Factors affecting the field infection of potato tubers of different cultivars by blight (Phytophthora infestans)

The relationships between rain and blight (Phytophthora infestans) were studied in unsprayed crops of cultivars differing widely in foliage and tuber susceptibility. The occasions when tubers were infected depended on rain and not cultivar, but numbers of tubers infected after rain was affected by the blight susceptibility of the cultivar. Infected tubers were first found when less than 5 % (BMS key) of the potato foliage was infected but few fresh infections occurred when 50–75% of the foliage had been destroyed. Some tubers were infected after 8 mm rain (tubers near the surface with even less) but large increases in numbers of tubers infected usually occurred only after 25 mm or more had increased soil moisture to above ‘field capacity’ around the tuber for at least 24 h. The most susceptible cultivars Ulster Ensign and Arran Banner had all plants with some tuber blight, and some plants with all tubers affected and often many lesions per tuber. Cultivars of intermediate susceptibility, King Edward and Up-to-Date, had some plants without blighted tubers, many with a few and very few with all. The more resistant cultivars Majestic and Arran Viking had many plants without infected tubers and many lesions that aborted while still necrotic threads, so that the fungus did not spread. Most infections occurred through tuber eyes, lenticels or sometimes growth cracks. The distribution of blight lesions on tubers differed in the different seasons, for example, lenticels were most commonly infected on Arran Banner and Ulster Ensign and eyes on King Edward, Majestic and Arran Viking. In late or slowly developing attacks, lesions on stems became more numerous and larger than in fast, early attacks and were prolific sources of spores on King Edward and Up-to-Date but not on Majestic and Arran Viking. Because much rain water runs down the stems of Up-to-Date and King Edward, stem lesions can provide an important source of inoculum for tubers.     

Potato tuber lenticels: development and structure

Stomata on growing potato tubers are changed to lenticels by about the third internode from the apical bud (A-3). In wet soil lenticels proliferate, while in dry soil the filling cells become suberized by about the fifth internode (A-5). Lenticels did not proliferate in the field after mid-August, and laboratory experiments on freshly harvested and stored tubers showed that lenticels proliferated less readily as tubers aged. Proliferation occurred more readily on stored tubers induced to sprout than on tubers from which the eyes had been removed, but nevertheless proliferation is unlikely during winter storage.     

Potato tuber lenticels: susceptibility to infection by Erwinia carotovora var. atroseptica and Phytophthora infestans

In laboratory tests, proliferation increased the susceptibility of lenticels on mature tubers to infection by Erwinia carotovora var. atroseptica. No differences in susceptibility to E. c. atroseptica and Phytophthora infestans were detected on tubers lifted at different stages of growth from soil kept wet or dry for 3 wk prior to each sampling. As tubers aged, fewer lenticels became infected by either pathogen or were penetrated by fluorescein dye, presumably because cells had been suberized and cork barriers formed.     

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.     

Effects of skin spot (Oospora pustulans) on potatoes

King Edward and Majestic seed potatoes selected as ‘clean’ (macroscopically symptomless), moderate and severe according to the extent of skin spot were planted in field experiments at Rothamsted between 1964 and 1968. Usually crops from ‘clean’ and moderately infected seed did not differ detectably in growth or yield. Plants from severely infected seed tubers emerged more slowly, had fewer stems and yielded less (King Edward 20 %, Majestic 13 %). Seed infection also affected tuber size distribution; severely infected seed of King Edward yielded almost 4 tons/acre less of 1 1/4-2 1/4 in tubers and Majestic, 1 ton/acre less of these and 2 tons/acre less 2 1/4-3 1/4 in tubers. However, the total yield from diseased seed stocks was only slightly less (King Edward, o-6 ton/acre and Majestic o-8 ton/acre) than the yields from the ‘clean’ tubers selected from them. Seed severely infected by Oospora pustulans often increased infection of the progeny tubers, and usually decreased their infection by Rhizoctonia solani and sometimes by Helmintho-sporium solani. Another series of experiments compared King Edward seed tubers classified according to the number of live eyes showing in March. Seed with one, two, three and more live eyes yielded equally. About half the tubers without live eyes in March eventually produced plants, but late, with few stems and giving only half the yield of seed with three or more live eyes. Surprisingly, the progeny tubers from seed without live eyes were least infected by O. pustulans, R. solani and H. solani. Progenies of King Edward and Majestic seed from a common source grown on seven widely separated farms were infected more in 1963 than in 1964, but in each year infection differed widely between farms. Often where O. pustulans was common, R. solani was scarce and vice versa. By contrast, when King Edward stocks very differently infected by O. pustulans were grown at Rothamsted their progenies were almost uniformly infected by O. pustulans and R. solani.     

