Selection for virulence in the potato cyst-nematode, Globodera pallida

The ability of six populations of Globodera pallida to multiply on Solanum vernei hybrids (Morag, 12380 2, Glenna and Santé) and S. tuberosum andigena hybrids (A 27/23, Paladin (ZB 35/29) and ZC 83/6) was studied in microplots and pots. Evidence was obtained of selection for virulence in one population after one year's cultivation of cvs Glenna or Morag. Similarly, after four years' continuous cultivation of cvs Morag or ZC 83/6 in microplots, three of five populations of G. pallida contained more virulent nematodes than the same populations maintained for the same period on cv. Pentland Crown (susceptible to G. pallida). These results are discussed in relation to earlier work and the use of partially resistant cultivars in the integrated control of G. pallida.     

The resistance of six potato cultivars to English populations of potato cyst-nematodes, Globodera pallida and G. rostochiensis

Six cultivars of potato (Santé, Morag, Paladin, Glenna and Fingal bred for resistance to both potato cyst-nematodes (Globodera rostochiensis and G. pallida) and Valiant bred for resistance to G. pallida alone) were exposed to 28 English populations of G. pallida and eight English populations of G. rostochiensis in pots. Susceptible cv. Désirée potatoes served as controls for all 36 populations. Inoculum (Pi) was 12000 eggs in cysts per 400ml pot of soil. Average increase of G. rostochiensis (Pf/Pi) on cv. Désirée was 23.5 but on cvs Sante, Glenna and Fingal it was < 1.0 and on cv. Morag it was 2.2. In contrast, cvs Paladin and Valiant were susceptible (average Pf/Pi = 17.4 and 26.5, respectively). Against G. pallida populations, average Pf/Pi for cv. Désirée was 21.7; on cvs Paladin, Santé and Glenna it was 2.9, 2.6 and 2.4, respectively; cvs Morag and Fingal were less resistant (7.4 and 5.6, respectively) and cv. Valiant was quite susceptible (11.0). Resistance to the different populations of G. pallida and G. rostochiensis varied but for the most resistant cultivars (Santé, Glenna and Paladin) the variation was usually small. The value of the six resistant cultivars studied to the integrated control of potato cyst-nematodes in England and the genetic diversity of the nematode populations to which they were exposed are discussed.     

Observations on the control of potato cyst-nematodes, Globodera rostochiensis and G. pallida in 35 English potato soils in pots by oxamyl and resistant potatoes

The increase of 35 English field populations of potato cyst-nematodes (Globodera rostochiensis and/or G. pallida) was measured on Désirée, Maris Piper, Caxton (A25/11), Cromwell (A27/20) and clone 11233 ab 22 in pots of sandy, silty or peaty loam soil. Désirée was susceptible to all populations tested and, as in field soils, the final population (Pf) was inversely related to the initial population of potato cyst-nematode eggs (Pi) in the soil. Maris Piper and Cromwell were resistant to all G. rostochiensis populations, with one possible exception. Maris Piper was susceptible to all G. pallida populations. Caxton was susceptible to some and fairly resistant to other populations of G. rostochiensis, indicating the existence either of two biotypes within the one pathotype (Rol) as yet encountered in Britain, or the existence of an additional pathotype. Caxton and Cromwell were fairly resistant to G. pallida. Clone 11233 ab 22 was only moderately resistant to both species. Resistance to potato cyst-nematode increase varied considerably, especially in Caxton (to G. rostochiensis) and in 11233 ab 22 (to both species). Oxamyl greatly reduced the increase of G. rostochiensis populations on Désirée potatoes, with the notable exception of one population but it generally had much less effect on G. pallida populations, regardless of soil type. The difference in effect on the two species may be due to a longer period of hatching in G. pallida than in G. rostochiensis and also perhaps to a second generation in G. pallida.     

