Assessment of the losses caused to potatoes by the potato cyst nematodes, Globodera rostochiensis and G. pallida

Nine field trials were carried out from 1967 to 1973 on sandy loam soils in Staffordshire and Worcestershire to study the relationship between potato yield and numbers of potato cyst nematodes (Globodera spp.). Three (probably five) of these trial sites appeared to be infested with G. rostochiensis only, whereas both species occurred on the remaining four sites which grew the resistant variety, Maris Piper. Although mixtures of the two species occurred on some plots they tended to have different distributions within each trial site. The results from this work have been analysed, together with those from earlier work in the Eastern Counties on peat and silt soils, over two population density ranges, 0–40 eggs/g (5 sites) and 0–160 eggs/g (16 sites). There is no evidence of any difference in the regression of yield on eggs/g amongst the five sites in the lower range (b =–0.90 + 0.11) nor amongst 15 of the 16 sites in the higher range (b = -0–40 ± 0–02). Analysis of 10 sites with sufficient data in the 40–160 eggs/g range gave b = -0.24 ± 0.06. Thus the regression lines are essentially parallel for each of the two ranges, covering several potato varieties, soil textures and different potential yields and suggest that the varieties used are equally tolerant. The losses are 6.25 t/ha/20 eggs/g for the 0.40 eggs/g range, 1.67 t/ha/20 eggs/g for the 40–160 eggs/g range, and 2.75 t/ha/20 eggs/g as a mean for the whole range. A maximum loss of 22 t/ha is indicated. Peat soils are less dense than mineral soils but there was no need for adjustment in nematode counts. Losses caused by potato cyst nematodes are better expressed as actual yield losses rather than as percentage decreases.     

The assessment of the tolerance of partially resistant potato clones to damage by the potato cyst nematode Globodera pallida at different sites and in different years

A range of potato cultivars and clones was grown at three sites infested with Globodera pallida in each of four years (1983 – 1986). Comparison of yields from nematicide treated plots with those of untreated plots provided estimates of yield losses due to G. pallida. The proportional yield losses were calculated for each genotype at each site as a measure of nematode tolerance. There were significant differences between genotypes with regard to tolerance of damage by G. pallida. There were also significant interactions between the different genotypes and the sites and years, but the interactions between years and genotypes represented only a small proportion of the observed variation. The interactions between sites and years indicate that not all of the genotypes behaved consistently between sites and years. The extent of yield losses caused by the nematodes varied significantly between sites. The implications of these results are discussed in relation to potato breeding programmes and assessing resistance and tolerance to G. pallida.     

Basal fertiliser application method, tuber initiation nitrogen, foliar NPK and the tolerance of potatoes to infection by the potato cyst nematodes Globodera rostochiensis and G. pallida

The effects of broadcast granular, placed liquid and foliar fertilisers on the tolerance of potatoes to infection by potato cyst nematodes were investigated. The tolerance of the potato cv. Pentland Dell was not significantly improved by fertiliser application type but placed liquid fertiliser, with or without foliar applications, increased the concentrations of N, P and K measured in whole plant dry matter of PCN infected plants. The tolerance of the potato cv. Sante was not statistically improved by altering the balance of fertiliser nitrogen applications between planting and tuber initiation or by applying foliar nitrogen. Nitrogen applications of 120 kg N ha^-1 at planting and a further 120 kg N ha^-1 at tuber initiation supplemented with foliar N, however, achieved a larger tuber yield than the same nitrogen programme without foliar N and gave a significantly greater yield than the application of 240 kg N ha^-1 at planting plus foliar N. The emergence of both cultivars was delayed in the absence of oxamyl. N, P and K concentrations within whole plant dry matter were significantly higher in plants from oxamyl treated plots and both N and K concentrations were significantly increased by increasing the quantity of N at planting, at 56 DAP. Splitting the fertiliser N between planting and tuber initiation appears to be important in maintaining the availability of this nutrient to PCN infected plants throughout the season.     

