSC stocks.     

Transmission of Fusarium solani var. coeruleum and F. sulphureum from seed potatoes to progeny tubers in the field

In experiments in three years, seed tubers were inoculated before planting with either Fusarium solani var. coeruleum or F. sulphureum to initiate a rot, or were contaminated by dipping in soil slurries containing spore suspensions of one or other of the pathogens. Transmission to progeny tubers was tested by uniformly wounding and incubating tubers and by dilution plating of soil samples. In two years, transmission of F. sulphureum was greater from highly contaminated than from rotting seed and was greater on cv. Pentland Crown than on cvs Desiree and Maris Piper. F. solani var. coeruleum appeared to be transmitted most readily from rotting seed and Maris Piper was the cultivar most extensively contaminated. In experiments with different harvest dates, transmission of both fungi from highly contaminated seed could be detected by late June or mid-July. More progeny tuber wounds rotted in F. sulphureum than in F. solani var. coeruleum plots and in one year, F. sulphureum caused more rots on cv. Record than on cv. Maris Piper. These differences between the pathogens may be related to their differing abilities to sporulate underground on the surface of seed tubers and on stem bases.     

Contamination of seed potato tubers grown from stem cutting or microplant-derived tubers by Phoma foveata and Polyscytalum pustulans in Scotland: the influence of site and fungicide treatment at harvest

A survey was made over 5 years of the incidence of contamination by Phoma foveata and Polyscytalum pustulans on 3-year-old virus-tested stem cutting (VTSC) stocks assessed on three occasions during the storage season, and on their daughter tubers at five farms. The contamination occurring on plants grown in small plots from stem-cutting/microplant tubers was also assessed at the farms.
Fungicide treatment soon after harvest reduced the incidence of both pathogens, especially P. pustulans . Tuber contamination was similar for samples collected after fungicide treatment and before planting. The incidence of contamination on daughter tubers, relative to that on the untreated mother tubers, was less with P. foveata but was similar or greater with P. pustulans . Contamination by both pathogens varied between farms and was reflected in the degree of contamination occurring on daughter tubers from stem-cutting/microplant tubers.
The incidence of P. foveata on stems, soil and tubers of seed stocks classified at Foundation Stock grade from various regions of Scotland was assessed over 2 years. The incidence of tuber contamination was correlated ( P < 0.05) with the amount of tuber-borne soil contamination for all comparisons, but was correlated with stem inoculum in only 7 out of 13 comparisons. The largest amounts of stem inoculum of P. foveata were found in stocks in Aberdeenshire.
These results are discussed in relation to the spread of both pathogens to healthy VTSC stocks.     

Factors affecting the occurrence of gangrene (Phoma exigua) in potatoes

The presence in soil from Scotland and England of Phoma exigua f. sp. exigua and P. exigua f.sp. foveata, which cause gangrene, is confirmed by isolation, and it is established that infection of tubers occurs before lifting, and after lifting from soil adhering to tubers. The distribution of the disease is related to soil moisture, gangrene being most prevalent in tubers from the north-eastern counties of Scotland where the moisture content of arable soils remains high throughout the growing season. The incidence of the disease may be affected by the haulm, either as a source of infection—though not an important source—or by its effect on the maturity of the tubers. The incidence of gangrene is less where haulm destruction is rapid. The incidence of gangrene in a crop is not related to its incidence in the seed tubers planted and, unless infection is severe, the effect on yield is not serious. Symptoms of skin necrosis were associated with tubers from acid soils, infected with P. exigua f.sp. foveata and stored at low temperature.     

Experiments with foliar sprays of thiabendazole and captafol for control of potato gangrene

In an experiment designed to measure the effect of foliar sprays of fungicides on gangrene in stored tubers thiabendazole but not captafol reduced yield by 30%. Although both fungicides reduced disease incidence in stems and tubers the extent of the reduction was so small that foliar sprays were considered unimportant as a method of control. Sprays of thiabendazole and captafol applied to the haulm 1 wk or 1 wk and 2 wk after desiccation did not reduce the incidence of gangrene in storage.     

Sampling potatoes for the incidence of tuber diseases and levels of inoculum

In three years, between three and six seed stocks of cv. Pentland Crown were planted in field experiments. Seed tubers, stem bases and progeny tubers from the growing crop and tubers at harvest and after storage were sampled intensively and estimates of inoculum and disease made for five principal tuber diseases: the blemishes skin spot (Polyscytalum pustulans), silver scurf (Helminthosporium solani) and black scurf (Rhizoctonia solani) and bacterial soft rot (Erwinia carotovora). The data were principally used to calculate sample variances after transformation of % values to logits. A variance of 0.010 was selected as giving acceptable accuracy for most purposes and to achieve such values substantial numbers of samples would have been needed. Five to ten 50-tuber samples of seed tubers or 15–20 samples of a harvested or stored crop would have been required for assessing disease symptoms and similar numbers of samples for assessing gangrene potential by a standard damage test. The eye plug test for assessing inoculum of the three blemishing diseases used 15 or 20-tuber samples and five such samples of seed or about ten from the growing crop would usually have been adequate. Variances were nearly always greater for black scurf assessments than for other blemishing diseases. The data also illustrated changes in inoculum and disease levels during the season and in store. Their significance for understanding the epidemiology of the diseases and for storage disease forecasting is discussed.     

