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.     

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.     

Stem canker (Rhizoctonia solani) on five early and seven maincrop potato cultivars. II Effects on growth and yield

In 1983 and 1984, potato seed tubers of five early and seven maincrop cultivars were inoculated with cultures of Rhizoctonia solani during planting in field experiments to simulate severe seed infection. The size of foliage was assessed during June-August and tuber yields recorded during growth and at harvest in October. Stem canker delayed shoot emergence, decreased the number and length of stems and caused increased variation in stem length; these effects were greatest with Maris Peer and Arran Comet (early cultivars) and King Edward and Pentland Squire (maincrop cultivars). Total weight of foliage was decreased, especially with earlies, dry matter of stems increased and the proportion of foliage on lateral stems increased. With the early cultivars, tuber yield from sprouted Maris Peer seed 11 wk after planting in 1983 was decreased by 24%, and 13 wk after planting in 1984 yields were decreased by 42% (Maris Peer), 40% (Ulster Sceptre), 34% (Estima), 30% (Arran Comet) and 17% (Ulster Prince) with sprouted seed and by 20, 29, 53, 39 and 28% respectively with non-sprouted seed. Decrease in total yield at harvest in October averaged 13% with sprouted seed and 10% with non-sprouted seed. In all cultivars the weight of small tubers was decreased and with Estima the weight of large tubers was increased. Tuber bulking was also delayed with all maincrop cultivars and at harvest yields from sprouted King Edward seed were decreased by 13% in 1983 and by 16% (sprouted seed) and 23% (non-sprouted seed) in 1984; yields of Pentland Squire were decreased by 5, 16 and 21% respectively. Yield losses with other cultivars ranged from 5–13% with sprouted seed and 0–16% with non-sprouted seed. The yields of small tubers were decreased with all cultivars and yields of large tubers were increased with Pentland Squire, Pentland Crown and Cara.     

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.     

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.     

Observations on blight (Phytophthora infestans) and resistant potatoes at Toluca, Mexico

A late blight epidemic studied at Toluca, Mexico, in 1962 may have started from stems infected at soil level by soil-borne Phytophthora infestans. Its severity was demonstrated by the large differences in the rate foliage was destroyed and in yield of tubers between fungicide-sprayed and unsprayed susceptible and resistant cultivars. The foliage resistances of some Mexican and European cultivars were compared using conventional blight keys and recording the destruction of marked leaves. Cultivars reacted in four ways: (1) Up-to-Date and Alpha leaves were infected and killed soon after plants emerged; (2) Bertita, Conchita and Florita abscissed many infected lower-canopy leaves, and many infections on upper leaves aborted; (3) Elenita leaves had a few lesions in which the fungus grew slowly but remained alive; (4) Greta showed no infection until the potato plants met between rows but then infections developed rapidly and the foliage soon died. Mexican cultivars, except for Elenita, had few blighted tubers; susceptible European cultivars were killed before many tubers formed. Most spores were released during the morning, as in Europe, and leaf infection seemed associated with days with rain when much of the night remained humid. Cool nights lengthened incubation and generation times.     

Susceptibility of potato tubers to infection by Phytophthora infestans

Phytophthora infestans infects King Edward potato tubers more readily through inoculated eyes than through lenticels, but more lenticels than eyes became infected when whole tubers were sprayed with inoculum. The resistance of lenticels but not of eyes increased as tubers aged. The spores did not infect through intact periderm. The likelihood of tubers on plants grown in pots becoming infected by sporangial suspension poured on to the soil increased the nearer the tubers were to the soil surface, the stem, or the side of the pot. Naturally infected tubers, and those sprayed with sporangial suspension, had most eyes infected at the rose end, and most lenticels infected on the middle region of the tuber. Of naturally infected tubers, on which the site of infection could be identified, most were infected through eyes at the rose end.     