Hatching of Globodera pallida juveniles by diffusate of potato genotypes, differing in tolerance to G. pallida

Hatching induced by root diffusate, obtained from various potato genotypes, and by standard potato root diffusate, was determined in vitro. The used potato genotypes differed considerably in tolerance to Globodera pallida. A three parameter logistic model was used to describe the numbers of hatched juveniles in relation to time of exposure to root diffusate. Clear differences in hatching characteristics between genotypes were found. Some tolerant genotypes induced hatching of G. pallida juveniles relatively slowly, compared to intolerant genotypes. Other tolerant genotypes, however, induced hatching as fast as intolerant genotypes, and no significant correlation between hatching parameters and tolerance was found.     

The identification of the potato cyst nematodes Globodera rostochiensis and G. pallida by isoelectric focusing of proteins on polyacrylamide gels

Isoelectric focusing on thin layers of polyacrylamide gel was used to separate proteins from the potato cyst-nematodes Globodera rostochiensis and G. pallida. General protein patterns could be used to identify the two species, even from. single cysts. Staining for the enzyme phosphoglucomutase (PGM) also produced species specific patterns and the presence of PGM variation among G. pallida populations was revealed. The potential of isolectric focusing as a routine research and advisory tool in nematology is discussed.     

Differences in the tolerance of potato cultivars to potato cyst nematodes (Globodera rostochiensis and G. pallida) in field trials with and without nematicides

In field trials Cara, Brio, Maris Piper and Pentland Javelin were consistently more tolerant of damage by Globodera rostochiensis yielding more than Corsair, Pentland Dell, Maris Anchor and Maris Peer, in untreated, heavily infested soil and giving the smallest increases to nematicide treatment. No yield or growth differences were found between plants in untreated and nematicide treated plots at a nematode-free site. The most tolerant cultivars all had a gene (H₁) for resistance to G. rostochiensis derived from Solanum tuberosum ssp. andigena and in soil infested with G. pallida the tolerance of at least one resistant cultivar (Maris Piper) appeared to be lessened. However, some resistant cultivars were comparatively intolerant, even to G. rostochiensis. Early cultivars were generally less tolerant than late maturing cultivars but there were exceptions. Amongst cultivars with resistance derived from Solanum vernei the early cultivar Guardian was more tolerant than the main crop cultivar Corsair. The effect on the yield of several cultivars of a range of densities of G. rostochiensis, produced either by applying different rates of a nematicide or by cropping in the previous year, was examined at two sites. The results indicated that the slope of the regression for yield in relation to nematode density was less for tolerant than intolerant cultivars. At sites infested with G. rostochiensis Maris Piper was found to be consistently more tolerant than Pentland Crown.     

Supplementary foliar N, P and K, applied individually or in combinations, and the tolerance of potatoes to infection by the potato cyst nematodes Globodera rostochiensis and G. pallida

The use of supplementary foliar N, P and K to ameliorate the reduced nutrient uptake of potato plants infected by potato cyst nematode (PCN) were investigated. The potato cv. Pentland Dell achieved yields in plots not treated with oxamyl similar to those found in plots treated with oxamyl when supplementary foliar N or N plus K was applied to plots infested with 13 eggs g^-1 soil of Globodera pallida. Yield improvements from foliar N applications were attributed to increased leaf area index but the reason for yield increases from foliar N plus K applications could not be clarified. In a second experiment, where PCN infestation was 76 eggs g^-l soil, the potato cv. Sante gave yields up to 19% higher than a standard fertiliser practice when supplementary foliar N was applied to plots not treated with oxamyl. Nutrient analysis showed that without oxamyl there were significantly lower concentrations of N, P and K in whole plant dry matter at 58 days after planting (DAP) but higher levels of N in the fourth leaf dry matter at 98 DAP. Emergence was significantly advanced by the use of oxamyl in both experiments. Sante dramatically reduced populations of Globodera rostochiensis from an average of 76 eggs g^-1 soil to 7 eggs g^-1 soil. Foliar application of nutrients is a promising method of ameliorating the effects on potatoes of PCN invasion but the nutrient concentrations and timing of individual sprays need to be more closely matched to crop requirement than was possible in our experiments     

Effects of rates of a nematicide and of fertiliser on the growth and yield of cultivars of potato which differ in their tolerance of damage by potato cyst nematodes (Globodera rostochiensis andG. pallida)

Potato plants growing in soil heavily infested with potato cyst nematode (PCN) contained less N, P and K in their leaf dry matter than plants growing in the same soil treated with a nematicide. These differences were less in tolerant than intolerant cultivars. Applying additonal fertiliser increased the growth of untreated plants more than that of nematicide-treated plants and nematicides increased growth most in plots receiving the lowest rate of fertiliser. Overall, the results are consistent with the hypothesis that damage by invading juveniles of PCN decreases the effectiveness of the potato root system leading to a chronic deficiency of one or more nutrients and a consequential reduction in the rate of top growth.     