The role of water in the spread of Phytophthora infestans in the potato crop

During falls of rain before 5% of the haulm was destroyed by blight, most rain water was deposited on soil at the bottom of the ridge, and least at the top in crops of King Edward, Majestic, and Up-to-Date potatoes. Afterwards less water was trapped half way up the side of the ridge than above or below. Water was also channelled down the stems and, while these were upright, was deposited in the channel often formed around the stem bases by wind movement. Fewer than 500 sporangia per ml. were seen in water collected when blight was present in the crop. Growing tubers of Ulster Ensign and King Edward, inoculated with P. infestans, infected healthy tubers less than 1–3 cm. away, when the soil water content was greater than 20%, and sporulation on the inoculated tubers was maximal. Similarly inoculated Up-to-Date and Majestic tubers sporulated less abundantly and failed to infect healthy tubers. Sporulation also occurred on inoculated seed tubers, although infection of the haulm from these tubers could not be confirmed. Water movement from the seed tuber region to the furrows may occur under suitable conditions.     

Development of potato blight Phytophthora infestans after planting infected seed tubers

About 1000 blight-infected seed potato tubers, usually of the cultivar King Edward, were planted for 9 yr and the subsequent plants examined until the disease had developed in the plots. Haulm infection originated each year from the seed tubers and occurred first on basal leaves. When tubers were inoculated with a complex race of P. infestans this race was recovered from the leaves and from the soil near the seed tuber. Transmission of infection from soil to leaves was demonstrated by splash of artificially contaminated soil to leaves suspended above the soil. In 4 yr, plants were grown on flat rows as well as on ridges. In 2 yr, when emergence was almost complete, infected stems were observed on otherwise normal plants. In the first year 0.6% grew on ridges and 3.0% on the flat and in the second all grew on the flat (5.3%). Only seven of the 43 plants had more than one infected stem. Flat plots had a significantly higher number of stemdplant than ridge plots, but this bore no relation to numbers of infected stems. When flat plots which had developed affected plants had soil replaced as ridge plots, no further infected stems were observed. Such stems continued to develop on flat plots. No prematurely dead stems were observed below soil level when all plants were dug. Underground portions of most infected stems showed little evidence of P. infestans which was found only at about soil level. Infection appeared to occur first in this area.     

Healthier seed potatoes. I. Effects of inoculating stem cutting stocks with Polyscytalum pustulans and Rhizoctonia solani on growth, yield and disease

Potato seed tubers of seven cultivars derived from stem cuttings in 1965 (healthier seed) were grown in 1969–72 at two sites, one clay with flints soil (Rothamsted) and the other sandy loam soil (Woburn). Inoculating sprouted tubers at planting with Polyscytalum pustulans did not affect the number of stems/plant or total yield but increased stem base and tuber infection. The yields of large tubers (57–83 cm) were increased and small tubers (>57 cm) decreased, indicating a decrease in tuber numbers. Rhizoctonia solani inoculated at planting decreased numbers of stems/plant and yield by up to 14% at Woburn but not at Rothamsted. At both sites, yields of large tubers were increased with cvs Majestic and Record and decreased with Pentland Crown. Stem canker and tuber infection were increased but infection was also prevalent on tubers from non-inoculated seed at Woburn. When both pathogens were inoculated together yields of large tubers were increased in cvs King Edward, Majestic and Record and decreased in Pentland Crown. Infection of stem bases and tubers was sometimes less than when either pathogen was inoculated singly.     

The effect on gangrene incidence of dusting seed potatoes with benomyl at time of planting

In 1 out of 2 years' field trials benomyl applied as a dust treatment at time of planting seed potatoes resulted in an increase in potato gangrene in the progeny of both gangrene-free tubers planted in land contaminated with Phoma exigua var. foveata and gangrene-diseased tubers planted in clean land. Gangrene-infected seed tubers treated with benomyl also produced more stems infected with var. foveata than untreated tubers. Two hypotheses are presented to account for this increase in gangrene which does not occur in seed potatoes treated shortly after lifting. In all cultivars tested an organo-mercury dip-treatment increased total numbers of tubers in the seed and chat-size grades without increasing total weight whereas benomyl dust increased the numbers in these grades in Majestic only. The treatment of gangrene-diseased seed with benomyl dust affected neither total weight nor total number of tubers.     

