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.     

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.     

Effect of the nematicide oxamyl on the multiplication of Globodera rostochiensis and G. pallida and on the haulm growth and yield of six potato cultivars with different levels of resistance and tolerance

The effect of a pre-planting application of oxamyl on the yields of six potato cultivars was studied in co-operative field trials in 1981. Two sites were ‘uninfested’ with potato cyst nematodes (PCN), two were lightly infested (<25 eggs/g soil) and six were moderately to heavily infested (three with Globodera rostochiensis and three with G. pallida). At the uninfested and lightly infested sites oxamyl had little effect on mid-season haulm weights or on final tuber yields. At sites moderately to heavily infested with G. rostochiensis the haulm growth of all cultivars tended to be increased by a similar amount on plots treated with oxamyl, Pentland Dell being least responsive. Yield was increased by different amounts, the increases being least for cvs Cara and Maris Piper and most for Corsair and Pentland Dell. At the sites moderately to heavily infested with G. pallida Cara was again tolerant, its yield being increased little by oxamyl compared with the other cultivars. Maris Piper gave the largest yield increase. Final populations of PCN on non-resistant cultivars were reduced by oxamyl at some sites but not at others. Resistant cultivars also decreased the final numbers of PCN at most sites. Two cultivars derived from Solanum vernei with different degrees of resistance, appeared to be almost equally effective in controlling G. rostochiensis and G. pallida.     

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.     

The effect of granular nematicide incorporation depth and potato planting depth on potatoes grown in land infested with the potato cyst nematodes Globodera rostochiensis and G. pallida

Field experiments at Harper Adams, Shropshire and Wisbech, Cambridgeshire investigated the effect of nematicide incorporation and seed tuber planting depth on the yield of the potato (Solamum tuberosum L.) cultivars Estima and Maris Piper and the population control of the potato cyst nematodes Globodera rostochiensis Woll. (Skarbilovich) and G. pallida (Stone). The nematicide fosthiazate was applied at 3 kg^?1 ha and either not incorporated, or incorporated to 20 cm or 35 cm. Potatoes were mechanically planted to three depths; approximately 10 cm, 15 cm and 25 cm. Incorporation to 20 cm with tubers planted at a depth of 10 cm or 15 cm, reduced root invasion compared with the other treatments. Incorporating nematicide to 20 cm also gave consistently higher ware yields and better nematode control than the other incorporation methods, which were not significantly different to the control. However, ware yield and nematode multiplication rate were not significantly affected by planting depth.     

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.     

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.     

Emergence of juvenile potato cyst-nematodes Globodera rostochiensis and G. pallida and the control of G. pallida

Speed of emergence of juveniles from cysts in potato root diffusate (PRD) in vitro differed between Globodera rostochiensis and G. pallida and between populations within each species. Early emergence in vitro was slower in most populations of G. pallida than in most populations of G. rostochiensis. Fewer G. rostochiensis juveniles emerged from 4 or 6 month old than from 4 yr old cysts. More G. rostochiensis emerged in solutions of sodium metavanadate at concentrations of 10^-2 and 10^-3 M than in PRD and as many G. pallida emerged in the same solutions as in PRD. In plots of bare fallowed sandy loam, emergence of G. pallida was stimulated by 10^--³ M sodium metavanadate. The emergence of two populations of C. pallida in PRD was stimulated by the addition of benomyl at 0.1 ppm (3.4 × 10^--⁷ m). In microplots, cv. Cara potatoes grown for 8 wk decreased four populations of G. pallida by up to 93%. During a 4 wk period in PRD, more than 20 juveniles per gravid female emerged from five of 25 populations of G. pallida. In root observation boxes in which cv. Désirée was grown, oxamyl applied to the top 15 cm of a peaty loam soil greatly increased G. pallida in soil 1545 cm deep. In another peaty loam, but not in a sandy loam, the same treatment appeared to increase the nematode in soil 15–30 cm deep. Oxamyl incorporated in the uninfested top 15 cm of all three soils largely prevented nematode increase from juveniles migrating upwards from untreated heavily infested soil 15–30 cm deep. These experiments suggest that inadequate control of G. pallida increase on susceptible potatoes by an oximecarbamate nematicide of short persistence, such as oxamyl, is primarily due to the slow rate of juvenile emergence in most populations of G. pallida, with a second generation and the upward migration of juveniles from deeper untreated soil later in the growing season as potential contributory factors.     