Observations on the distribution of potato cyst-nematodes, Globodera rostochiensis and G. pallida on two fenland potato farms

The distribution of potato cyst-nematodes, Globodera rostochiensis and G. pallida was surveyed in two widely separated fenland potato farms in Eastern England. Evidence was obtained of the suppression of G. rostochiensis and increase of G. pallida populations when Maris Piper and other potatoes with the same gene (H₁) for resistance to G. rostochiensis are grown. Numbers of G. rostochiensis and/or G. pallida in the soil could not be related directly to recent potato cropping histories of the fields. Identification of the two species of Globodera by the presence and colour of females on the roots of Désirée, Maris Piper, Cromwell (A27/20) and A25/11 potatoes were confirmed by agarose gel isoelectric focussing of general proteins. Cromwell exhibited little or no resistance to G. pallida at Woodwalton. A25/11 was more resistant to this species at Woodwalton but was susceptible to G. rostochiensis. The results of these surveys emphasise the need for carefully planned and integrated control programmes based on the use of resistant cultivars of potato, crop rotations and appropriate nematicide use to combat the growing problem of G. pallida, especially in organic soils.     

A method for estimating the contribution of seed potatoes, machinery and soil tare in field infestations with potato cyst nematodes on a national scale

In order to make a cost benefit analysis for the management of the potato cyst nematodes Globodera rostochiensis and G. pallida, we developed a method to estimate the relative importance of three basic distribution channels of potato cyst nematodes: seed potatoes, machinery and soil tare. The baseline is determined by the area planted with potatoes, the area infested with potato cysts, the proportion of resistant potato cultivars and the distribution of cysts trough different channels. This quantification forms a basis for the evaluation of the effects of different control measures for potato cyst nematode on a national scale. The method can be useful as an example for application in other countries.     

Effects of potato genotype, oxamyl and the numbers of potato cyst-nematodes, Globodera rostochiensis or G. pallida on tuber yields and nematode increase

A new technique is described for establishing different numbers of the potato cyst-nematode Globodera rostochiensis in field soil, which leaves the soil homogeneous in nutrient status. Field plots established in this way were used to compare yield losses in four potato cultivars (Maris Piper, Pentland Crown, Pentland Dell and Désirée) associated with different numbers of G. rostochiensis. Over the range of 7.4 to 148.4 eggs g^-1 soil at planting, yield losses were 18.7% (Maris Piper), 53.2% (Désirée), 55.7% (Pentland Crown) and 63.5% (Pentland Dell). Similar results were obtained in another experiment on the same field in a different year using only lightly and heavily infested plots. Treating the seedbed soil with oxamyl before planting prevented significant injury to potatoes by G. rostochiensis but increased the yield of Pentland Dell and perhaps Désirée (but not Maris Piper or Pentland Crown) more than expected from nematode control alone. Treating heavily infested soil with such a nematicide cannot therefore be recommended as part of a valid procedure for establishing lightly and heavily infested plots for comparing tolerances of attack by potato cyst-nematodes in a range of potato genotypes. In peaty loam soils moderately or heavily infested with G. pallida, oxamyl at 5.6 kg a.i. ha^-1 incorporated into the seedbed before potatoes were planted generally increased tuber yields, though the effects varied considerably with the cultivar grown. Increase of G. pallida in these soils was controlled better by growing potatoes bred for resistance to it (ZB 35 – 29, Caxton, Santé, Morag, 11233 ab 22, Fingal, A27/23, Cromwell). Increase of G. pallida on susceptible cultivars varied greatly and Romano increased G. pallida no more than the resistant Morag. G. pallida is probably controlled best in peaty loam by growing a resistant cultivar in soil treated with a granular (non-fumigant) nematicide.     

