Effects of Zygorrhynchus moelleri on the growth of potato and reproduction of potato cyst nematodes

Laboratory, pot and field experiments investigated the effects of the fungus Zygorrhynchus moelleri on the growth of potato and on the reproduction of the potato cyst nematodes (PCN), Globodera pallida and G rostochiensis. Preliminary laboratory tests showed that Z. moelleri growth was favoured by temperatures and pH ranges commonly present in field soils. The fungus colonised potato roots in vitro and in compost or field soil. It also stimulated in vitro root growth of three potato cultivars. In pot experiments Z. moelleri stimulated potato growth, particularly in the presence of PCN attack. In field plots infested with a mixture of G pallida and G. rostochiensis, tuber yields were not increased after application of the fungus but, in G pallida‐infested plots, yields were significantly increased after drills were inoculated with Z. moelleri. The application of Z. moelleri had no apparent effects on nematode reproduction. Factors influencing the interactions between Z. moelleri, potato and potato cyst nematodes are discussed and the potential role of the fungus as a plant growth promoter in organic potato production considered.     

Effect of potato used as a trap crop on potato cyst nematodes and other soil pathogens and on the growth of a subsequent main potato crop

A field experiment in which main‐crop potatoes were grown every other year was conducted on a sandy soil from 1994 to 1999. The aim of the experiment was to control soil‐borne pathogens of potato with ecologically sound methods. Potato grown as a trap crop from the end of April to the end of June (8 wk) was used to control potato cyst nematodes (PCN) (Globodera pallida), and its effects on other important soil pathogens and on the growth of a subsequent potato crop were also assessed. Additional experimental treatments were a potato crop from which the haulm was removed and a green manure crop. Three potato cultivars with different degrees of resistance to PCN were grown as the main crop. Duplicate sets of the experiment were run concurrently. The PCN were effectively controlled by the potato trap crop. When a highly resistant potato cultivar was grown as a main crop after the trap crop, the post‐harvest soil infestation was very low. When a moderately resistant cultivar was grown after the trap crop the soil infestation also remained low. When the trap crop was alternated with a susceptible potato cultivar as a main crop, soil infestation increased slightly, but the degree of control when compared with no trap crop averaged 96%. Soil infestation with root‐knot nematodes (mainly Meloidogyne hapla) increased when potato was grown as a trap crop, but soil infestation with the root‐lesion nematode Pratylenchus crenatus was not affected. Stem canker caused by Rhizoctonia solani was not affected by the trap crop but black scurf (sclerotia of R. solani) on tubers was reduced. Soil infestation with Verticillium dahliae declined in one of the duplicate sets of the experiment but not in the other. However, stem infections by V. dahliae were significantly decreased in both sets, although the effect depended on the PCN‐resistance level of the potato cultivar. When a highly resistant potato cultivar was grown Verticillium stem infections were not significantly affected, they were decreased with a moderately resistant cultivar but the decrease was most pronounced with a PCN‐susceptible cultivar. Senescence of a following potato crop was not influenced by the trap crop when a highly PCN‐resistant cultivar was grown, but it was delayed in the case of a moderately resistant or a susceptible cultivar, resulting in higher tuber yields for those cultivars. The experiment proved that a trap crop can be an alternative to chemical soil disinfection but, for several reasons, the potato itself is not an ideal crop for this purpose; a trap crop other than potato must be developed.     

Co-limitation of potato growth by Potato Cyst Nematode (Globodera rostochiensis) and Rhizoctonia solani

Globodera rostochiensis and Rhizoctonia solani are the most important growth limiting factors influencing potato production in Iran. The effects of inoculation with Potato Cyst Nematodes (PCN) (0, 50, 75 and 100 cysts/3.5?kg soil) and R. solani (with or without inoculation) on potato growth and development were investigated in cultivars Santé and Marfona. Inoculation with R. solani induced severe damage, especially when inoculation was accompanied with high density of PCN. The damage caused by R. solani tended to increase with an increase in PCN density, especially in Marfona. In Santé, number of stems or branches per plant significantly increased by inoculation with R. solani, while in Marfona it was significantly affected either by R. solani inoculation or PCN density. In Santé, number of stolons per plant was significantly increased by PCN, but not by R. solani. In Marfona, however, the number of stolons per plant was significantly affected either by R. solani inoculation or by presence of PCN, but not affected by PCN density. The general effect of R. solani or PCN inoculation treatments on shoot, below-ground and total dry weight of potato was significant, but strongly affected by cultivar. In general, our study supports the synergistic interaction between R. solani and PCN and its moderation by the use of a resistant cultivar such as Santé.     