Potato cyst nematodes in England and Wales - occurrence and distribution

Potato cyst nematodes (PCN) have been known to occur in the UK for nearly a hundred years. They are the most problematic pests of potatoes and can cause severe yield losses. Previous work has shown the two species, Globodera rostochiensis and G pallida, to be distributed throughout the UK. This paper reports the results of the first structured and statistically unbiased survey undertaken to assess their occurrence and distribution in the potato growing land of England and Wales. The survey showed that PCN were present in 64% of sites sampled. Of the populations found, 67% were G pallida, 8% were G rostochiensis and 25% contained both species. The results show an increase in the incidence of PCN since previous studies were completed and confirm the perceived shift towards G pallida as the predominant species. Of the infestations found, 62% had a population density of less than 10 eggs g^?1 soil.     

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.     

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.     

Characterisation of a 34 kD protein from potato cyst nematodes, using monoclonal antibodies with potential for species diagnosis

Two monoclonal antibodies, which differentially recognise the two species of potato cyst nematodes (PCN), Globodera pallida and G. rostochiensis, are described. They have been shown to have potential for quantification of these two species, recognising proteins of the same molecular weight (34 kD) in both species. Further investigation showed these proteins to have isoelectric points at pH values of 5.7 in G. pallida and 5.9 in G. rostochiensis, in common with the proteins used by Fleming & Marks (1983) to differentiate the species of PCN. They are likely to be structurally very similar, with the same physiological function (and therefore similar concentrations) in the two species. In cross-reactivity tests with a wide range of soil nematode species, the antibodies reacted strongly only with species of the genus Globodera, and thereby confirmed their potential as the basis of a quantitative immunoassay likely to be useful in management of PCN populations.     

The effect of partially resistant potato clones and aldicarb on the multiplication of the potato cyst nematode Globodera pallida in field trials

Potato cultivars with different degrees of resistance to Globodera pallida (PCN) were trialled at three infested sites in 1983, 1984 and 1985. These trials were primarily intended to assess the effects of PCN damage on the yield of tubers and consequently the sites chosen had high initial population densities of G. pallida. The population density of G. pallida was determined from samples taken both pre-planting and post-harvest and results showed that there were significant differences between the clones in the nematode multiplication rates. Although actual rates of multiplication varied between sites the relative differences between clones were maintained across sites. Under the susceptible control cultivars the populations generally increased even in the presence of nematicide. The partially resistant clones gave best control, as measured by nematode multiplication, at the sites with the highest initial population densities but gave good control at all sites when treated with nematicide irrespective of initial population density. The role of partial resistance in integrated control systems and the effectiveness of nematicides against G. pallida is discussed.     

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.     

Site-Specific Management of Nematodes Pitfalls and Practicalities

The greatest constraint to potato production in the United Kingdom (UK) is damage by the potato cyst nematodes (PCN) Globodera pallida and G. rostochiensis. Management of PCN depends heavily on nematicides, which are costly. Of all the inputs in UK agriculture, nematicides offer the largest potential cost savings from spatially variable application, and these savings would be accompanied by environmental benefits. We mapped PCN infestations in potato fields and monitored the changes in population density and distribution that occurred when susceptible potato crops were grown. The inverse relationship between population density before planting and multiplication rate of PCN makes it difficult to devise reliable spatial nematicide application procedures, especially when the pre-planting population density is just less than the detection threshold. Also, the spatial dependence found suggests that the coarse sampling grids used commercially are likely to produce misleading distribution maps.     

Comparative Responses of Globodera rostochiensis and G. pallida to Hatching Chemicals

Globodera rostochiensis and G. pallida responded similarly to hatch stimulation by potato root leachate, but proportionally more second-stage juveniles (J2s) of G. rostochiensis hatched than of G. pallida in response to picrolonic acid, sodium thiocyanate, alpha-solanine, and alpha-chaconine. Fractionation of the potato root leachate identified hatching factors with species-selective (active toward both species but stimulating greater hatch of one species than the other), -specific (active toward only one species), and -neutral (equally active toward both species) activities. In a comparison of two populations of each of the two potato cyst nematode (PCN) species, however, greater similarity in response to the individual hatching factors was observed among populations of different species produced under the same conditions than among different populations of the same PCN species. Smaller numbers of species-specific and species-selective hatching factor stimulants and hatching inhibitors than of hatching factors were resolved. In a study to determine whether the different hatching responses of the two species to the same root leachate were associated with different ratios of species-selective and species-specific hatching factors, G. rostochiensis pathotype Ro1 exhibited greater hatch than did G. pallida pathotype Pa2/3 in response to leachate from older plants (more than 38 days old), while G. pallida exhibited greater hatch in response to leachate from younger plants (less than 38 days old); the response of G. pallida pathotype Pal with respect to plant age was intermediate between the other two populations. Combined molecular exclusion-ion exchange chromatography of the root leachates from plants of different ages revealed an increase in the proportion of G. rostochiensis-specific and -selective hatching factors as the plants aged.     

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.     

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.     

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.     

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.