The effects of length of the growth period and of nutrition upon potato-tuber susceptibility to dry rot (Fusarium caeruleum)

Standardized inoculations of the potato varieties Catriona and Doon Star over 6 years have shown that shortening the growth period, either by deferring planting or by removing the haulm prematurely, may considerably reduce susceptibility of the tubers to dry rot (Fusarium caeruleum). Although these treatments reduce dry-matter content of the tubers, no direct relationship between this factor and susceptibility was established. High pre-maturity susceptibility was confirmed at the time of flowering; as the haulm matured susceptibility decreased: when the haulm was dead tubers were resistant. Within a fortnight after any time of premature haulm removal, tubers became completely resistant and showed a decrease in sucrose content but not in content of reducing sugars. It is suggested that susceptibility in the immature tuber is closely related to the content of sucrose, which accumulates because of rapid translocation from the vigorously growing haulm. After harvest, tuber susceptibility slowly increased during storage, with an increase mainly in reducing sugars, but these two factors were not directly related. An application of 6 cwt/acre (753 kg/ha) of 12:12:18 NPK fertilizer to Catriona significantly reduced liability to infection. Additional nitrogen (6 cwt/acre ammonium sulphate) raised tuber susceptibility to that in a non-fertilized plot: when sulphate of ammonia was the only treatment given, the tubers were significantly more susceptible than those receiving complete fertilizer.     

DRY-ROT DISEASE OF THE POTATO

Inoculation of wounded tubers at intervals throughout the storage season showed that, after harvesting, resistance to infection by F. avenaceum was maintained for a longer period than resistance to F. caeruleum , although eventually the tubers became equally susceptible to both fungi. Tubers were less readily infected through clean-cut than through scarified wounds. Resistance to infection was greater when an interval elapsed between wounding and inoculation than when the wound was inoculated immediately; this was related to an increase in the intrinsic ability of the cells near the wound to resist infection rather than to suberization of the wounded surface or wound periderm formation.
Histological studies showed that F. caeruleum grew through the intercellular spaces and that the adjacent host cells remained alive, often for considerable periods, whereas F. avenaceum killed and penetrated the cells with which it came into contact. Restriction of incipient or established lesions caused by F. caeruleum and of incipent lesions caused by F. avenaceum was associated with suberin deposition on the host cell walls and in the intercellular spaces; in established lesions caused by F. avenaceum , restriction was attributed to an increase in the intrinsic resistance of the adjacent host cells, similar to that found near a wounded surface.     

Varietal susceptibility of potatoes to wireworm herbivory

1 Wireworms, the soil dwelling larvae of click beetles, Agriotes spp., have recently become a more prevalent pest of potatoes. The present study investigated whether potato varieties showed variable susceptibility to wireworm herbivory, and also tested whether increased susceptibility was associated with lower concentrations of glycoalkaloids. Twelve varieties were originally screened across a range of experimental scales, including laboratory and tunnel experiments and a large‐scale field trial involving over 2000 tubers. 2 In laboratory no‐choice tests, Maris Peer, Marfona and Rooster varieties were significantly more susceptible to wireworm attack, with 63% of tubers showing damage, compared with just 15% of the less susceptible varieties of King Edward, Nadine and Maris Piper. There was also greater tissue consumption and weight gain when wireworms were reared on the most susceptible varieties. 3 In choice tests, wireworms showed a significant preference for those varieties previously identified as being the most susceptible to wireworm herbivory (4.2 holes per tuber) compared with the least susceptible (1.2 holes per tuber). Similar patterns of susceptibility were seen in the field trial, although there was generally more variation in susceptibility. 4 In a tunnel experiment, Marfona and Maris Peer were significantly more susceptibile to wireworm attack (47% of tubers showing damage) compared with Nadine, King Edward and Maris Piper (27% of tubers showing damage). Although Nadine, in particular, had the highest glycoalkaloid concentrations (309.33 mg/kg) and lowest amounts of wireworm herbivory, the relationship between susceptibility and glycoalkaloid concentrations was weak, suggesting that this is unlikely to be the sole mechanism underpinning varietal susceptibility.     

