Enhancement of plant growth by Zygorrhynchus moelleri

Cucumber, tomato, two flowering annuals and a flowering perennial produced plants of greater weight in soilless compost infested with Zygorrhynchus moelleri than in uninfested compost. In Z. moelleri -infested compost the annuals commenced flowering 4 to 10 days earlier, cucumbers produced mature fruits more rapidly and a greater weight (20%) of fruit was harvested from 15-wk-old tomato plants. Leaves of 11 and 15-wk-old tomato plants grown in infested compost contained more chlorophyll than leaves from plants in uninfested compost. Z. moelleri was isolated from 99% of segments cut from roots of tomato plants grown in infested compost and it suppressed the growth of Trichoderma spp. at the root surface. Propagules of Z. moelleri remained viable in air-dried compost for at least 12 months. In axenic culture, Z. moelleri enhanced lateral root development in tomato and cucumber seedlings.     

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

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

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

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

The management of potato cyst nematodes using resistant Solanaceae potato clones as trap crops

The concept of using a range of Solanaceae potato clones as trap crops for potato cyst nematode (PCN) management was investigated. A series of field trials were undertaken from 1999 to 2002 that evaluated 10 clones of either wild Solanum potato species, breeder’s hybrid lines or commercial cultivars. All had high resistance to all known PCN pathotypes (both Globodera rostochiensis and Globodera pallida) and the ability to stimulate high levels of PCN hatch. Investigations showed potential for the development of some clones as a means of reducing high PCN field population levels and for use by organic potato producers.     

Glyphosate applied to genetically modified herbicide-tolerant sugar beet and 'volunteer' potatoes reduces populations of potato cyst nematodes and the number and size of daughter tubers

Glyphosate, applied early or later or twice to genetically modified glyphosate‐tolerant sugar beet, gave excellent control of planted ‘volunteer’ potatoes growing within the crop compared to conventional herbicide programmes with or without clopyralid. In three out of four trials, this resulted in significant reductions in the numbers of eggs and cysts of potato cyst nematodes (Globodera rostochiensis and G. pallida) where infestations were moderate (23–89 eggs g^?1 soil). In the fourth trial, which had very high initial populations (130 eggs ^?1 soil), none of the herbicide treatments had any significant effect on numbers of nematode eggs or cysts. This was probably due to competition for feeding sites, and the early death of the potatoes in all treatments caused by feeding damage by the nematodes and infection by blight, which prevented the nematodes from completing their life cycle. Glyphosate also significantly reduced the number and size of daughter tubers produced, thus helping to prevent a further volunteer problem in the next crop in the rotation. This was achieved by one or two applications of one chemical compared to 2–5 applications of cocktails of conventional herbicides.     

Potato cyst nematodes Globodera rostochiensis and G. pallida

TaxonomyPhylum Nematoda; class Chromadorea; order Rhabditida; suborder Tylenchina; infraorder Tylenchomorpha; superfamily Tylenchoidea; family Heteroderidae; subfamily Heteroderinae; Genus Globodera.BiologyPotato cyst nematodes (PCN) are biotrophic, sedentary endoparasitic nematodes. Invasive (second) stage juveniles (J2) hatch from eggs in response to the presence of host root exudates and subsequently locate and invade the host. The nematodes induce the formation of a large, multinucleate syncytium in host roots, formed by fusion of up to 300 root cell protoplasts. The nematodes rely on this single syncytium for the nutrients required to develop through a further three moults to the adult male or female stage. This extended period of biotrophy—between 4 and 6 weeks in total—is almost unparalleled in plant–pathogen interactions. Females remain at the root while adult males revert to the vermiform body plan of the J2 and leave the root to locate and fertilize the female nematodes. The female body forms a cyst that contains the next generation of eggs.Host rangeThe host range of PCN is limited to plants of the Solanaceae family. While the most economically important hosts are potato (Solanum tuberosum), tomato (Solanum lycopersicum), and aubergine (Solanum melongena), over 170 species of Solanaceae are thought to be potential hosts for PCN (Sullivan et al., 2007).Disease symptomsSymptoms are similar to those associated with nutrient deficiency, such as stunted growth, yellowing of leaves and reduced yields. This absence of specific symptoms reduces awareness of the disease among growers.Disease controlResistance genes (where available in suitable cultivars), application of nematicides, crop rotation. Great effort is put into reducing the spread of PCN through quarantine measures and use of certified seed stocks.Useful websitesGenomic information for PCN is accessible through WormBase ParaSite.     