Laboratory studies of potato tuber resistance to infection by Phytophthora infestans

Eyes and lenticels on tubers of cvs King Edward, Record, Maris Peer and Majestic gradually became more resistant to infection by Phytophthora infestans as the growing season progressed. Lenticels of Record and Majestic, and eyes of Maris Peer and Majestic were more resistant than those of King Edward. Differences in resistance between cultivars were apparent at all concentrations of inoculum.     

Effects of water potential on the infection of potato tubers by Streptomyces scabies in soil

When soil was maintained at a mean water potential (Slatyer & Taylor, 1960) of the order of -80 J kg^-1 at 25 cm depth throughout the growing period, much infection of potato tubers by Streptomyces scabies occurred, but when soil was irrigated to maintain it at potentials greater than - 13 J kg^-1 at 10 cm depth infection was negligible. Until about 5 weeks after initiation, tubers were very susceptible to infection; irrigation during this period reduced scab considerably, but subsequent irrigation reduced it only slightly. At the low water potential, actinomycete populations on lenticels were high and bacterial populations low; these effects were reversed at the high water potential. Since also actinomycetes were more frequently isolated in the absence than in the presence of bacteria it was deduced that there was an interaction between these two groups. It is suggested that irrigation may decrease the population of S. scabies in tuber lenticels by increasing populations of bacteria antagonistic to it.     

The influence of potato harvesting and grading machinery on contact spread of Phoma exigua var. foveata on tubers

The incidence of wounds infected by Phoma exigua var. foveata was increased if freshly damaged tubers (recipients) were shaken in a bag with diseased tubers (donors) to simulate the tuber-to-tuber contact that occurs during potato handling. An increase in the number of gangrene rots on damage points also occurred if the recipient tubers were wounded after contact with the diseased tubers, rather than before, and when the donor tubers were heavily infested with P. exigua var. foveata but were free of gangrene lesions. Increasing the proportion of donor to recipient tubers increased the percentage of infected wounds on recipients. Increased incidences of infection in recipient tubers also occurred after they had been passed over an elevator digger when it was lifting stocks of tubers heavily infested with P. exigua var. foveata. When spores of an E +ve isolate of P. exigua var. foveata were sprayed onto the webs of manned potato harvesters, tubers harvested immediately after developed gangrene rots from many of which the E +ve isolate was cultured. An E +ve isolate was also used to demonstrate the transfer of P. exigua var. foveata inoculum from tubers onto soil on riddles of a potato grader and from these soil-coated surfaces onto other tubers during grading.     

SOME FACTORS AFFECTING THE OCCURRENCE AND DEVELOPMENT IN POTATOES OF GANGRENE CAUSED BY PHOMA SOLANICOLA PRILL. & DELACR.

It is shown that gangrene is a soil- and haulm-borne disease and that infection of the tubers can take place through cuts, bruises, 'eyes' and lenticels. Low temperature and dry, dark storage were found to be most favourable to development of the disease, but rotting was more severe if the tubers had been kept at a higher temperature before infection took place. Storage at 21° C. for 10 days was sufficient to arrest rotting. Gangrene developed on all the potato varieties tested, but different varieties showed differences in susceptibility to the disease. No difference was observed in the susceptibility of early and late varieties but susceptibility increased with advancing maturity of the tubers.     

Studies on the lenticel development, surface microflora and infection by common scab (Streptomyces scabies) of potato tubers growing in wet and dry soils

The mechanisms of control of potato common scab (Streptomyces scabies) by irrigation were investigated in 5 years by studying the growth and surface microflora of young potato tubers developing in wet or dry soils. Direct examination of the tuber surface by light or scanning electron microscopy showed that the newly formed internodes near the apex, which carry stomata, had a very sparse microflora. In dry soil the older internodes, which carry lenticels at a susceptible stage, were increasingly colonised by actinomycete hyphae and discrete, although sometimes large, bacterial colonies. In wet soil, actinomycetes were rarely seen on tuber surfaces but bacteria were generally scattered over them, differences not always shown by isolation from periderm pieces onto water agar. When dry scab-infested soil was wetted, scabs did not subsequently develop on the two youngest tuber internodes (A-1, A-2), which may be an example of disease escape rather than inherent resistance of the stomata. The scab control achieved in wet soil was probably caused by some form of microbial antagonism, but whether through competition or antibiosis was not established.     