Effects of black scurf (Rhizoctonia solani) on potatoes

King Edward and Majestic seed tubers, selected as ‘clean’ (macroscopically symptomless), moderate and severe according to the extent of black scurf, were planted in field experiments at Rothamsted between 1964 and 1968. Seed infection sometimes delayed plants emerging but did not affect final plant populations. Crops from severely diseased seed yielded, on average, 7% less than ‘clean’ tubers (King Edward 6–8% less and Majestic 0–20% less). Seed infection affected tuber size distribution; compared with ‘clean’ seed, severely infected King Edward seed yielded slightly more chats (< 1 ½ in, 3.8 cm) and 1.5 ton/acre (3.8 t/ha) less large tubers (2 ¼–3 ¼ in, 5.7–8.3 cm). The effects were similar with Majestic although differences were smaller. However, total yields from diseased stocks (unselected) seldom differed significantly from the ‘clean’ tubers selected from them. Crops from moderately and severely diseased seed had more Corticium on stems and black scurf on tubers and usually less Oospora pustulans than from ‘clean’ seed.     

ROGUING POTATO CROPS FOR VIRUS DISEASES

Removing virus-infected plants from plots of Majestic potatoes at Rothamsted on 2 July 1947 did not reduce the spread of leaf roll but reduced rugose mosaic (potato virus Y) to about one-fifth of that in plots rogued on 21 July or left unrogued. Roguing Arran Pilot potatoes on 16 June or 2 July reduced leaf roll to five-sixths of that in unrogued plots; roguing on 16 June reduced rugose mosaic to about half that in plots rogued on 2 July, and about a quarter of that in unrogued plots. Lifting Arran Pilot potatoes in mid-August reduced virus diseases to about two-thirds.
Roguing flattened the gradient (decrease in percentage plants diseased with increasing distance from the source of infection) with rugose mosaic, but had little effect with leaf roll. Evidently any plants prevented by roguing from contracting virus Y were near the initially infected plants.
In 1948, Majestic and King Edward potatoes at three places were rogued during 22–24 June and tubers were dug during 28–30 July and again at the end of the season. Leaf roll spread more in Majestic than in King Edward, and rugose mosaic spread more in King Edward. Roguing reduced the spread of both by about one-fifth at Rothamsted, but had no effect at Sutton Bonington. At Bretton, in the Derbyshire hills, roguing had no effect on leaf roll, but prevented the spread of rugose mosaic.
The small benefit occasionally achieved by roguing in the ware-growing districts of England does not make the practice economically worth while.     

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.     

Black scurf and stem canker of potato (Corticium Solani Bourd. & Galz.)

Black scurf and stem canker of the potato was investigated in field trials on infected soil at Warburton, Cheshire, to study the effect of planting clean and contaminated seed, and the relation between the dates of planting and lifting and the contamination of the crop tubers.
Black scurf was prevalent on crops grown from clean seed but was more severe on crops raised from contaminated seed. Under conditions favourable for the disease the yields from clean and contaminated seed were satisfactory and were not significantly different. Young shoots of plants from contaminated seed were severely attacked and tuber formation was checked; these results were not obtained on clean seed plots.
More black scurf occurred on late-dug crops; on early-dug plots the disease, although almost absent in 1941, was prevalent in 1942. Late planting failed to reduce appreciably the amount of black scurf.
Variety trials included Arran Banner, Kerr's Pink (two vigorous varieties), King Edward and Majestic (two less vigorous varieties). All were heavily infected and each recovered well from an attack on the young shoots.
Inoculation of the seed at planting time did not affect the results; in all trials, misses and wilted shoots caused by Corticium were rare and there was no relation between the yield and the amount of black scurf on the crop tubers.
The results indicate that the disease causes little, if any, loss under farm conditions.     

