The effect of pyramiding two potato cyst nematode resistance loci to Globodera pallida Pa2/3 in potato

Globodera pallida is a major nematode pest causing severe constraints in many potato production regions worldwide. The most prevalent G. pallida pathotypes are Pa2 and Pa3, which exist in mixed populations referred to as Pa2/3. Due to heterogeneity for avirulence genes within these nematode populations, so far, breeders have failed to identify a single resistance gene source which offers complete resistance. Currently, there are two quantitative trait loci (QTL) available, $ GpaIV_{adg}^{s} $ and Gpa5, conferring partial levels of resistance to G. pallida pathotype Pa2/3. The objective of this research was to pyramid $ GpaIV_{adg}^{s} $ and Gpa5 through marker-assisted selection to investigate whether or not pyramiding provides increased resistance to G. pallida Pa2/3. We developed a population segregating for both resistance QTLs and, using diagnostic genetic markers (Contig237 and HC), we identified groups of individuals from this population containing each individual QTL, both QTLs simultaneously, and neither QTL. These individuals were assessed for their resistance levels against G. pallida Pa2/3 population Chavornay. We demonstrated that individuals carrying both QTLs showed a significant reduction in the number of cysts formed in comparison to genotypes carrying $ GpaIV_{adg}^{s} $ or Gpa5, indicating an additive effect. Overall these results show that MAS-based pyramiding of these QTLs is an effective strategy for breeding cultivars exhibiting very high levels of 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).     

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.     

Observations on the population dynamics of Globodera pallida under single and double cropping conditions

The population dynamics of Globodera pallida were studied on susceptible potato plants. Oxamyl treated and untreated plots were planted with a G. pallida susceptible crop, from the first early maturity class, under polythene at the end of February and immediately followed, after harvest in June, by a susceptible second crop. Other nematicide treated and untreated plots were sown in April with a G. pallida susceptible cultivar. The results were variable but showed that reproduction of G. pallida under double cropping conditions was greater than on a single main crop.     

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.     

Analyses of the effects of potato cyst nematodes (Globodera pallida) on growth, physiology and yield of potato cultivars in field plots at three levels of soil compaction

Field experiments were carried out in 1991 and 1992 on sandy soil highly infested with the potato cyst nematode Globodera pallida. Half the trial area was fumigated with nematicide to establish two levels of nematode density. Three levels of soil compaction were made by different combinations of artificial compaction and rotary cultivation. Two potato cultivars were used in 1991 and four in 1992. Both high nematode density and soil compaction caused severe yield losses, of all cultivars except cv. Elles which was tolerant of nematode attack. The effects of the two stress factors were generally additive. Analysis of the yield loss showed that nematodes mainly reduced cumulative interception of light while compaction mainly reduced the efficiency with which intercepted light was used to produce biomass. This indicates that nematodes and compaction affect growth via different damage mechanisms. Nematodes reduced light interception by accelerating leaf senescence, by decreasing the specific leaf area and indirectly by reducing overall crop growth rate. Partitioning of biomass between leaves, stems and tubers was not affected by nematode infestation but compaction decreased partitioning to leaves early in the growing season while increasing it during later growth stages. The effects of nematodes and compaction on root length dynamics and nutrient uptake were also additive. This suggests that the commonly observed variation in yield loss caused by nematodes on different soil types is not related to differences in root system expansion between soils of various strength. Cv. Elles, which showed tolerance of nematodes by relatively low yield losses in both experiments, was characterised by high root length density and thick roots. These characteristics did not confer tolerance of soil compaction, since compaction affected root lengths and tuber yields equally in all cultivars. In the first experiment only, high nematode density led to decreased root lengths and lower plant nutrient concentrations. The yield loss which occurred in the second experiment was attributed to the effects of nematodes on other aspects of plant physiology.     

The effect of oxamyl and the growing of susceptible and resistant potato cultivars on the population dynamics of Globodera rostochiensis throughout the soil profile

Comparisons are made between the population dynamics of potato cyst nematode and root growth of Pentland Crown and Maris Piper potato cultivars. Large changes in the number of eggs occurred to a depth of 48 cm, particularly in a peaty loam and in plots treated with oxamyl. Oxamyl delayed the hatch of eggs in the peaty loam but not in the sandy loam, giving the protected plants several weeks without invasion damage during which they became bigger with their roots better established deeper in the soil. Oxamyl killed or impaired the movement of second stage juveniles but appeared to have little systemic activity to hinder juvenile development within roots.     