Effects of potnetial trap crops and planting date on soil infestation with potato cyst nematodes and root-knot nematodes

In 1997 and 1998 the stimulation of hatch of potato cyst nematodes (PCN) by a trap crop was studied at various times during the growing season in a container and a field experiment. Solanum nigrum‘90‐4750‐188’was used as the trap crop in both experiments and was sown on 1 May, 16 June or 1 August in two successive years on different plots. Neither experiment revealed much seasonal variation in hatchability of PCN juveniles under a trap crop. In the container experiment, the hatch of the Globodera pallida Pa3 population was equally and strongly stimulated (89%) at all sowing dates in both years, except for the 1 August sowing in 1998 (when the hatch was 77% under extremely wet soil conditions). In the control treatment with non‐hosts (flax followed by barley) the total spontaneous hatch was 50% over 2 yr. In the field experiment, the hatch of PCN, averaged over the four populations, was also equally stimulated (71%) at all sowing dates in both years. In the control treatment with non‐hosts (flax‐barley) the total spontaneous hatch was 36% over 2 yr. Total hatch under the trap crop over 2 yr varied between the four PCN populations from 63% to 80%. In 1998 and 1999, control of potato cyst nematodes (PCN) by the potential trap crops Solanum sisymbriifolium and S. nigrum‘90‐4750‐188’was studied in the field. Potato was also included as a trap crop. In the 1998 experiment, potato, S. sisymbriifolium and S. nigrum strongly stimulated the hatch of PCN compared with the non‐host white mustard (Sinapis alba). Roots of potato and white mustard were mainly found in the top 10 cm of soil, whereas roots of S. sisymbriifolium and S. nigrum were also abundant at depths of 10–20 cm and 20–30 cm. In the 1999 experiment, soil infestation with PCN decreased markedly with potato and S. sisymbriifolium as trap crops. In plots moderately to severely infested with 2‐yr old cysts (2–29 juveniles ml^?1 air dried soil), potato reduced soil infestation by 87% and S. sisymbriifolium by 77%. In plots moderately to severely infested with 1‐yr old cysts the reductions were 74% and 60%, respectively. The reduction was least on plots very severely infested with PCN (110–242 juveniles ml^?1 soil): 69% and 52% for potato and S. sisymbriifolium, respectively. Soil infestations of plots that were initially slightly to severely infested with the root‐knot nematode Meloidogyne hapla were greatly reduced under fallow and S. sisymbriifolium but increased under potato. From these and previous experiments it was concluded that, for several reasons, S. sisymbriifolium is a promising trap crop.     

The effect of resistant potato cultivars on a field population of Globodera pallida Pa2 over three years of cropping

Six potato cultivars with different levels of resistance to the white potato cyst nematode (PCN) Globodera pallida Pa2 were grown for three seasons in field plots to which G. pallida Pa2 cysts had been introduced earlier. There were two planting times, corresponding to early and maincrop commercial planting times, and two initial PCN population densities, high and low. The effect of cultivar on PCN population density was far greater than the effect of planting time or initial nematode population. The final PCN populations for the cultivars Ilam Hardy, Wha, 4696A(2), Sovereign, D40/6 and V390 were 151, 74, 27, 1.4, 0.2 and 0.06 eggs per g of soil respectively. It is concluded that resistant potato cultivars can be very effective in controlling G. pallida Pa2 in the field.     