The development of silver scurf (Helminthosporium solani) disease of potato

The main source of inoculum of Helminthosporium solani was the seed tuber; the fungus was not detected in soils in which infected potatoes had been grown the previous year. Lesions spread rapidly on seed tubers after planting, so that within a few weeks the entire surface was covered. As lesions aged they lost the capacity to sporulate, so most inoculum was produced in the first few weeks after planting. Infection of progeny tubers was first detected at the heel (stolon attachment) end soon after their initiation. Once established, lesions spread slowly on the surface of progeny tubers when in the soil; more rapid spread took place during storage. Potatoes which seemed unaffected at lifting became diseased during storage.     

Effects of treating seed potatoes from commercial and stem cutting stocks with benomyl, thiabendazole and 2-aminobutane on yield and disease

Potato seed tubers of six cultivars from commercial stocks and from stocks derived from stem cuttings (healthier seed) were fumigated with 2-aminobutane 2 wk after lifting or treated with benomyl or thiabendazole in January. 2-aminobutane prevented skin spot and gangrene developing on treated tubers. Experiments were planted at Rothamsted (clay with flints soil) and at Woburn (sandy loam soil) in 1973–75. Healthier seed produced more stems/plant than commercial stocks and yielded on average 8% more at Rothamsted in 1973 and 1974 and respectively 5 and 10% more at Woburn in 1973 and 1975. Seed treatments did not consistently affect stem numbers or increase yield although all treatments tended to decrease tuber size. Infection of stem bases and tubers by Polyscytalum pustulans and Rhizoctonia solani was usually less from healthier than from commercial seed and was decreased by benomyl and thiabendazole in 1973 and 1974. Infection by Helminthosporium solani of the skin around tuber eyes was greater from healthier than from commercial seed but was decreased by benomyl and thiabendazole. 2-aminobutane sometimes decreased infection of tubers by P. pustulans and R. solani but neither of stem bases nor of tubers by H. solani. Gangrene on tubers uniformly wounded at lifting was not consistently affected by seed source or seed treatment. Treating seed with benomyl or thiabendazole in 1975 decreased skin spot and black scurf in tubers stored until March 1976. These treatments also decreased silver scurf on the produce of commercial seed at Rothamsted but gangrene was not consistently affected by seed treatments.     

Effects of common scab (Streptomyces scabies) on potatoes

Seed tubers with severe (71% cover) or slight (12% cover) common scab selected from a badly infected crop of cv. Maris Piper were boxed for sprouting in November and February and planted in field experiments at Woburn and Rothamsted in 1980. The severely scabbed seed lost more moisture than the slightly scabbed seed during sprouting and sprouted from a larger number of eyes but total stem numbers were greater on plants from the slightly scabbed seed. Plant growth during the first 6–9 wk after planting was less from the severely scabbed than from the slightly scabbed seed. This was shown by records of ground cover, leaf area index, fresh weights of stems and leaves, total tuber yield (18% lower at 9 wk) and tuber number. Subsequently these differences became non-significant. Seed with slight scab boxed in November produced smaller numbers of progeny tubers than that boxed in February but there were no other consistent effects of boxing date on growth and yield. Scab incidence on progeny tubers was very slight and unrelated to the severity of disease on the seed tubers. It seems unlikely that severe scab infection of seed tubers would significantly decrease tuber yields except perhaps in early potato production.     

The phenology of skin spot (Oospora pustulans Owen &Wakef.) and other fungal diseases of potato tubers

Underground stems and roots of the potato varieties King Edward and Majestic became more severely infected by Oospora pustulans (skin spot) as the growing season advanced. Tubers became infected at their initiation in June and July but the proportion of infected eyes usually increased during both the growth of the crop and bulk storage. Some buds on tubers in bulk stores died in December, but few were killed until after January, when attacks increased until by March about a quarter of Majestic eyes were usually dead. Boxing (chitting) not only prevented most eyes from dying but also prevented much skin spotting on tubers. At High Mowthorpe, Majestic seed unboxed or boxed in March yielded, respectively, 3 and 1 ton/acre less than seed boxed by January. Early boxing increased the yield of seed–sized tubers but did not affect the infection of progeny tubers. Dipping Majestic seed tubers in organo–mercurial fungicide and then boxing soon after lifting decreased their infection by O. pustulans; it also lessened infection on plants they produced and increased the yield of progeny seed–sized tubers. However, dipping, even in successive years, had little effect on the infection of progeny tubers by O. pustulans. Rhizoctonia solani and Helminihosporum atrovirens became increasingly prevalent on tubers during growth after July and during bulk storage. H. atrovirens was decreased by early boxing or dipping tubers in fungicide. Verticillate conidiophores often grew on incubated tuber plugs but their prevalence was affected by past storage. The results suggest that some of the benefit for chitting may be from disease control, it was not possible to measure the extent to which benefits from chitting depend on effects on disease or on the physiology of the tuber.     

