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.     

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.     

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.     

Sites of contamination and numbers of Erwinia carotovora present in stored seed potato stocks in Scotland

Forty-eight bulk and crate stored commercial seed potato stocks from the east of Scotland were examined for contamination by Erwinia carotovora in 1967 and 1968. Both E. carotovora var. carotovora and E. carotovora var. atroseptica were present, the latter, the blackleg pathogen, being detected four times more frequently. Most contamination was located on the tuber surface and in the lenticels; it was rare in the vascular ring and the organism was never detected in the cortex. The level of contamination of lenticels was only slightly affected after winter storage, which, however, substantially reduced surface contamination. More than 10³ cells were detected on the surface of tubers in over half the stocks and in half the stocks each lenticel sampled contained more than 10² bacteria.     

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.     

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.     

Effect of Time of Inoculation with Xanthomonas campestris pv. mangiferaeindicae on Mango Fruits Susceptibility Epiphytic Survival of X. c. pv. mangiferaeindicae on Mango Fruits in Relation to Disease Development

Susceptibility of mango fruits to bacterial black spot disease was related to the stage of fruit development and to climatic factors (rainfall and temperature). The highest percentages of infected fruits occurred when inoculations were performed during the month just before harvest. Wounds and lenticels are effective entry sites for the bacterium. The ability of lenticels to take up bacterial inoculum was estimated using safranin. The susceptibility of lenticels to entry of the pathogen was directly related to the fruit age. X. c. pv. mangiferaeindicae can survive epiphytically on immature mango fruits. Bacterial populations around 10⁵ c.f.u./fruit are commonly detected. Epiphytic X. c. pv. mangiferaeindicae were not detected on symptomless mature fruits. Those resident populations are an important source of inoculum for fruit infection. The number of bacterial spots occurring on the fruits is directly related to epiphytic populations.     

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.     

Tuber surface microorganisms influence the susceptibility of potato tubers to late blight

Potato tubers (cvs Cara and Bintje) were grown in compost in a glasshouse and immature tubers harvested 57, 68 and 78 days after planting. Two moisture levels were imposed after the first harvest by disconnecting the water supply to one of the treatments and allowing the soil in that treatment to dry naturally. Tubers from wetter compost (59.4% moisture holding capacity) were more resistant to Phytophthora infestans than those from drier compost (14.7% moisture holding capacity) 78 days after planting. The potential causes of this difference were investigated. Aqueous extracts of wet compost did not inhibit the growth of P. infestans. The susceptibility of the internal tuber tissue, from which the periderm had been removed, was different to whole tuber susceptibility. The internal tissue of tubers from wet compost was more susceptible (cv. Cara), or as susceptible (cv. Bintje) as that of tubers from dry compost 78 days after planting. Fungi were isolated from the surface of whole tubers and there were no differences between the populations of potentially antagonistic fungal genera on tubers from wet and dry compost. As the experiment progressed, the number of bacteria per gram fresh weight on tubers grown in wet compost increased, whereas that on tubers from drier compost decreased (cv. Bintje) or remained similar (cv. Cara). There were significantly (P= 0.008) more bacteria on the surface of tubers from wet compost 78 days after planting. When P. infestans was co-cultured in Petri dishes with randomly selected tuber surface bacteria, some isolates (≤ 16.7%) inhibited the growth of the fungus. The percentage of the total bacterial population that was antagonistic to P. infestans was not significantly affected by soil moisture level (P= 0.368). The greater numbers of bacteria, of which a proportion are antagonistic to P. infestans, on the surface of tubers grown in wet compost may account for the greater resistance to tuber blight in that instance.     

Age-specific bioassays and fecundity of Phthorimaea operculella (Lepidoptera: Gelechiidae) reared on cry1Ac -transgenic potato plants

Variation in the susceptibility of lepidopterous pest larvae of different ages to transgenic crops and the potential for survivors to reproduce could have important consequences for the development of resistance in such pests. Experiments were undertaken in the laboratory to determine if larvae of the potato tuber moth, Phthorimaea operculella, of different ages (0 (< 1 day old), 3, 5, 7 days) varied in their susceptibility to cry1Ac9–transgenic potato (Solanum tuberosum) foliage grown in the glasshouse or field. The survival and fecundity of larvae reared on transgenic tubers was also determined in the laboratory. There were no apparent differences in susceptibility of larvae of different ages to transgenic foliage. Larvae fed glasshouse or field‐grown non‐transgenic foliage had significantly larger relative growth indices and more larvae pupated, than those fed transgenic foliage, regardless of larval age. Eggs from a laboratory colony were placed on transgenic or non‐transgenic tubers to measure survival and fecundity. Between 6% and 15% of eggs placed on transgenic tubers developed into pupae for three of the four transgenic potato lines tested. On one transgenic line, only six adults emerged from 1300 eggs. In contrast, between 71% and 97% of the eggs placed on non‐transgenic tubers developed into pupae. Male and female pupae from transgenic lines weighed less than those from non‐transgenic lines. The fecundity of females from two of four transgenic lines was lower than from the non‐transgenic parent cultivar. Although larvae of different ages did not exhibit any overall age‐dependent pattern of increasing or decreasing susceptibility to transgenic foliage of glasshouse or field‐grown plants, the ability of larvae to survive and reproduce on transgenic tubers suggests this pest has the ability to evolve resistance to the transgenic plants used in the present study.     