Spread of potato leafroll virus is decreased from plants of potato clones in which virus accumulation is restricted

Tubers of eight potato clones infected with potato leafroll luteovirus (PLRV) were planted as ‘infectors’ in a field crop grown, at Invergowrie, of virus-free potato cv. Maris Piper in 1989. The mean PLRV contents of the infector clones, determined by enzyme-linked immunosorbent assay (ELISA) of leaf tissue, ranged from c. 65 to 2400 ng/g leaf. Myzus persicae colonised the crop shortly after shoot emergence in late May and established large populations on all plants, exceeding 2000/plant by 27 June. Aphid infestations were controlled on 30 June by insecticide sprays. Aphid-borne spread of PLRV from plants of the infector clones was assessed in August by ELISA of foliage samples from the neighbouring Maris Piper ‘receptors’. Up to 89% infection occurred in receptor plots containing infector clones with high concentrations of PLRV. Spread was least (as little as 6%) in plots containing infectors in which PLRV concentrations were low. Primary PLRV infection in guard areas of the crop away from infectors was 4%. Some receptor plants became infected where no leaf contact was established with the infectors, suggesting that some virus spread may have been initiated by aphids walking across the soil.     

Dynamics and management of the white potato cyst nematode Globodera pallida in commercial potato crops

Eight trials were conducted in commercial potato fields infested with the white potato cyst nematode (wPCN, Globodera pallida) and one in a field infested with the yellow PCN (yPCN, Globodera rostochiensis). Our aims were to produce data to validate and refine a computer‐based program (The Model) for the long‐term management of PCN, to determine nematicide effectiveness and to assess rates of PCN population decline between potato crops. Prior to planting, each farmer applied an overall nematicide treatment to his field, except for ten untreated plots that were widely spaced to encompass a range of PCN population densities. Each untreated plot was paired with a similar plot in the adjacent treated area and all plots were intensively sampled for PCN population densities at planting (P_i) and again at harvest (P_f) when tuber yields were determined. Four trials were re‐sampled 2–4 years later to determine PCN population decline rates. Regressions that form the basis of ‘The Model’ and described the relationship between P_i and tuber yield and PCN population density at harvest were fitted to the results from both the untreated and nematicide treated plots. These regressions also enabled us to estimate the yield potential at each site in the absence of PCN and showed that nematicide treatment generally did not increase yield potential and that both tuber yield and PCN multiplication decreased with increasing P_i. However, there were major differences between sites and cultivars. When untreated, the yield of cv. Maris Piper was hardly affected in a highly organic soil with P_i > 200 eggs g^?1 whereas the yield of partially resistant cv. Santé was decreased from a potential of c. 60 t ha^?1 to c. 20 t ha^?1 in a light silt with P_i = 20 egg g^?1 soil. Similarly, untreated wPCN multiplication rates at a low P_i ranged from 46‐fold to >100‐fold. Nematicide effectiveness was estimated from the regressions and, at several sites, yield was decreased despite nematicide treatment. Control of wPCN multiplication was even poorer. In only two of seven trials planted with susceptible cultivars was more than 50% control achieved – maximum populations in treated plots usually exceeded 250 eggs g^?1. Partially resistant Santé decreased the multiplication rate of wPCN in the two trials where it was planted. An alternative analysis using Genstat indicated that The Model tended to underestimate the maximum multiplication rate and overestimate the maximum population density. When four sites were re‐sampled 2–4 years after harvest the populations of wPCN had declined by between 15% and 33.5% per annum with a mean of 26% per annum. Modelling indicated that rotations longer than 8 years were required to control wPCN unless other effective control measures, such as growing a partially resistant cultivar, were used.     

Integrated control of potato cyst nematodes using small amounts of nematicide and potatoes with partial resistance