THE USE OF TETRACHLORONITROBENZENE ISOMERS ON POTATOES

In laboratory and field experiments from 1952 to 1955, 2:3:5:6-tetrachloro-I-nitrobenzene (tecnazene) delayed the development of sprouts on treated potato tubers, and 2:3:4:6-TCNB retarded sprout development in two laboratory experiments. At the usual commercial rates of application the 2:3:4:5 isomer had no effect on sprout development and very little effect at higher rates. The crop from 2:3:5:6-TCNB-treated seed tubers usually contained a higher proportion of small tubers than that from untreated or 2:3:4:5-TCNB-treated seed. Marketable yield from 2:3:5:6-TCNB-treated seed was sometimes below that from untreated seed, but when this happened the yield from 2:3:4:5-TCNB-treated seed was higher.
Both isomers produced good control of dry rot, Fusarium caeruleum (Lib.) Sacc., on five varieties of potatoes, but did not produce a commercial control on Doon Star. To achieve control, the isomers had to be present at the time of infection. There was some evidence that the development of disease symptoms was delayed in treated, infected tubers. Since 2:3:4:5-tetrachloro-1-nitrobenzene has no sprout depressant action it merits further trial as a fungicidal treatment for seed potatoes.     

Control of Fusarium solani rot of potato tubers with fungicides

Tecnazene (up to 33 ppm) and dichloran (up to 500 ppm) had little effect on germination of spores or growth of Fusarium solani isolated from and causing a rot of potato tubers; they also did not decrease rotting when applied to wounds later inoculated with the pathogen. Benomyl and thiabendazole (up to 500 ppm) also had little effect on spore germination but did greatly decrease growth at 5 ppm. A pronounced pink coloration developed in cultures growing slowly in the presence of benomyl; a similar though less striking effect appeared in agar cultures containing thiabendazole. Benomyl suspended in water or diluted with Fuller's earth gave good control of rotting when applied to wounds inoculated later with F. solani. Still better control was obtained with thiabendazole; dusts containing 1% a.i. substantially decreased rots and those containing 10 % a.i. gave almost complete control when applied to wounds shortly before inoculation. Thiabendazole was also very effective when used 24 h after inoculation and a fair measure of control was obtained when it was applied 24 h later. Benomyl and thiabendazole placed on apparently intact surfaces of tubers caused tissue 5 mm deep to become toxic to F. solani 10 days later, and, unexpectedly, this tissue prevented spore germination. Fuller's earth alone substantially decreased rotting. The results obtained suggest that dusts containing thiabendazole have some promise for the control of Fusarium rots of potato tubers, especially of early crops.     

The resistance of potato varieties to Heterodera schachtii Schmidt, the potato-root eelworm

The potato varieties Epicure and Doon Star permitted the production of fewer and smaller cysts of Heterodera schachtii on their roots than the varieties Golden Wonder and Majestic, when all were grown under standard conditions. Under the stimulus of potato-root excretion the number of larvae which emerged from Epicure and Doon Star cysts was less than that from Majestic or Golden Wonder cysts of the same volume. Accepting Lapage's (1938) criteria of resistance to nematode infestation in animals and applying them to plants, it is concluded that this deleterious effect of the variety on the eelworm is probably of the nature of a degree of resistance to eelworm attack specific to each variety and is physiological and not anatomical in action. Of the varieties tested, Epicure is judged the most resistant, closely followed by Doon Star. Majestic is fairly susceptible but not so much so as Golden Wonder on which the eelworm forms large cysts that are extremely productive of larvae.     

Comparisons between Phytoalexin Concentrations and the Extent of Rotting of Potato Tubers Inoculated with Erwinia carotovora sub sp. atroseptica, E. carotovora sub sp. carotovora or E. chrysanthemi

Isolates of Erwinia carotovora sub sp. atroseptica, E. carotovora sub sp. carotovora and E. chrysanthemi were compared for their ability to rot potato tubers and to elicit the accumulation of phytoalexins. With three incubation temperatures (15, 22 and 30°C) tubers were more susceptible at higher temperatures and mean rot weights were inversely correlated with mean rishitin concentrations. However, within treatment variation between tubers in the extent of rotting was not correlated with rishitin concentration suggesting that other factor(s) also affect resistance. Phytoalexins, including 15-dihydrolubimin, 3-hydroxylubimin, phytuberin and solavetivone, were measured by high pressure liquid chromatography using a silica column.     

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.     

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.     