Mapping QTLs for resistance to the cyst nematode Globodera pallida derived from the wild potato species Solanum vernei

Resistance to the potato cyst nematode (PCN) species Globodera pallida, derived from the wild diploid potato species Solanum vernei, has been investigated. This source of resistance, which is effective against all of the major pathotypes of G. pallida and Globodera rostochiensis, has been assumed to be due to several genetic factors, but it has proved difficult to deploy effectively in breeding strategies for potato cultivars. Diploid and tetraploid potato populations segregating for 'vernei' resistance were analysed. At the tetraploid level, a bulk segregant analysis (BSA) approach was employed and detected AFLP markers linked to a resistance QTL on potato linkage group V. Conventional linkage analysis of a diploid population identified QTL on linkage groups V and IX. A marker linked to a QTL on linkage group V has been converted to a single-locus PCR-based marker, which can be used to detect the presence of the QTL in diploid and tetraploid potato germplasm. Moreover, there is evidence that one of the AFLPs detected by BSA appears to be specific to an introgressed segment of DNA from S. vernei. These results are compared with those obtained from other studies on resistance to the PCN species G. pallida.     

Enhancement of the efficacy of a carbamate nematicide against the potato cyst nematode, Globodera pallida, through mycorrhization in commercial potato fields

Two experiments were conducted over 2 years in commercial potato fields in Shropshire, UK, to evaluate the compatibility of the nematicide aldicarb with commercial inocula of arbuscular mycorrhizal fungi (AMF) in the control of the potato cyst nematode Globodera pallida. The AMF used were Vaminoc (mixed-AMF inoculum), Glomus intraradices (BioRize BB-E) and G. mosseae (isolate BEG 12). In the absence of AMF, the in-soil hatch of G. pallida increased 30% (P < 0.01) from wk-2 to wk-4 after planting. Inoculation of physiologically-aged potato (cv. Golden Wonder) tubers with AMF eliminated this delay in G. pallida hatch by stimulating a mean increase of 32% (P < 0.01) in hatch within 2 wk after planting. In the aldicarb-treated plots in Experiment 1, G. pallida multiplication rate was 38% lower (P < 0.05) in roots of AMF-inoculated than noninoculated plants, but in Experiment 2, this effect was slightly lower (P = 0.07). In these plots, the single AMF inocula showed also a weak trend (P = 0.10) towards greater tuber yields relative to their noninoculated counterparts. Mycorrhization therefore appears to enhance the efficacy of carbamate nematicides against G. pallida and consequently more research is proposed to validate these findings and fully explore the potential of this model.     

Control of potato pale cyst-nematode, Globodera pallida, with a granular nematicide and partially resistant potatoes

The effects of oxamyl applied to the seedbed and growing partially resistant potatoes in controlling potato pale cyst-nematode, Globodera pallida, were assessed in eleven field experiments on sandy, peaty and silty loam soils in England from 1986 to 1988. Standardised procedures allowed valid comparisons to be made between data from the three experiments in 1987 and from the seven experiments in 1988. In soil moderately or heavily infested with G. pallida, oxamyl frequently increased tuber yields of susceptible cv. Désirée and of partially resistant potatoes and lessened crude nematode increase (Pf/Pi) significantly at some sites but not at others. Although 2.8 kg oxamyl ha“¹ often increased tuber yields significantly, it was significantly less effective than 5.6 kg in controlling G. pallida at one site and at another site G. pallida increased more than in untreated plots. Potatoes partially resistant to G. pallida were more effective that oxamyl in controlling nematode increase at some sites but not at others. Combined use of oxamyl and partially resistant potatoes was generally more effective in controlling G. pallida than either measure alone. As measured by a ‘control coefficient’ (weight of tubers over 40mm diameter (t ha“¹) 4-crude nematode increase (Pf/Pi)), the most effective integrated control of G. pallida was obtained by growing cvs Glenna, Morag or Santé in soil treated with 5.6 kg oxamyl ha^-1.     