Population decline of potato cyst nematodes (Globodera rostochiensis, G. pallida) in field soils in Northern Ireland

The decline rates of potato cyst nematodes (Globodera rostochiensis, G. pallida) in agricultural soils in Northern Ireland were monitored over 13 years. Initial decline rates appeared erratic due to variable cyst ages and high Standard Errors at low population levels, but were, generally, slower than those previously reported with less than 10% annual decline within each category of assessment (cysts sample^-1, eggs g^-1 and eggs cyst^-1). Rate of decline was not affected by the range of soil types studied, but regular cultivation of infested land appeared to accelerate it. Juveniles present within cysts over 30 years old were either dead or inactive and such field populations were considered non-viable.     

Tolerance of potato to potato cyst nematodes (Globodera rostochiensis and G. pallida) in relation to the growth and efficiency of the root system

Potato cyst-nematode (Globodera rostochiensis) was shown to damage potato plants in several ways. A major cause of damage, affecting all cultivars to a similar extent, was a reduction in the top to root weight ratio. Intolerant cultivars also suffered a reduction in the weight and length of their root systems when grown in heavily infested soil, the combined damage resulting in a marked decrease in nutrient uptake and top growth. In addition intolerant cultivars tended to senesce prematurely when heavily infested, further decreasing their leaf area duration and yield. Cultivars tolerant of potato cyst-nematode (PCN) differed from intolerant cultivars in that their root systems tended to grow larger in heavily infested soil than in lightly infested or nematicide-treated soil, so partly compensating for the reduction in the top/root ratio. In a growth cabinet experiment Maris Anchor was more severely damaged at a soil temperature of 10 than at 15 °C. In a glasshouse, without temperature control, differences were obtained between cultivars in small pots (10 cm) in the effect of PCN on root growth which correlated well with differences in tolerance obtained in field trials.     

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.     

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.     

The identification and fitness of virulent potato cystnematode populations (Globodera pallida) selected on resistant Solarium vernei hybrids for up to eleven generations

Selection of Globodera pallida populations on resistant Solanum vernei-hybrids resulted in distinct virulent strains after eleven generations. Some of these virulent populations were assessed on their environmental fitness under field-type conditions. All reproduced less well in unsterilised soil, but virulent populations were less affected by environmental variation than their avirulent counterparts. Evaluation of their reproductive ability could not equate virulence to overall enhanced or reduced genotypic fitness compared with their avirulent counterparts. These populations were shown to be genetically distinct from their unselected counterparts using isoelectric focusing and specific enzyme staining. The control and management of virulent G. pallida populations is discussed.     

Management of the potato cyst nematode (Globodera pallida) with bio-fumigants/stimulants

Field trials evaluated the effect of four plant-based bio-fumigants/stimulants on population levels of G. pallida and the resulting potato yields and quality. Three formulations contained seaweed biostimulants (Algifol, Nutridip and Metastim) and one bio-fumigant containing mustard and chilli pepper extracts (Dazitol). These were compared with the fumigant nematicide Nemathorin and untreated control plots. The effect of G. pallida on growing potato crops was assessed by recording haulm characteristics which indicated that the nematicide treatment gave most protection. Levels of PCN juveniles and migratory nematodes were assessed during the trial. Plots treated with Nemathorin and Dazitol had fewest PCN, whilst the highest number of migratory nematodes occurred in fallow plots. Sixteen weeks after planting the nematicide treatment produced highest yield and tuber numbers. Dazitol treatment produced a lower yield but the largest tubers.     

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.     