Fungal Assciations: I. Synergistic Relation between Rhizoctonia solani Kuhn and Fusarium solani Snyder and Hansen in causing a Potato Tuber Rot

Rhizoctonia solani, Fusarium solani, and Phoma foveata werechosen for the study of disease caused by these fungi in differentcombinations in potato tubers. An initial Rhizoctonia infection,when followed by a Fusarium infection, gave an extensive rottingwith external pimple-like formations in some cases. This typeof rotting could not be brought about by individual infectionswith either of the two fungi, or jointly by them when Fusariumwas inoculated first. Microscopic observations of infected matureand young potato tubers showed that Rhizoctonia grew intracellularlywhen infected alone, whereas it grew inter- as well as intra-cellularlyin the successive double infection. Fusarium formed more haustorium-likestructures when inoculated alone that when it followed Rhizoctonia.The length of these structures in the double infection was greaterin mature than in young tubers. Atmospheric humidity affectedthe amount of rotting, the shape and colour of the rot, andthe morphology of the fungus in the tissue.     

Dry rot of potato Fusarium caeruleum (Lib.) Sacc.; effect of planting infected and contaminated sets on plant establishment

The effect on plant establishment of planting infected and contaminated sets of susceptible varieties has been studied in field trials from 1942 to 1945.
Healthy whole sets, and whole sets inoculated internally 3 days before planting, yielded good stands, whilst whole sets with obvious dry-rot lesions present at planting time produced gappy crops.
Healthy cut sets gave satisfactory stands except in one test with Doon Star, whilst sets cut with a contaminated knife at planting time yielded only a few misses in 1943 but many in the unfavourable 1944 season and in 1945.     

Effects of Temperature on Viability of Pathogenic Fusarium Species

The viability of three species of Fusarium pathogenic to winter cereals was differently affected by temperature. F. culmorum survived in vitro in soil for 4 months at 8, 20 and 30 °C, the number of colony forming units (CFU) recorded at these temperatures after incubation being 125, 185 and 624 % respectively when compared with the number present at the beginning of the experiment. F. avenaceum and F. heterosporum barely survived at 8 °C after incubation for 4 months but at 20 °C the numbers of CFU were 314 and 380 % respectively, while at 30 °C the corresponding number tor each of these species was 200 %.
At natural soil temperatures in winter, the number of CFU of F. culmorum after 4.5 months decreased to 60, 70 and 87 % of the number present at the beginning of this experiment when infested soil was buried at depths of 7–10, 15—20 and 30 cm respectively. In the upper soil layer where winter temperatures were lowest, survival of F. avenaceum and F. heterosporum did not occur. Survival rates for these two species at a depth of 15—20 cm were 20 and 5 % respectively. In the deepest layer of soil (30 cm) survival of these two species was 30 % for F. avenaceum and 10 % for F. heterosporum of the number present at the beginning of this experiment.
These results demonstrate that during the early spring stages of growth of winter wheat, inoculum of F. culmorum , unlike that of F. avenaceum and F. heterosporum , is also a source of infection in the top soil layer in our climatic conditions.     

Fungus diseases on potato seed tubers planted in England and Wales, 1963-76

During 1963-76 samples of potato tubers from commercial seed stocks of cvs King Edward (14 yr), Pentland Crown (9 yr), Majestic (7 yr), Pentland Dell (3 yr), Record and Arran Pilot (2 yr) were received from farms in England and Wales. Fifty tubers from each sample were examined macroscopically for fungus diseases and eyes were excised from a 20-tuber sub-sample, incubated and examined for pathogenic fungi; 50 tubers were stored on trays to sprout and examined for diseases and sprouting in May and in most years samples of 50 tubers were wounded by dropping onto expanded metal, stored at 5° C and examined for gangrene and dry rot after 12 wk. Amounts of disease varied between years and during 14 yr black scurf and powdery scab on King Edward tended to increase and skin spot and late blight decrease. On average 44% of King Edward tubers were affected with skin spot, 25% with black scurf and 16% with powdery scab. Gangrene affected 5% of tubers and 97% of the isolates from rots were identified as Phoma exigua var. foveata. Wounding tubers increased the incidence of gangrene three-fold. During 1963-69 late blight affected 2% of King Edward tubers but fewer in later years and in other cultivars. Majestic had most common scab (44% tubers) and Arran Pilot most dry rot (9% tubers) and this disease was increased by wounding tubers. Conidiophores of Helminthosporium solani (silver scurf) were more common on excised eyes of Pentland Crown, Record and Arran Pilot than of other cultivars, and isolations from verticillate conidiophores that developed on the side of incubated eye plugs of King Edward and Majestic stocks gave pure cultures of Verticillium tricorpus (78%), V. nigrescens (9%) and V. nubilum (3%). Proportions of tubers with different diseases were affected by their country of origin; Scottish seed had most skin spot and gangrene, Irish seed most powdery scab and English seed most common scab, late blight and H. solani. There was also evidence of differing disease incidence in seed from different geographical areas in Scotland and England. Up to half the King Edward and Pentland Crown stocks examined in 1975 and 1976 were derived from stem cuttings and average amounts of diseased tubers were similar to those in stocks not derived from stem cuttings. Annual and cultivar differences in disease incidence and effects of date of receipt of seed on farms are discussed.     