The broad-spectrum potato cyst nematode resistance gene (Hero) from tomato is the only member of a large gene family of NBS-LRR genes with an unusual amino acid repeat in the LRR region

The Hero gene of tomato is a broad spectrum resistance gene that confers a high level of resistance to all pathotypes of the potato cyst nematodes Globodera rostochiensis and partial resistance to G. pallida. The gene was identified by map-based cloning, sequencing and complementation analysis of two susceptible tomato lines with an array of 13 overlapping cosmids spanning a total distance of 135 kb. Hero encodes a protein with a nucleotide-binding site (NBS) and a leucine-rich-repeat (LRR) domain and is a member of a gene family of 14 highly homologous genes, which are clustered within a continuous 118-kb region. The isolated Hero gene displayed resistance to various G. rostochiensis pathotypes and partial resistance to G. pallida pathotype Pa2/3 in transgenic tomato lines. None of the Hero homologues conferred resistance to G. rostochiensis pathotypes. Hero can be distinguished from its homologues by the length of a compound hexanucleotide microsatellite, which codes for a charged and repetitive amino acid domain within the LRR. We propose that the expansion of this microsatellite may be involved in the evolution of the Hero resistance gene.     

Influence of temperature on the life-cycle of Globodera spp

The potato cyst nematodes (PCN) G. rostochiensis (Woll.) and G. pallida (Stone) are the most economically important pests of potato (Solanum tuberosum L.) in the UK and are widespread in ware potato growing regions in Europe. The new EU directive 2007/33/EC which came into effect July 1, 2010 aims to control their spread and limit further increases in populations. We are investigating the role of temperature in the life cycle of PCN to assess how this effects population multiplication in relation to managing PCN. Hatching and nematode development are stages in the life cycle that are affected by temperature and thus are important life stages that can be examined to determine the impact of temperature on the length of time required for one generation to be completed and the potential for final populations to increase on different potato genetic backgrounds. In some conditions a partial or complete second generation has also been observed within the growing season. Females have been observed on the surface of tubers and "pecking" skin damage can occur which may be a result of a second generation. We are investigating the influence of temperature on the potential for a second generation or the induction of diapause.     

Genetic variability and differentiation of three Russian populations of yellow potato cyst nematode Globodera rostochiensis as revealed by nuclear markers

Genetic variability of yellow potato cyst nematode G. rostochiensis from three Russian populations (Karelia, Vladimir oblast, and Moscow oblast) was investigated using two types of nuclear markers. Using RAPD markers identified with the help of six random primers (P-29, OPA-10, OPT-14, OPA-11, OPB-11, and OPH-20), it was possible to distinguish Karelian population from the group consisting of the populations from two adjacent regions (Moscow oblast and Vladimir oblast). Based on the combined matrix, containing 294 RAPD fragments, dendrogram of genetic differences was constructed, and the indices of genetic divergence and partition (P, H, and G(st)), as well as the gene flow indices N(m) between the nematode samples examined, were calculated. The dendrogram structure, genetic diversity indices, and variations of genetic distances between single individuals in each population from Karelia and Central Russia pointed to genetic isolation and higher genetic diversity of the nematodes from Karelia. Based on polymorphism of rDNA first intergenic spacer ITS1, attribution of all populations examined to the species G. rostochiensis was proved. Small variations of the ITS1 sequence in different geographic populations of nematodes from different regions of the species world range did not allow isolation of separate groups within the species. Possible factors (including interregional transportations of seed potato) affecting nematode population structure in Russia are discussed.     

Differential control of potato cyst-nematodes, Globodera rostochiensis and G. pallida by oxamyl and the yields of resistant and susceptible potatoes in treated and untreated soils

In sandy loam infested with Globodera rostochiensis (2–95 eggs g^-1 soil) the yield of Desiree potatoes was decreased by 8·2 t ha^-1 for every increment of 20 eggs g^-1 soil. Oxamyl incorporated in the seedbed at 5 kg ha^-1 before planting prevented significant loss of yield and damage to the tubers and minimised nematode increase. Cara and Maris Piper potatoes, which were resistant and tolerant to G. rostochiensis usually responded less to oxamyl than did susceptible cultivars. In a range of cultivars, yield responses to oxamyl treatment of soil infested with G. rostochiensis often differed from those in soil infested with G. pallida. In field experiments, oxamyl controlled G. pallida less than G. rostochiensis. In pots, such differential control of the two species by oxamyl was not observed.     