The control of potato cyst nematode (PCN) by less than approved amounts of nematicide combined with partially resistant potato clones was studied in a series of field experiments. On a site heavily infested with Globodera pallida only the most resistant clone (12380ac2) decreased the population density in untreated soil. With aldicarb at its full approved rate (3·36 kg ha^-1) numbers of PCN were decreased under all the genotypes, including the non-resistant Maris Piper. Aldicarb at 1·68 kg ha^-1significantly decreased populations on all clones except 12380ac2. Aldicarb at 0·84 kg ha^-1still significantly decreased population densities and multiplication rates of G. pallida on two clones with intermediate resistance (12243acl and 11233ab22). At two G. rostochiensis sites with light infestations nematode multiplication rates were greater and the control given by aldicarb and partially resistant genotypes of potato was not as great as that at the site with G. pallida. Tuber yields were not increased by the application of aldicarb at the G. rostochiensis infested sites. However, at the site heavily infested with G. pallida the yield of the most intolerant genotype (12380ac2) was increased seven-fold by the full rate of aldicarb (3·36 kg ha^-1) and four-fold by the quarter rate (0·84 kg ha^-1)-Yield of the most tolerant genotype (12243acl) was unaffected by the application of aldicarb.     

Effect of the fungus Pochonia chlamydosporia and fosthiazate on the multiplication rate of potato cyst nematodes (Globodera pallida and G. rostochiensis) in potato crops grown under UK field conditions

This is the first report of the successful use of Pochonia chlamydosporia as a biological control agent against potato cyst nematodes (PCN) (Globodera pallida and G. rostochiensis) in potato crops grown under commercial field conditions and represents an important step in the development of biological control for PCN. Two field experiments were established in consecutive years (2006 and 2007) at different field sites in Shropshire, England. Treatments comprised of (1) untreated control, (2) P. chlamydosporia, (3) P. chlamydosporia with the nematicide fosthiazate and (4) fosthiazate alone. In both experiments, significant reductions in the nematode multiplication rate (Pf/Pi) for P. chlamydosporia treated plots were observed (48% and 51% control, respectively). The P. chlamydosporia treatment did not differ significantly from both fosthiazate treatments in terms of Pf/Pi in spite of the trend towards increased control particularly in Experiment 1. P. chlamydosporia therefore provided similar levels of nematode population control as fosthiazate. The combined treatment did not provide any additional reduction in Pf/Pi but demonstrated that P. chlamydosporia was compatible with fosthiazate. Over the different developmental stages of the juvenile nematodes, there was evidence of parasitism of adult females on the plant root by P. chlamydosporia. Root colonization by P. chlamydosporia was higher in the P. chlamydosporia treatment due to increased levels of nematodes in plant roots. Results from both experiments demonstrated the efficacy of P. chlamydosporia as a biological control agent of PCN and indicate its potential for use as part of an integrated pest management strategy.     

The use of foliage assessment to improve the identification of tolerance to damage by nematodes (Globodera pallida) in potatoes

Six potato trials, two in each of three years, were conducted in collaboration with the Agricultural Development and Advisory Service (ADAS) at sites infested with potato cyst nematodes (G. pallida Pa 2/3). The trials were part of a selective screen to identify PCN tolerant and intolerant clones with each trial consisting of four blocks divided into nematicide treated and untreated sub-blocks. A range of partially resistant and susceptible material was assessed for yield losses due to PCN damage to the roots and for the effect on the foliage by comparison between the nematicide treated and untreated areas. The relationships between the foliage symptoms, untreated yields, treated yields, proportional yield loss, initial PCN population and the post-harvest PCN population levels are examined. Significant correlation coefficients were obtained between foliage symptoms and yield of clones in PCN infected soil and also between foliage symptoms with percentage yield loss due to PCN infestation. The conclusions were that the assessment of PCN damage to foliage vigour/development can contribute positively to a more accurate identification of tolerant or intolerant potato genotypes.     

Control of potato cyst nematodes and economic benefits of application of 1,3-dichloropropene and granular nematicides