Hydrolytic Activities of Fusarium solani and Fusarium solani f. sp. eumartii Associated with the Infection Process of Potato Tubers

The development of dry rot caused by Fusarium solani f. sp. eumartii was evaluated in susceptible (Huinkul) and resistant (Spunta) potato cultivars. Fungal proteolytic and polygalacturanase activities were measured at different days postinoculation either with the pathogenic F. solani f. sp. eumartii, isolate 3122 or with the non‐pathogenic F. solani, isolate 1042. After inoculation with the pathogenic fungus, proteolytic and polygalaturonase activities were higher in the susceptible than in the resistant cultivar. In addition, we found a correlation between the levels of proteolytic activity detected in the intercellular washing fluids with the size of the lesion area caused by F. solani f. sp. eumartii in Huinkul tubers. The action of the proteolytic activity over cell wall proteins of both potato cultivars was assayed. An extracellular potato protein with homology to proteinase inhibitors of the Kunitz family was identified as a substrate of the proteolytic activity in the susceptible cultivar. A microscopic study revealed differences between the potato genotypes in the rate of response to infection by F. solani f. sp. eumartii. In addition, the cell wall alteration caused by F. solani f. sp. eumartii in cortical cells of susceptible tubers was evaluated. The data with respect to the correlation between the course of cyto‐ and biochemical events of the two host–pathogen interactions were discussed.     

Effects of potato leafroll virus on the crop processes leading to tuber yield in potato cultivars which differ in tolerance of infection

The effects of infection with potato leafroll virus (PLRV) on the four crop processes leading to tuber fresh weight yield were examined in field plots of four cultivars (Montana, Pentland Crown, Maris Piper and King Edward) differing in tolerance to infection. Averaged across cultivars, infection decreased yield by 50%. This decrease was equally due to less light (total solar radiation) being intercepted, a lower efficiency with which intercepted light was converted into dry weight, and a smaller proportion of dry weight being partitioned to tubers. Dry matter content of the tubers was also diminished but to a lesser extent. The main difference between the cultivars in their response to infection was in the partitioning of dry weight. In Montana and Pentland Crown, harvest index was decreased by 15% in infected plants, whereas in the less tolerant cultivars, Maris Piper and King Edward, it was decreased by 25%. The decline in photosynthetic performance of Montana, a cultivar with slightly earlier maturity than the other three, was delayed in PLRV-infected plants. Effects on number of daughter tubers essentially reflected those on yield to the extent that average tuber weight did not change in Maris Piper, was one third less in King Edward, and the change was intermediate in Montana and Pentland Crown.     

A method for assessing the field susceptibility of potato cultivars to blackleg (Erwinia carotovora subsp. atroseptica)

In experiments to develop a method for assessing the field susceptibility of potato cultivars to blackleg (Erwinia carotovora subsp. atroseptica) seed tubers were stab-inoculated near the stolon (attachment end), with a suspension of the bacterium, or with water, before planting. Disease symptoms were recorded in three years (1980–1982) and plant growth and yield in 1982. Estima and Maris Bard were the most susceptible cultivars with many plants failing to emerge and most of those that did showing disease symptoms. Pentland Crown was the most resistant: few plants failed to emerge and few showed blackleg. Nevertheless compared with water-inoculated plants bacterial inoculation of the seed tubers of this cultivar caused loss of yield and differences in tuber size distribution. Cara, Wilja and King Edward showed intermediate reactions.     

Stem canker (Rhizoctonia solani) on five early and seven maincrop potato cultivars I. Infection of shoots, stolons and tubers