The inhibitory effects of rose powdery mildew infection on the oviposition behaviour and performance of beet armyworms

In this study, we investigated whether the oviposition behaviour and performance of the beet armyworm, Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), on the rose cultivar Rosa chinensis Jacq. (Rosaceae) were affected when the plants were infected by rose powdery mildew, Podosphaera pannosa (Wallr.: Fr.) de Bary (Erysiphales). The bioassays revealed that the moths significantly avoided ovipositing on mildew‐infected rose leaves when compared to healthy leaves. Pupal weights, emergence rates, and fecundity decreased when the caterpillars were fed mildewed rose leaves. Further laboratory bioassays aimed to elucidate the effects of two volatile headspace extracts (separately collected from healthy and mildewed rose plants) on the oviposition behaviour and performance of the moths. The moths clearly preferred to oviposit on healthy rose leaves that were not sprayed with additional volatiles rather than on healthy leaves sprayed with the volatile extracts from mildewed plants. The mean number of eggs laid on the former leaves was more than six times higher than that laid on the latter leaves. Olfactory bioassays demonstrated that ovipositing moths were significantly more attracted to volatiles emitted by healthy rose leaves than to those emitted by mildew‐infected leaves. Similar results were obtained when comparisons were made between the volatile extracts collected from healthy and mildewed rose plants. Thus, volatiles from mildew‐infected roses have a strong inhibitory effect against the moths. These results indicated that rose volatiles play a role in the oviposition behaviour of the moths, and that the volatiles induced by powdery mildew might be used for insect control.     

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.     

The behaviour of potato mop-top virus in soil, and evidence for its transmission by Spongospora subterranea (Wallr.) Lagerh.

Potato mop-top virus (PMTV) was best detected in field soils by air-drying them for more than a week before remoistening and growing seedlings of Nicotiana tabacum or N. debneyi for a 6–10 week period. Infection of N. tabacum was assessed by inoculating sap from roots and shoots to Chenopodium amaranticolor. Similar inoculations from N. debneyi were far less convenient for detecting PMTV than recording leaf symptoms, but slightly more efficient. Air-dry soil retained PMTV infectivity for 9 months, when passed through a 50 μ sieve or when diluted with 10³ but not 10⁴ parts of steamed soil. Tobacco seedlings were not infected when their roots were steeped in PMTV-containing tobacco sap. Infective soils contained Spongospora subterranea, spore balls of which resisted air-drying for more than a year and passed a 50 μ sieve. Roots of susceptible seedlings were infected with PMTV when exposed to spore balls of S. subterranea taken from powdery scabs on PMTV-infected potato tubers, or to suspensions obtained by steeping, in nutrient solution, roots infected with virus-carrying cultures of S. subterranea. Plants in several families were hosts of S. subterranea, but probabilities of infection when exposed to spore balls differed greatly between families and only species of Solanaceae were good hosts. The ten species infected with PMTV when grown in infective soil or when exposed to spore balls of S. subterranea taken from PMTV-infected potato tubers are all members of this family. PMTV seems to be carried internally in S. subterranea spore balls and survived in them for at least a year. PMTV was transmitted by S. subterranea to Arran Pilot potato, causing yellow blotches in some leaves and spraing in many tubers. However, when newly infected with PMTV in the field, not all Arran Pilot tubers developed spraing. Also, although many spraing-affected or symptomless but PMTV-infected tubers carried PMTV-containing spore balls of S. subterranea, powdery scabs were rarely found near the centres of the rings of primary spraing. PMTV became established in virus-free soil when PMTV-infected tubers carrying S. subterranea were planted as ‘seed’ but not when virus-free tubers bearing powdery scabs were used. 5. subterranea seems the main, and possibly the only, vector of PMTV in the soils examined. S. subterranea did not transmit potato aucuba mosaic virus from potato to N. debneyi or Capsicum annuum.     

Field and glasshouse experiments on the control of potato mop-top virus

Field observations during 3 yr on a stock of potato cv. Red Craigs Royal partially infected with potato mop-top virus (PMTV) confirmed that the virus was passed by an infected mother plant to only a proportion of its progeny tubers, and showed that in this cultivar symptomless plants gave rise only to symptomless progeny. The elimination of PMTV from stocks can therefore be greatly accelerated by removing symptom-bearing plants. Infected potato tubers were not freed from PMTV by treating them at 37 °C for up to 8 wk. Treating ‘seed’ tubers bearing powdery scabs that contain PMTV-carrying resting spores of Spongospora subterranea with formaldehyde or organo-mercurial fungicide greatly decreased PMTV establishment when the tubers were planted in previously uninfective soil, but fumigation with 2-aminobutane was ineffective. Decreasing the pH of infective soil to 5-0 by applying sulphur greatly decreased the infection of potato cv. Arran Pilot with PMTV and S. subterranea in field experiments, but this treatment did not eliminate either; when the pH of treated soil was raised the transmission of PMTV resumed. Treating infective soil with a range of fungicides greatly decreased the infection of Nicotiana debneyi bait seedlings in glasshouse experiments but only calomel at 75 kg/ha controlled spread of PMTV and 5. subterranea to potato in field experiments. In other field experiments, applying zinc frit, zinc sulphate or zinc oxide to infective soil greatly decreased the spread of both to potato. The amount of zinc required increased with increase in clay content of the soil. However, treatment with zinc compounds did not eliminate PMTV-carrying vectors from soil, and when treated soil was diluted with autoclaved soil many of the bait seedlings planted in the mixture became infected. The zinc frit was phytotoxic because of its boron content but zinc sulphate and zinc oxide caused little or no decrease in tuber yield. The zinc content of potato tubers was increased but not doubled in zinc-treated plots, and during the first year after treatment the zinc content of topsoil decreased greatly. The zinc content of ryegrass grown after potatoes was greater than of potato tubers but did not reach a level considered dangerous to livestock. Treatment of soil with sulphur, zinc oxide or calomel may be useful for small plots used in the early stages of propagation of virus-tested potato clones where there is risk of infection with PMTV.     