The two species of the potato cyst nematodes (PCN) Globodera pallida and G rostochiensis are the most problematic pests of the potato crop in the UK. There are no commercially available cultivars with full resistance to G. pallida and both crop rotation and granular nematicides are less effective at controlling this species than G. rostochiensis. In situations of very high PCN levels it may be possible to reduce populations and yield losses by using an autumn application of the soil fumigant 1,3-dichloropropene (1,3-D) followed by a spring application of a granular nematicide. Two field experiments were done to look at the integration of methods for the control of PCN. The Common Field experiment (G. rostochiensis infested) compared the use of 1,3-D with the granular nematicides aldicarb, oxamyl and fosthiazate when growing the susceptible cv. Estima. The Four Gates experiment (infested with both PCN species but mainly G rostochiensis) compared the performance of cv. Santé (partially resistant to G. pallida, fully resistant to G. rostochiensis) with that of the susceptible cv. Estima when treated with 1,3-D and oxamyl at full and half-rates. The results of the experiments show that an integrated approach to nematode control on heavily infested sites, including granular and fumigant nematicides and cultivar resistance, can lead to significant decreases in nematode population densities and reduce yield losses. An economic evaluation of the experiments modelled the gross margins from the different nematicide treatments. In Common Field, the highest gross margins were achieved with the combined use of fumigant and granular nematicides. In Four Gates, there was a clear economic benefit for both cultivars from the use of 1,3-D. In this experiment, oxamyl was of economic value to Estima but not to Sante and full-rate oxamyl was of more benefit than half-rate to Estima.     

Effects of a mixed-isolate mycorrhizal inoculum on the potato – potato cyst nematode interaction

Inoculation of microplants of potato cv. Golden Wonder with Vaminoc, a mycorrhizal inoculum of three arbuscular mycorrhizal fungi (Glomus spp.), resulted in an increase in in‐sand hatch of Globodera pallida, but not G. rostochiensis, within 2 weeks. By this time, mycorrhized plants also supported a larger number of feeding nematodes of both PCN species (50% higher for G. rostochiensis) than did non‐mycorrhized plants, with a higher proportion of the G. pallida population being fertilised females than for G. rostochiensis. After 12 weeks, the multiplication rate of G. rostochiensis on mycorrhized plants was significantly greater than on non‐mycorrhized plants, whereas no such difference was observed for G. pallida. The principal component of PCN multiplication affected by mycorrhization was increased cyst number per plant from 6 to 12 weeks. Over this period, there was no increase in cyst number per plant for either PCN species on non‐mycorrhized plants, whereas the value increased on mycorrhized plants for both G. rostochiensis (by almost 200%) and G. pallida (57%). Mycorrhization resulted in significant increases in the root and shoot dry weights of plants grown in the absence of PCN. Although mycorrhized plants carried a larger PCN burden than non‐mycorrhized plants when grown on PCN‐infested medium, as a result of the increased PCN multiplication rate, they produced larger root systems than did nonmycorrhized plants, suggesting increased tolerance to PCN of the mycorrhized plants, particularly to G. rostochiensis. Of morphological characters investigated in the absence of PCN, only stem height (increased) was significantly affected by mycorrhization. Colonisation by mycorrhizal fungi resulted in increased tuber yield both in the absence (significant increase) and presence (non significant) of PCN, as a result of increased tuber number per plant. These results are discussed in the light of the possible use of AMF as part of an integrated PCN management plan.     

Aspects of resistance to sweet potato virus disease in sweet potato

In field trials during the first and the second rainy season of 1996 in Uganda, whiteflies were similarly abundant and aphids were absent on three clones of sweet potato (NIS-93–63, cv. Tanzania and cv. New Kawogo) although the three clones differed considerably in their resistance to sweet potato virus disease (SPVD), a complex disease resulting from infection by both the aphid-borne sweet potato feathery mottle virus (SPFMV) and the whitefly-borne sweet potato chlorotic stunt virus (SPCSV). This suggests that vector resistance does not determine the relative SPVD resistance of these genotypes. SPFMV alone had only a low virus titre in sweet potato cvs Tanzania and New Kawogo, became increasingly difficult to detect in plants of these cultivars and was seldom acquired by aphids. However, this resistance to SPFMV was not apparent in plants which were also infected with SPCSV. Plants then had a high SPFMV titre, appeared unable to eliminate SPFMV and provided good sources for aphids to acquire it.     