In 1983 and 1984, potato seed tubers of five early and seven maincrop cultivars were inoculated with cultures of Rhizoctonia soluni during planting to simulate severe seed infection. Shoot and stolon infection was assessed in June-August and black scurf on tubers recorded after harvest in October. Almost all shoots of all cultivars had stem canker in both years and disease on shoots, stolons and tubers was more severe in 1984 than in 1983. In 1983 similar amounts of disease developed on all early cultivars and between 11% (Ulster Sceptre) and 32% (Maris Peer) shoots were pruned off. Maris Peer had a stem canker score lower than other cultivars in 1984 but more than half the shoots were pruned off. Shoot pruning on Estima, Ulster Prince and Ulster Sceptre was more common on plants from sprouted than non-sprouted seed. Between 30 and 50% of stolons were pruned off. After harvest in 1985, black scurf was least prevalent on Arran Comet and Maris Peer tubers and in 1984 on Arran Comet and Estima tubers from non-sprouted seed. Of the maincrop cultivars, King Edward plants from sprouted seed had many shoots pruned off in both years. Shoot pruning was also prevalent on Maris Piper and Pentland Squire plants from non-sprouted seed. Record had fewest pruned shoots and stolons and the lowest stem canker score. The disease was more severe on Pentland Crown and Maris Piper plants from non-sprouted than sprouted seed. Black scurf was most common on Cara and King Edward tubers in 1983 and on King Edward and Record tubers in 1984. In both years few shoots but many tubers were infected on plants from non-inoculated seed and the significance of this is discussed.     

Factors affecting the development of spraing in potato tubers infected with potato mop-top virus

A larger proportion of tubers of Arran Pilot potato growing at the surface of soil infested with potato mop-top virus (PMTV) showed spraing symptoms (brown rings) at harvest than of tubers from below the surface. Infected tubers with or without spraing developed a spraing ring when stored in darkness, first for 1–2 wk at 18 d?C and then for 1–2 wk at any constant temperature between 5 and 13 d?C. Only a faint surface ring developed when either of these periods was decreased to 1 day; 4-day periods were needed to induce distinct symptoms. Internal tuber symptoms developed more slowly than surface symptoms, and their formation was favoured by cutting the tubers in half. Additional pigmented surface rings were produced outside the first ring by successive cycles of treatment at 18 and 9d?. Spraing did not develop when the first stage of treatment was at 22–25d?, when the tubers were kept first at 10d? and then at 5d?, when the treatment at 5–13d? preceded that at 18d?, or when the tubers were kept at constant temperatures ranging from 5 to 25d?. When tubers of six potato varieties were grown in PMTV-infested soil and then stored at temperatures designed to induce symptoms, the varieties known to be the most susceptible in the field were those which had the greatest tendency to develop spraing during storage. When infected tubers were exposed to light, typical spraing symptoms were not induced, but greening of the tuber surface was much delayed in localized ring-shaped areas, so that pale weals appeared. Spraing symptoms were produced, in favourable conditions, by the reaction of cells at the periphery of the PMTV-invaded zone. Internal spraing did not prevent PMTV invading tissue outside the brown arcs; its rate of spread was about 10 μm/h at 14–18d?.     

FIELD EXPERIMENTS ON POTATO SKIN-SPOT DISEASE CAUSED BY OOSPORA PUSTULANS OWEN & WAKEF

In experiments on skin-spot disease of potatoes, caused by Oospora pustulans Owen & Wakef., Kerr's Pink proved the most susceptible of twenty-four commercial varieties; Arran Banner, Majestic and King Edward were highly susceptible; Home Guard and Golden Wonder were highly resistant.
Thymol and tetrachlornitrobenzene, applied as dusts to pits of potatoes at lifting, did not give satisfactory control of the disease. The efficacy of an organo-mercury dip treatment at lifting time was confirmed.
The incidence of the disease was considerably decreased by storing tubers in boxes instead of in pits, by digging the crop about a month before normal harvest, or by cutting the haulms at this time and digging at the normal time.
Ware tubers showed a significantly higher infection than seed tubers of the same crop, indicating that susceptibility of tubers increases towards maturity.
The difference in eye infection between varieties may not be related to that of general superficial infection. It is suggested that, in future, the assessment of varietal susceptibility should be based upon the degree of eye infection, which is in practice the measure of the economic importance of the disease. Tubers having pustules at or near all the eyes gave markedly reduced sprout emergence and plant stand.     

DRY-ROT DISEASE OF THE POTATO

A laboratory method is described in which a modified hypodermic syringe is used to inoculate potato tubers with Fusarium caeruleum. This injection method gives consistent results and permits reliable assessments of factors such as varietal susceptibility.
The occurrence of arrested lesions is noted.
Suspensions of very high spore concentration or of spores already germinated did not greatly increase the amount of dry rot above that given by the standard too spores per inoculation, provided that spore germination was adequate.
The size of tuber and site of inoculation were found to have a considerable effect upon the results of inoculation experiments, large tubers being more susceptible than small ones and the heel end being more susceptible than the rose end. The necessity for uniformity of material used in inoculation experiments is emphasized.