Effects of rotation length and oxamyl on potato yield and the potato cyst-nematode, Globodera rostochiensis, in a sandy loam soil

In sandy loam infested with golden potato cyst-nematode, Globodera rostochiensis, oxamyl at 5.6 kg a.i. ha^-1 incorporated in the top 15 cm of the soil just before planting potatoes greatly reduced nematode population increase on susceptible cv. Désirée grown six, seven or eight years after the last susceptible potato crop, but did not significantly increase tuber yields. In four-course and two-course rotations, oxamyl also controlled increase of G. rostochiensis and greatly increased yields of both cv. Désirée and resistant cv. Maris Piper. Oxamyl maintained tuber yields in a four-course rotation at the same level as in a six to eight-course rotation. Decline of G. rostochiensis in the soil was much faster under barley in some two-course rotations than under barley in four-course rotations.     

Effects of rotation length, fungicide treatment of seed tubers and nematicide on diseases and the quality of potato tubers

In 1982 – 88, potatoes were grown in 2-, 4- and 6-course rotations with spring barley on a field infested with Globodera rostochiensis. Severity of stem canker and black scurf increased with increasing frequency of previous potato crops, and seed tuber treatment with tolclofos-methyl became less effective in controlling diseases. This suggested that previous crops had increased the amounts of soil-borne inoculum of Rhizoctonia solani. Oxamyl soil treatment increased stem canker in one year and decreased black scurf in four years. Seed tuber treatment with imazalil or prochloraz decreased stem base infection by Polyscytalum pustulans and skin spot and silver scurf on tubers. Black dot was prevalent on tubers in all years and was not affected by seed tuber treatment or previous cropping. Oxamyl increased black dot and common scab in five years and decreased % tuber dry matter in six years. Cysts of G. rostochiensis were found attached to Désirée but not to Maris Piper tubers in August. At harvest tubers of both cultivars were affected by superficial pitting and its severity was related to soil populations of G. rostochiensis at planting. This damage was controlled by oxamyl. It is suggested that the pitting developed from holes made in the tuber skin at larval invasion. In 1989, Désirée seed tubers and healthy mini tubers were planted in all plots and severity of stem canker and black scurf increased with increasing proximity of previous potato crops and with the number of previous crops. Black dot on stems and tubers was not affected by previous cropping but was much less severe in a plot that had not grown potatoes during the seven years of the experiment. The severity of common scab generally decreased as the number of preceding potato crops increased.     

Characterization of susceptibility and resistance responses to potato cyst nematode (Globodera spp.) infection of tomato lines in the absence and presence of the broad-spectrum nematode resistance Hero gene

The tomato Hero A gene is the only member of a multigene family that confers a high level (>80%) of resistance to all the economically important pathotypes of potato cyst nematode (PCN) species Globodera rostochiensis and G. pallida. Although the resistance levels of transgenic tomato lines were similar to those of the tomato line LA1792 containing the introgressed Hero multigene family, transgenic potato plants expressing the tomato Hero A gene are not resistant to PCNs. Comparative microscopy studies of in vitro infected roots of PCN-susceptible tomato cv. Money Maker, the resistant breeding line LA1792, and transgenic line L10 with Ro1 pathotype have revealed no statistically significant difference in the number of juveniles invading roots. However, syncytia (specialized feeding cells) induced in LA1792 and L10 roots mostly were found to have degenerated a few days after their induction, and a few surviving syncytia were able to support only the development of males rather than females. Thus, the ratio between males and females was biased towards males on LA1792 and L10 roots. A series of changes occur in resistant plants leading to formation of a layer of necrotic cells separating the syncytium from stellar conductive tissues and this is followed by degradation of the syncytium. Although the Hero A gene is expressed in all tissues, including roots, stems, leaves, and flower buds, its expression is upregulated in roots in response to PCN infection. Moreover, the expression profiles of the Hero A correlates with the timing of death of the syncytium.     

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.     