The effect of the potato cyst nematode Globodera pallida on in vitro root growth of potato genotypes, differing in tolerance

Roots of eighteen potato genotypes, differing in tolerance of G. pallida, were grown from tuberpieces on agar in Petri dishes. Juveniles of G. pallida were inoculated directly onto root tips. Root length was measured at various times after inoculation. Inoculation reduced root growth within one day. At later stages, genotypes differed strongly in growth of inoculated roots. Between four and seven days after inoculation, growth of inoculated roots was not significantly correlated with growth of untreated roots, and was only poorly correlated with tolerance assessed in the greenhouse or in the field. However, multiple regression analysis revealed that the tolerance of the tested genotypes was associated with both the rate at which they induced hatching and the growth of roots after inoculation. The combination of these two variables accounted for high percentages explained variance.     

Differentiation of Closely Related Globodera pallida (Stone) Populations by Means of DNA Restriction Fragment Length Polymorphisms (RFLPs)

A small genomic library from DNA of G. pallida population Delmsen (pathotype 3) was constructed. A number of clones were tested as hybridization probes with 21 G. pallida and 6 G. rostochiensis populations. The two species were easily distinguished by their hybridization patterns. A total of 32 RFLP markers were identified which permitted the differentiation of all G. pallida populations. By means of cluster analysis of the RFLP data the genetic distances between the 21,G. pallida populations were calculated and a dendrogram was constructed. The arrangement of the 21 G. pallida populations in the dendrogram suggests a partial correlation between genetic distance and virulence according to biological pathotyping.     

Development of diagnostic markers for use in breeding potatoes resistant to Globodera pallida pathotype Pa2/3 using germplasm derived from Solanum tuberosum ssp. andigena CPC 2802

Quantitative resistance to Globodera pallida pathotype Pa2/3, originally derived from Solanum tuberosum ssp. andigena Commonwealth Potato Collection (CPC) accession 2802, is present in several potato cultivars and advanced breeding lines. One genetic component of this resistance, a large effect quantitative trait locus (QTL) on linkage group IV (which we have renamed GpaIV _adg ^s ) has previously been mapped in the tetraploid breeding line 12601ab1. In this study, we show that GpaIV _adg ^s is also present in a breeding line called C1992/31 via genetic mapping in an F₁ population produced by crossing C1992/31 with the G. pallida susceptible cultivar Record. C1992/31 is relatively divergent from 12601ab1, confirming that GpaIV _adg ^s is an ideal target for marker-assisted selection in currently available germplasm. To generate markers exhibiting diagnostic potential for GpaIV _adg ^s , three bacterial artificial chromosome clones were isolated from the QTL region, sequenced, and used to develop 15 primer sets generating single-copy amplicons, which were examined for polymorphisms exhibiting linkage to GpaIV _adg ^s in C1992/31. Eight such polymorphisms were found. Subsequently, one insertion/deletion polymorphism, three single nucleotide polymorphisms and a specific allele of the microsatellite marker STM3016 were shown to exhibit diagnostic potential for the QTL in a panel of 37 potato genotypes, 12 with and 25 without accession CPC2082 in their pedigrees. STM3016 and one of the SNP polymorphisms, C237(119), were assayed in 178 potato genotypes, arising from crosses between C1992/31 and 16 G. pallida susceptible genotypes, undergoing selection in a commercial breeding programme. The results suggest that the diagnostic markers would most effectively be employed in MAS-based approaches to pyramid different resistance loci to develop cultivars exhibiting strong, durable resistance to G. pallida pathotype Pa2/3.     

Use of resistant and susceptible potato cultivars in the trap cropping of potato cyst nematodes, Globodera pallida and G. rostochiensis

The effects of planting date and growing period of potato cultivars on their efficiency as trap crops for potato cyst nematodes (PCN) were studied. Plots were planted with susceptible or resistant cultivars in April, June and August and these were grown for 5, 6 or 7 wk before removal of the plants by hand lifting. Crops planted in June provided the best overall reductions in PCN population density of up to 95%, with cv. Santé significantly more effective than the other cultivars. Population reductions from the August planting were only slightly less than from planting in June but the tuber yields obtained were much greater: Maris Piper and Maris Bard produced 16.4 and 21.4 t ha^-1 respectively, with 37% and 43% respectively, of a size useful for canning (i.e. between 20 and 40 mm diameter).