Control of potato cyst nematode (Globodera pallida) by host plant resistance and nematicide

Seven trials conducted over four years on sites naturally infested with the white potato cyst nematode established that potato clones bred for resistance to Globodera pallida allowed significantly less nematode multiplication than conventional cultivars under field conditions. Nematode multiplication was inversely related to initial infestation level. The nematicide, aldicarb, significantly reduced nematode multiplication. However, nematode multiplication on nematicide treated susceptible cultivars was greater than on untreated partially resistant clones, indicating that resistance may offer more effective control of G. pallida than chemical treatment. Integration of host plant resistance and nematicide treatment is discussed.     

Glucose phosphate isomerase polymorphism in field populations of the potato cyst nematodes, Globodera rostochiensis and G. pallida

The isozymic variation of glucose phosphate isomerase (GPI) was compared in 20 field populations of potato cyst-nematode (PCN) and related to variation in reference pathotypes of PCN. Populations pathotyped as G. rostochiensis Ro 1 by conventional differential cultivar tests were found to be different from populations of G. pallida. Variation within populations pathoptyped as G. pallida was seen and three populations were found to be similar to the reference G. pallida Pal pathotype. This seemed to be confirmed for two populations using cysts formed on a cultivar resistant to the Pa 1 pathotype when an isoenzyme pattern characteristic of G. palida Pa2 and Pa3 was found. The feasibility of the use of isozyme identification of PCN pathotypes is discussed in the light of these results.     

Superoxide dismutase polymorphism in field populations of the potato cyst nematodes Globodera rostochiensis and G. pallida

Superoxide dismutase (SoDase) polymorphism in reference populations of potato cyst nematodes (PCN) was studied by isoelectric focusing and compared with the banding pattern obtained from 26 English field populations of PCN. Qualitative and quantitative differences were found between the reference populations: a band detected only in the G. rostochiensis Ro4 and Ro5 reference populations was found in four field populations of varying pathotype composition and two field populations of G. rostochiensis Rol possessed extra bands that were not detected in the other field populations. Analysis of the band differences showed little correlation between pathotype and banding pattern in the field populations.     

Use of seedling progeny tests for genetical studies as part of a potato (Solanum tuberosum subsp. tuberosum) breeding programme

A diallel set of crosses, including selfs and some reciprocal crosses, was made between 15 parents chosen for their male fertility from those included in a tetraploid potato (Solanum tuberosum subsp. tuberosum) breeding programme at the Scottish Crop Research Institute. Seedling progeny tests were used to evaluate the progenies for non-race-specific resistance to late blight (Phytophthora infestans) in both foliage and tubers, quantitative resistance to the white potato cystnematode (PCN) (Globodera pallida) and the commercial worth of their tubers as judged by breeders' visual preference. No reciprocal differences were found. Comparisons of the selfs and crosses revealed inbreeding depression for breeders' preference, which varied among the parents from negligible to severe, whilst there were also statistically significant differences for foliage and tuber blight, but not for PCN. When the selfs were omitted from the combining ability analyses, large differences in general combining ability (GCA) were found for all four traits, and smaller differences in specific combining ability for tuber blight and breeders' preference. The only statistically significant correlation between GCAs for different traits was a favourable one of r = 0.56 between foliage and tuber resistance to late blight. It was concluded that prospects were good for simultaneously improving all four traits by multitrait genotypic recurrent selection.     

Differential accumulation of potato tuber mRNAs during the hypersensitive response induced by arachidonic acid elicitor

Accumulation of messenger RNAs in potato tuber discs was analysed during the hypersensitive response induced by treatment with the biotic elicitor arachidonic acid. In vitro translation of polysomal poly(A)⁺ RNAs indicated that the accumulation of some sixteen mRNAs varied following treatment with arachidonic acid, and that the level of thirteen of these was increased. Two cDNA closes (pSTH-1 and-2) were isolated from a library of elicitor-treated tissue cDNAs. Northern blot analysis using these clones as molecular probes indicated that the levels of at least two mRNAs were markedly increased after elicitor treatment. In hybrid-released translation experiments, each of the cDNA clones selected more than one mRNA. Translation of these mRNAs yielded two polypeptides of M_r 45 000 (for the pSTH-1 clone), and three polypeptides of M_e 17 000 (for the pSTH-2 clone). The low molecular weight polypeptides may correspond to potato pathogenesis-related (PR) proteins.