The Role of Polyphenol Oxidase and Peroxidase in Potato Tuber Resistance to Soft Rot Caused by Erwinia carotovora

Tubers from somatic hybrids produced by protoplast fusion between Solanum brevidens, a diploid, non-tuber-bearing wild species, and a tetraploid S. tuberosum showed resistance to decay caused by soft rot Erwinia. Tubers of the S. tuberosum fusion parent and potato cultivar Russet Burbank are susceptible to bacterial soft rot. Tubers of somatic hybrids indicated higher levels of activities of polyphenol oxidase and peroxidase than tubers of the parental line of S. tuberosum and cultivar Russet Burbank. This is true for intact tubers and also for injured or inoculated tubers. Polish commercial potato cultivars indicated a higher susceptibility to soft rot than somatic hybrids. However, there were some differences in susceptibility to soft rot between Polish commercial potato cultivars, only slight differences were observed in the activities of the polyphenol oxidase and peroxidase between Polish cultivars. A relation between soft rot resistance and the activity of each enzyme was not found for intact, injured or inoculated tissue of commercial cultivars. On the contrary, the activities of both enzymes were significantly higher in the periderm than in the medullary tissue of somatic hybrids, the parental line and the commercial cultivars.     

<Emphasis Type="Italic">Strboh A</Emphasis> homologue of NADPH oxidase regulates wound-induced oxidative burst and facilitates wound-healing in potato tubers

During 30-months of storage at 4°C, potato (Solanum tuberosum L.) tubers progressively lose the ability to produce superoxide in response to wounding, resist microbial infection, and develop a suberized wound periderm. Using differentially aged tubers, we demonstrate that Strboh A is responsible for the wound-induced oxidative burst in potato and aging attenuates its expression. In vivo superoxide production and NADPH oxidase (NOX) activity from 1-month-old tubers increased to a maximum 18–24 h after wounding and then decreased to barely detectable levels by 72 h. Wounding also induced a 68% increase in microsomal protein within 18 h. These wound-induced responses were lost over a 25- to 30-month storage period. Superoxide production and NOX activity were inhibited by diphenylene iodonium chloride, a specific inhibitor of NOX, which in turn effectively inhibited wound-healing and increased susceptibility to microbial infection and decay in 1-month-old tubers. Wound-induced superoxide production was also inhibited by EGTA-mediated destabilization of membranes. The ability to restore superoxide production to EGTA-treated tissue with Ca⁺² declined with advancing tuber age, likely a consequence of age-related changes in membrane architecture. Of the five homologues of NOX (Strboh A-D and F), wounding induced the expression of Strboh A in 6-month-old tubers but this response was absent in tubers stored for 25–30 months. Strboh A thus mediates the initial burst of superoxide in response to wounding of potato tubers; loss of its expression increases the susceptibility to microbial infection and contributes to the age-induced loss of wound-healing ability.     

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.     

The significance of tuber damage and inoculum concentration of Phoma exigua var. foveata in the development of gangrene in stored potato tubers

The susceptibility to gangrene infection of wounds of various shapes and depths on potato tubers was studied by inflicting wounds using differently-shaped brass teeth and rods of different diameters. Inoculating wounds with spore suspensions or damaging tubers which had been previously contaminated with Phoma exigua var. foveata or which had been recently lifted from plots of field experiments showed that wounds in which tissue was crushed were most susceptible to infection. Over a wide range of inoculum concentrations and in experiments using several different cultivars the incidence of infection of any wound type was compared to that of the standard severe cut and crush wound. Using a probit transformation a linear relationship was established, the slope of the line indicating the relative susceptibility of the wound. In 1977 and 1978, crops of cv. Pentland Dell were surveyed for damage incidence, inoculum and inoculum potential on arrival at a commercial bulk store. Nets of tubers buried among the tuber bulk were recovered after storage and gangrene incidence compared with damage and inoculum assessments. Inoculum potential and incidence of severe damage both influenced disease development but damage incidence was of greater importance, showing that priority should be given to decreasing damage and to curing to promote rapid wound healing in endeavours to control the disease.     