Potato blight control in east and south-east Scotland 1959–68

Routine protective spray application to commercial Majestic potato crops on two farms in south-east Scotland for 9 and 10 years respectively gave highly significant average increases in total yield of 1.2 and 2.1 tons per acre (3.0 and 5.3 tonnes per ha) respectively in rows not affected by tractor wheels. In 6 and 7 years respectively the increases were more than 1 ton per acre. These increases were composed almost entirely of ware tubers. There was no average difference in yield whether the first spray application was made when the haulm closed in the rows or after the first general forecast based on the Beaumont period was announced. The average total yield reduction in the four rows affected by tractor wheels tended to offset the benefit given by spray application, and, where an eight-row spray boom was employed, halved the potential overall yield increase. Reduction was again confined to the ware fraction. Similar experiments with cv. King Edward over a shorter period in eastern Scotland generally showed little consistent effect on yield, possibly partly because of the late appearance of blight in some of these crops. Considerable yield gains were obtained in Majestic crops even when the disease first appeared in mid-August in some seasons which could not be classified as ‘blight years’. Over the 10 years of the investigation, there were 5 blight years as defined by Large (1959) compared with the 1 in 10 which Large mentioned for the north of England. Spread of blight from local sources, such as discard heaps outside the crop and before the occurrence of the field epidemic phase, was found to interfere with adequate timing of spray applications and with the effectiveness of the general forecast based on the Beaumont period. Infection of Majestic tubers was generally low but at one Majestic site and in the King Edward series the overall average tuber infection was significantly decreased by spraying. Tuber blight was shown to vary inversely with the size of tubers of Majestic as well as of King Edward.     

SOME OBSERVATIONS ON DAMAGE TO POTATOES BY SLUGS

The garden slug, Arion hortensis Fér., and the keeled slugs, Milax spp., are the slugs mainly responsible for primary damage to potato tubers in the south-west of England. A variety trial clearly demonstrated the existence of significant differences in susceptibility to slug attack between the three varieties tested. Arran Banner was the most damaged variety, Arran Peak the least, while Majestic occupied an intermediate position. Varietal differences in the amount and time of damage to tubers were associated with differences in the dates of maturity, and little damage occurred prior to the maturing of the tuber. The results confirm the need for prompt lifting of the potato crop on maturity, in order to avoid damage by slugs.     

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.     

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.     

The induction of top-roll symptoms on potato plants by the aphid Macrosiphum euphorbiae

In the field, caged potato plants of King Edward and Majestic cultivars infested with the potato aphid Macrosiphum euphorbiae developed top-roll symptoms, the proportion of affected plants increasing with the size and persistence of the aphid population. Yield of tubers from plots in which 90% of the plants had top-roll symptoms was 40% less than that from control plots; yield of saleable ware was even less. Foliage produced after the aphids had been killed was symptomless even when it arose from the axil of an affected leaf. Caged field plants treated with phorate granules to prevent aphid attack did not develop top-roll. Prolonged infestation of Pentland Crown, Majestic and King Edward plants by M. euphorbiae in a glasshouse induced rolling of upper leaves similar to top-roll of field plants. Experimental results suggest that rolling was directly attributable to heavy attack by M. euphorbiae, not to an aphid-transmitted pathogen.     

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.     

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.     

A comparison of Fusarium avenaceum and Fusarium caeruleum as causes of wastage in stored potato tubers

Fusarium avenaceum is reported for the first time as a cause of rotting of potato tubers in Britain. The progress of rotting in tubers infected with F. avenaceum has been compared with dry rot due to F. caeruleum in the laboratory, clamp and potato store. Of the four varieties, Majestic, King Edward, Doon Star and Arran Pilot, tested for susceptibility, King Edward was the most susceptible to F. avenaceum and Doon Star to F. caeruleum.
Optimum temperatures for growth on potato-dextrose agar were 20-25 C. for F. avenaceum and 20 C. for F. caeruleum ; maximum temperatures were > 30 and 30 C. respectively. For infection of wounded potato tubers, cardinal temperatures for F. avenaceum were similar to those on agar, but for F. caeruleum the optimum was 15 C. and the maximum 25 C. The optimum temperature for rotting tended, with both species, to be higher in the more susceptible potato varieties. At low temperatures F. caeruleum caused quicker rotting than did F. avenaceum , even though its rate of growth on agar was scarcely more than half that of the latter.
High humidity favoured rotting especially by F. avenaceum; F. caeruleum was more tolerant of relatively low humidity. Both species caused quicker rotting in the clamp than in store, even though there was no appreciable difference in mean temperature between the clamp and the store. This was attributed to the higher atmospheric humidity in the clamp.     

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.