The white potato cyst nematode (Globodera pallida) – a critical analysis of the threat in Britain

Over the last 30 years, there has been an epidemic of the white potato cyst nematode (wPCN, Globodera pallida). It has progressively replaced the yellow species (yPCN, G. rostochiensis) throughout most of England and Wales and is now a widespread problem. As damaging populations of wPCN are enormous (>10⁹ eggs ha^?1), several crops of potato cultivars resistant only to yPCN were required to produce this change. The threat it poses is reflected in an increase in the numbers of soil samples being tested and in nematicide use, which has increased to > 25 000 ha of potatoes being treated annually. Computer modelling shows that current management of wPCN is mostly ineffective and populations will continue to increase. The multiplication rate of wPCN is inversely related to its population density at planting and, because of this, modelling shows that sufficient eggs are likely to survive to enable large populations of wPCN to “rebound” following nematicide treatment. This is supported by recent trial results showing that wPCN population increase was almost as great in nematicides‐treated plots as in the untreated. Modelling also showed that current rotations (typically potatoes once every 5 or 6 years) are too short to prevent wPCN populations from progressively increasing, even when used in conjunction with a nematicide. Similarly, except with avirulent populations, the partially resistant cultivars currently available will not prevent wPCN from increasing. However, as the effectiveness of partially resistant cultivars is independent of population density, they can be very effective when integrated with a nematicide. Unfortunately, only c. 8% of the potato area is planted with partially resistant cultivars, and much of that is in land not known to be infested with wPCN. Consequently, the current epidemic of wPCN is likely to become progressively more serious. However, many farmers are failing to recognise and respond to this threat until it is too late because of the slow rate of increase of wPCN, the difficulties of detecting small populations and the costs of nematicides. To respond to the current epidemic of wPCN, the greatest priority is to have available an increased number of commercially‐attractive partially resistant cultivars.     

Control of potato cyst nematode (Globodera rostochiensis) by nematicides and a resistant potato cultivar at four sites in Scotland

The control of potato cyst nematode (Globodera rostochiensis) by the oxime-carbamates aldicarb and oxamyl was tested in four fields in Scotland. Dazomet was tested in three of these fields and carbofuran in one. In untreated plots in the three most heavily infested fields Maris Piper (resistant) yielded better than Pentland Crown (non-resistant). All nematicides increased the yields of both potato cultivars but had a greater effect on the yield of Pentland Crown. Dazomet increased yields of tubers most. Heavy nematode infestation reduced yield of tubers more in a sandy soil than in two sandy loams. In a field with few potato cyst nematodes nematicides did not significantly affect tuber yields. Although the nematicides greatly increased yields, they were not completely effective in controlling potato cyst nematodes. In treated plots in the lightly infested field, there were more nematode eggs following a crop of Pentland Crown than before. In contrast, Maris Piper markedly decreased post-cropping populations and except at one site, where dazomet further decreased nematode numbers, combining nematicides with the resistant cultivar failed to decrease nematode numbers further. Nematicides decreased the numbers of larvae invading potato roots by up to 95%, oxamyl at 5–6 kg/ha being consistently the best treatment.     

A northward colonisation of the Andes by the potato cyst nematode during geological times suggests multiple host-shifts from wild to cultivated potatoes

The cyst nematode Globodera pallida is a major pest of potato in South America where this specialist parasite is native. To investigate its phylogeography, we have genotyped individuals from 42 Peruvian populations using mitochondrial and nuclear molecular markers. A clear south-to-north phylogeographical pattern was revealed with five well-supported clades. The clade containing the southern populations is genetically more diverse and forms the most basal branch. The large divergence among cytochrome b haplotypes suggests that they diverged before human domestication of potato. As the nematodes studied have been sampled on cultivated potato, multiple host-shifts from wild to cultivated potatoes must have occurred independently in each clade. We hypothesise that this south-to-north pattern took place during the uplift of the Andes beginning 20 My ago and following the same direction. To our knowledge, this is the first study of a plant parasite sampled on cultivated plants revealing an ancient phylogeographical pattern.