Control of contamination of potatoes with air-borne Erwinia carotovora by foliar application of copper oxychloride

In field experiments in 1983 and 1984, micropropagated plants of cv. Désirée planted in soil tested and found free of erwinias became contaminated with airborne erwinias, mostly E. carotovora pv. carotovora, in September when rain was more abundant, but not earlier. Control of contamination of progeny tubers harvested in September and October was achieved by spraying plants with copper oxychloride (‘Cuprokylt’; 5 g litre^-1) twice weekly from June or July but not from August nor by weekly or fortnightly treatments from June. Bactericidal activity of copper in the effective spray regimes was not expressed on the leaves or in the soil and contamination was similar in the sprayed and unsprayed treatments. In contrast, multiplication of erwinias in senescent leaves on the soil surface (leaf debris) was inhibited in the sprayed treatment but not in the unsprayed treatment and population numbers of c. 10³ cells g^-1 fresh weight were detected. It is suggested that fewer erwinias would be washed from leaf debris by rain in the sprayed than in the unsprayed treatment to contaminate the progeny 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.     

The control of common scab of potatoes by irrigation

Irrigation applied early during tuberization significantly decreased scab on tubers of Majestic, King Edward and Record, but not on Pentland Crown, a resistant cultivar, where tubers had little scab even from the driest soil. There was little benefit from keeping soils wetter than 0.6 in (15.24 mm) soil moisture deficit (S.M.D.) for more than 3 weeks after tuber initiation, and irrigation to 0.8 in (20.32 mm) S.M.D. gave economic control. Irrigating only after the ‘marble stage’ increased yield but the tubers were scabbed. Irrigation did not alter the rate eyes (nodes) separated from the apical bud but speeded the swelling of tubers.     

The effect of Erwinia carotovora subsp. atroseptica (blackleg) on potato plants. I. Growth and yield of different cultivars

The effect of planting seed tubers inoculated either near the stolon attachment (heel end), among the eyes at the apex (rose end) or mid-way along the tuber with Erwinia carotovora subsp. atroseptica, was assessed in terms of growth of the plant, disease symptoms and yield. Invariably rose-end inoculation had the greatest and heel-end the least effect in decreasing yield when compared with uninoculated plants. Cultivars Majestic and King Edward were the most susceptible, Pentland Crown showed some resistance to invasion of stems (blackleg) although plant vigour, expressed in terms of plant height and stem number was affected and Maris Piper was the most resistant.     

Healthier seed potatoes. II. Effects of sprouting and benomyl treatment of stem cutting and commercial stocks on growth, yield and disease

Potato seed tubers of seven cultivars derived from stem cuttings in 1965 (healthier seed) were compared with samples from commercial stocks in 1969–72 at two sites, one clay with flints soil (Rothamsted) and the other sandy loam soil (Woburn). Plant emergence, usually slower from non-sprouted seed, was unaffected by seed source or seed treatment with benomyl. Sprouted healthier seed produced more stems/plant than sprouted commercial seed. Healthier seed yielded 6% (Rothamsted 5%, Woburn 8%) more than commercial when seed was sprouted and 7% (Rothamsted 6%, Woburn 9%) when not sprouted. The increased yields of cvs Record (10% sprouted, 23% not sprouted), Majestic (8% sprouted, 11% not sprouted) and King Edward (5% not sprouted) mostly comprised small ware (44–57 cm) and chats (>44 cm), indicating an increase in tuber numbers. Commercial Pentland Crown seed yielded as much as healthier but the healthier produced less large ware (57–83 cm). Benomyl treatment of commercial seed, especially when not sprouted, and sometimes of healthier seed also decreased tuber size. Infection of stem bases and tuber eyes by Polyscytalum pustulans was less from healthier than commercial seed and was decreased by benomyl. Stem canker (Rhizoctonia solani) was also decreased by benomyl but fungicide treatment of seed did not greatly decrease the high incidence of R. solani hyphae on eyes of tubers at Woburn. Helminthosporium solani, equally prevalent on the produce of commercial and healthier seed, was decreased by benomyl. There was slightly more gangrene (Phoma exigua) on the produce of commercial than healthier seed.     

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.