Morphological variability of potato cyst nematodes (Globodera spp.) in the Canary Islands

Thirty populations of potato cyst nematode (Globodera spp.) from the Island of Tenerife and two populations from the UK were assessed for several morphometric and non-morphometric characters thought to discriminate between the species G. rostochiensis and G. pallida. Also 200 cysts from each population were analysed by isoelectric focusing of soluble proteins. Correlation analysis, analysis of variance and principal component analysis were used to investigate relationships between the morphometric characters, how the relationships varied between species and between populations, and which characters were most useful for discriminating between species. The two species differed significantly for each of the four morphometric characters: stylet length, fenestra length, anus-fenestra distance and the number of ridges. The stylet length and fenestra length also showed differences between populations of G. rostochiensis while stylet length and number of ridges showed differences between populations of C. pallida. In general, populations of G. pallida showed greater variation than populations of C. rostochiensis. Principal component analysis of the population means indicated that over 73% of the variation in the characters could be explained by the contrast of stylet and fenestra lengths against the anus-fenestra distance and number of ridges. A plot of the first two principal components separated the two species. Stepwise discriminant analysis provided a linear combination of these four variables which discriminated between the species. Stylet length was found to be the most useful characteristic for distinguishing the species whilst anus-fenestra distance was the least useful.     

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.     

Screening of non-tuber bearing Solanaceae for resistance to and induction of juvenile hatch of potato cyst nematodes and their potential for trap cropping

Ninety accessions of non‐tuber bearing Solanaceae were screened for (i) resistance to and (ii) stimulatory effect on juvenile hatch of potato cyst nematodes, and (iii) their growth under temperate climatic conditions. All plant species belonging to the genus Solanum tested induced hatching but this effect was most pronounced for plant species of the Solanum nigrum complex. Hatching of juveniles was hardly or not stimulated by other plant genera of the Solanaceae. Solanum sisymbriifolium combined a high hatching effect with complete resistance to both Globodera rosiochiensis and G pallida. Two S. nigrum varieties showed full resistance to G rostochiensis and a high level of resistance to G pallida. Moreover, S. sisymbriifolium and the two varieties of S. nigrum performed very well under Dutch field conditions and, therefore, they are suggested as candidate trap crops for the control of potato cyst nematodes.     

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 compatibility of the fungicide azoxystrobin with Pochonia chlamydosporia, a biological control agent for potato cyst nematodes (Globodera spp.)

The biological control potential of Pochonia chlamydosporia against root-knot ( Meloidogyne spp.) and cyst-forming ( Heterodera and Globodera spp.) nematodes is widely appreciated. In spite of this, little has been undertaken to determine the compatibility of this fungus with modern fungicides. A series of experiments were undertaken to investigate the effect of azoxystrobin on P. chlamydosporia . Initial Petri dish experiments found a significant reduction in the growth of P. chlamydosporia . EC₅₀ values were calculated at 7 and 14 mg L⁻¹, respectively, for hyphal growth and chlamydospore germination. A microcosm experiment based on counts of the colony-forming units of the re-isolated fungus showed a reduction in EC₅₀ values for azoxystrobin in soil after 12, 20 and 40 days incubation (1.25, 1.00 and 3.00 mg kg⁻¹ soil, respectively). There was evidence the fungus recovered over time in response to azoxystrobin application. This was also demonstrated in a glasshouse experiment where EC₅₀ values of 1.32 and 4.4 mg kg⁻¹ soil were obtained for 35 and 49 days after planting (DAP), respectively.     