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.     

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.     

Effects of three rates of aldicarb and of different degrees of resistance and tolerance in potato cultivars on yield and post-harvest population densities of the potato cyst nematodes Globodera rostochiensis and G. pallida

Three field experiments were made to determine the effectiveness of small-plot trials in detecting differences between potato cultivars/clones in their tolerance of damage by potato cyst-nematodes. A nematicide (aldicarb) was applied at three rates to decrease nematode damage. The largest rate of aldicarb increased tuber yields most but the relationship between yield response and nematicide rate was not linear. The yield increases of the cultivars and clones differed, indicating that they have different degrees of tolerance of potato cyst nematodes. The results were analysed in several ways and the untreated yield as a proportion of the treated provided the best means of expressing and comparing tolerance; but whichever method was used the tolerance rankings of the cultivars and clones were similar. At two sites infested with Globodera rostochiensis, the rankings of the 10 cultivars and clones were similar but at a third site, heavily infested with G. pallida, they were different. Aldicarb decreased the nematode population density after harvest at the G. pallida site but was less effective at the G. rostochiensis sites, which were less heavily infested. Growing resistant or partially resistant potatoes usually prevented nematode increase, and the more resistant cultivars and clones decreased population densities markedly.     

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.     

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.     

Influence of nematicide, resistant and susceptible potato cultivars and bare fallow on the population dynamics of Globodera rostochiensis Woll. Rol under field conditions in Spain

The susceptible potato cultivar Désirée and the resistant cv. Diamant were cultivated on untreated and nematicide-treated plots on two fields in northern Spain infested with Globodera rostochiensis Rol. Initial and final infestations of cysts and viable eggs were assessed. Multiplication rates differentiated more accurately the effects of the different treatments than final number of cysts and eggs, due to the elimination of some of the errors associated with variability of the initial infestation. Many new cysts were observed on all plots where susceptible potatoes had been grown, whether they had been treated with nematicide or not, although significantly fewer cysts were actually found on the nematicide-treated plots. With the resistant cultivar no significant effects of the nematicide were detected even though there was a noticeable reduction of viable cyst contents over and above the effect of the resistant cultivar.     

The effect of Solanum tuberosum×S. vernei hybrids on hatching of the potato cyst nematode Globodera pallida

The hatching activity of partially resistant Solanum tuberosum×S. vernei clones and susceptible cultivars was investigated in a series of tests with G. pallida conducted in pots in the glasshouse. Differences in hatching were apparent, with the partially resistant clones generally stimulating a lower total hatch than the S. tuberosum cultivars. It is concluded that reduced hatching activity can make a small contribution to overall resistance and perhaps also to the tolerance of a clone to G. pallida.     

A field study of the relationship between pre-planting density of Globodera pallida and the growth and yield of two potato cultivars of differing tolerance

Non-resistant but tolerant cv. Cara and non-resistant but relatively intolerant cv. Pentland Dell were grown in split plots encompassing a range of population densities of potato cyst nematode, Globodera pallida. Light interception and its efficiency of conversion were estimated by regular ground cover measurements and plant harvests. It was concluded that increasing levels of infestation with G. pallida only slightly decreased the efficiency of utilisation of intercepted radiation. Heavy infestation of G. pallida initially decreased the top growth and light interception of both cultivars by similar proportions, but in later harvests, this adverse effect markedly decreased for Cara whereas it slightly increased for Pentland Dell. This difference was due to the heavily infested Cara eventually achieving and maintaining 100% ground cover whereas the equivalent Pentland Dell never exceeded 75% ground cover. Consequently, final tuber yields were decreased much more for Pentland Dell than for Cara though the decreases in tuber yield were less than those for top growth. The importance of nematode effects on top growth, and hence on light interception, with regard to both yield losses and tolerance differences, were clearly demonstrated. Both linear and logarithmic models were used to describe the relationship between the initial population density of G. pallida and yield, and the implications of differences in tolerance on the parameters in the logarithmic model are discussed.     

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.     

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.