Development of black dot disease (Colletotrichum coccodes (Wallr.) Hughes) and its effects on the growth and yield of potato plants

Seed tubers of cvs Désirée and Pentland Crown with different severities of black dot were planted in 1988 and 1989 at Rothamsted in fields in 4– or 7-course rotations, respectively. Tubers treated with prochloraz (1988) or imazalil (1989) were planted in some plots, and in others Colletotrichum coccodes inoculum was added to the soil at planting. In further experiments at Mepal, Cambridgeshire in 1989 and 1990 and at Rothamsted in 1990 on sites where potatoes had not been grown for more than 15 years, large amounts of inoculum were added to the soil around disease-free seed tubers of two (1989) or three (1990) cultivars at planting. In all experiments plants were sampled during the season and the effects of treatments on disease development, growth and yield were recorded. Disease on roots, stem bases and tubers was found early in the season and was more severe on Désirée than on Pentland Crown plants from fields in 4– or 7-course rotations. Severity increased throughout the season and with increasing amounts of disease on the seed tubers, especially with Desiree. Disease was also found on plants from disease-free tubers and was more severe in 1988 than 1989. At harvest, black dot on tubers was significantly more severe from severely affected than from disease-free seed, and was most severe where inoculum, especially large amounts, had been added at planting. Fungicide treatment decreased disease early in the season but had no effect on tuber infection at harvest. In 1989 the weight loss of seed tubers during sprouting increased with increasing amounts of black dot, but the disease had little effect on plant size through the season. At harvest the yield of ware tubers (>50 mm) decreased with severe disease but total tuber yields were not significantly affected. However, at harvest in 1988 severely affected seed yielded significantly less than healthy seed. Plants grown from mini-tubers were free from disease on sites where potatoes had not been grown for at least 15 years. Inoculum applied at planting caused severe disease on all cultivars in both years, whereas disease was slight on uninoculated plants. Inoculated plants senesced early at Mepal in 1990, but there were no significant differences in total tuber yield in any experiment. However, yields of ware tubers (>50 mm) were sometimes decreased and the total tuber number per plant increased.     

Black scurf and stem canker of potato (Corticium Solani Bourd. & Galz.): Further field studies on the use of clean and contaminated seed potatoes and on the contamination of crop tubers

Black scurf and stem canker of the potato were investigated in field trials in contaminated soil at Warburton, Cheshire, to determine the effect of planting clean and contaminated seed (vars. Arran Banner and Majestic) on crop yield and on the contamination of the crop tubers.
Black-scurf contamination was prevalent on crops grown from clean seed and was not significantly different from that occurring on plots planted with contaminated seed. It was severe on early-dug tubers, but was more severe on late-dug tubers. Produce grown with and without stable manure was heavily contaminated.
Contaminated seed caused a check to tuber formation and an appreciable increase in stem canker and in the number of primary shoots killed. Nevertheless, the yields from the manured, contaminated seed plots were satisfactory and were not lower than those from the clean-seed plots. There was no relation between the yield and the amount of black scurf on the produce.
The results of a trial made in a field which had been in grass for at least 43 years, suggested that the soil contained little, if any, Corticium Solani. Heavy contamination occurred on tubers grown under relatively dry soil conditions.
In all the trials misses and wilted shoots caused by Corticium were rare and there was no premature yellowing or death of the haulms on the clean or contaminated seed plots.
The evidence obtained in the four seasons from 1941 to 1944 indicates that in this country satisfactory yields of early maincrop and maincrop varieties may be obtained despite the prevalence of C. Solani in the soil and on the seed, provided the soil and cultural conditions are reasonably good.     

The effects of various chemical and physical haulm treatments on the incidence of potato gangrene

In studies of the influence of haulm defoliants on gangrene incidence in storage the use of diquat dibromide was consistently found to exacerbate levels of Phoma exigua vzr.foveata infection particularly on tubers harvested 3–4 wk after burning down. Intermediate levels occurred where the haulm was chemically defoliated with dinoseb or was physically cut and removed and least where the defoliants were sodium chlorate or sulphuric acid. Pycnidia of P. exigua var. foveata developed within 10 days on stems desiccated with diquat dibromide, sulphuric acid or dinoseb and most prolifically on those treated with diquat dibromide but tuber infection was not always related to their abundance. Speed of kill was not considered to be important in determining effect on gangrene incidence.