The systematic significance of surface features of theBalanophora tuber (Balanophoraceae)

The surface ofBalanophora tubers consists of a nonepidermal layer made up of two distinctive types of cells, armature cells and stellate wart cells. Both cell types are provided with a heavy wall, and are dead at maturity. Stellate warts in the three species investigated,B. elongata, B. fungosa, andB. hansenii, séem to be uniform in appearance, but armature cells are extremely distinctive for each species. They are present in large, agglomerate masses in the first, singly or in very small clusters in the second, and as completely free, individual, acicular cells in the third species. Such differences are believed to be significant systematically, and the separation ofB. hansenii is thus probably justified. Notwithstanding superficial similarities, stellate warts do not seem to be comparable to the lenticels of other plants.     

Effects of delaying fungicide treatment of wounded potatoes on the incidence of Fusarium dry rot in store

Potato tubers artificially inoculated with Fusarium solani var. coeruleum or F. sulphureum 3 months after harvest were uniformly wounded and held at 5, 10 or 15°C for up to 21 days before immersion in fungicide suspensions. Holding tubers for 14 days at 15°C (curing conditions) or at 5°C did not alter the incidence of dry rot subsequently developing on tubers stored at 10°C, and holding tubers for up to 21 days at 15°C slightly increased disease caused by both pathogens. Thiabendazole, imazalil and prochloraz applied to tubers immediately after wounding almost completely prevented dry rot. Treatment after 3 days was less effective and the amount of disease increased with further delay; fungicides were more effective on tubers held at 5°C than at 10 or 15°C before treatment and storage, and efficacy of the fungicide was decreased by increasing the amount of inoculum on tubers. Wounds became less susceptible to infection by F. solani var. coeruleum and F. sulphureum when tubers were held at 15°C before inoculation, and the incidence of rots was decreased by 70–80% by delaying inoculation for 7 days. Treating tubers with dichlorophen immediately after wounding slightly increased the disease. The effects of fungicide treatment, curing conditions and wound healing are discussed.     

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.     

Tracing the Path of Oxygen into Birdsfoot Trefoil and Alfalfa Nodules using Iodine Vapor

Although physiological control of nodule 0₂ permeability is an active area of research, the gas diffusion pathway between the atmosphere and the infected zone has not been firmly established. Previous studies have used infiltration of ink or dyes to identify points of entry, but such water-soluble tracers could give a misleading picture of gas diffusion pathways. We therefore used iodine vapor (and its reaction with starch) to trace gas-phase pathways into the infected zone of determinate birdsfoot trefoil (Lotus corniculatus) and indeterminate alfalfa (Medicago sativa) nodules. We also used histochemical methods to identify suberized or lignified layers that could act as barriers to gas diffusion. Birdsfoot trefoil nodules were surrounded by a suberized periderm, but nonsuberized cells and intercellular spaces were observed in the periderm between lenticels and their associated vascular bundles. Iodine entered birdsfoot trefoil nodules only through lenticels. The periderm appears to provide a significant barrier to gas diffusion. Although airspaces were rare in the nodule parenchyma (also referred to as the “inner cortex”), we found some evidence that a few air-filled pathways cross this secondary barrier, also in the vicinity of vascular bundles. Alfalfa nodules were cylindrically surrounded by a suberized endodermis which ended near the meristematic tip; iodine entered principally at the end of the endodermis near the meristem. Future research on physiological control of nodule O₂ permeability should concentrate on strategic “choke points”, associated with lenticels in determinate nodules, or in the zone proximal to the meristem in indeterminate nodules.     

Effect of certain cultural practices on susceptibility of potato tubers to soft rot disease caused by Erwinia carotovora pv. carotovora

Erwinia soft rot causes destructive and serious damage to many vegetable crops including potato in the field, transit and storage periods. The effect of certain cultural practices on the susceptibility of potato tubers to soft rot bacteria was studied and the results of this work can be summarised in the following: potato tubers harvested on 1 May first exhibited the highest disease incidence compared with those harvested on 15 May or 30 May. Harvesting on 15 June resulted in the lowest disease infection. The application of high levels of nitrogen fertiliser as urea (46.5%), ammonium nitrate (31%) and ammonium sulphate (20.5%) resulted in an increase of the susceptibility of potato tubers to bacterial soft rot disease. In contrast, the addition of phosphorous as superphosphate (15.5%) fertiliser caused the reverse effect. The addition of potassium as potassium sulphate (48%) alone at any of the tested levels showed no effect. The susceptibility of potato tubers to bacterial soft rotting disease was increased by increasing storage periods at 4°C for 1, 2, 3 and 4 months. Spraying copper sulphate exhibited the highest decrease in soft rot incidence disease followed by manganese, zinc and iron. However, spraying of boron increased susceptibility to the disease. Potato tubers obtained from plants sprayed with copper and stored for different periods showed the lowest susceptibility to disease incidence. Tuber sprayed with zinc, iron, manganese and finally boron came next.     

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.