Growth and development of plants with potential for use as trap crops for potato cyst nematodes and their effects on the numbers of juveniles in cysts

Growth and development of three plant accessions with potential for use as trap crops for potato cyst nematodes (PCN), Solanum sisymbriifolium and two varieties of S. nigrum, were studied under 12 h and 17 h photoperiods. In pot experiments, rate of plant emergence, plant height, and shoot and root mass were greater for the S. nigrum varieties ‘90‐4750‐188’ and ‘88‐4750‐061’ than for S. sisymbriifolium and markedly greater than for a S. nigrum variety found as a weed of arable fields in The Netherlands. However, the last mentioned S. nigrum variety produced the most berries. Plant height and shoot weight of all the S. nigrum varieties were greater under the longer photoperiod, whereas the root mass was hardly affected. Plant height and shoot weight of S. sisymbriifolium also were greater under the longer photoperiod but the root weight was less. Under field conditions, with sowing dates from the end of March to mid August, S. sisymbriifolium and S. nigrum‘90‐4750‐188’ grew better than S. nigrum‘88‐4750‐061’. In contrast to S. nigrum, S. sisymbriifolium appeared resistant to night frosts in autumn. The stubbles of both S. sisymbriifolium and S. nigrum showed good regrowth after cutting the plants 5 or 10 cm above the soil surface 11 wk after sowing. In a pot experiment, all the plant accessions strongly reduced the numbers of juveniles in cysts compared with flax. Tolerance to Globodera rostochiensis of S. sisymbriifolium and S. nigrum‘90‐4750‐188’ was investigated in pot experiments under glasshouse conditions in sandy soil at pH 4.8 and 6.0. At soil infestation levels ranging from 0 to 56 juveniles ml^?1 soil, S. sisymbriifolium appeared much more tolerant than S. nigrum‘90‐4750‐188’. Shoot yield of S. nigrum decreased markedly with increasing soil infestation and root weight also decreased, except at pH 4.8 and light infestation levels. Both S. sisymbriifolium and S. nigrum grew better at soil pH 4.8 than 6.0. The proportion of lateral roots in the total root mass increased in both species with increasing PCN infestation and soil pH. However, although the proportion of lateral roots in plants grown at soil pH 6.0 was greater at PCN infestations up to 14 juveniles ml^?1 soil, the proportion of laterals in S. nigrum was considerably less at PCN infestations of 56 juveniles ml^?1 soil. The proportion of PCN juveniles hatching was similar for the two species and decreased slightly with increasing initial nematode population densities.     

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.     

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.     

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.     

Resolution of natural hatching factors for golden potato cyst nematode, Globodera rostochiensis

The hatching responses of Globodera rostochiensis (golden potato cyst nematode) to purified and partially-purified preparations of natural (including the potato glycoalkaloids solanine and α-chaconine) and artificial hatching factors (HFs) were bimodal. At least 10 HFs, mostly anionic, were resolved from potato root leachate by a combination of gel permeation and ion-exchange chromatography. Whereas potato roots were the principal source of HFs, haulm leachate also contained such chemicals. Root leachate from aseptically-grown potato plants lacked several HFs which were present in conventionally-produced leachate.     

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.     

Field performance of Solanum sisymbriifolium, a trap crop for potato cyst nematodes. II. Root characteristics

Hatching of potato cyst nematodes is induced by root exudates of Solanaceae, such as Solanum sisymbriifolium, and is therefore related to root length distribution of this crop. A mathematical model was derived to relate the hatching potential to root length density (RLD). A series of field experiments was carried out to study actual root length distribution of S. sisymbriifolium in relation to shoot properties and to provide input into the model. Using a modified Poisson distribution formula for the three‐dimensional distribution of roots in a volume of soil, the relation between the zone of influence of hatching agents and the RLD could be derived. On this basis, the minimal RLD was estimated, which is needed to expose 75%, 90% or 95% of cysts to root exudates, as a function of the length of the zone of influence of hatching agents on cysts. The logarithm of the total root length showed a linear relation with the logarithms of above‐ground biomass and with leaf area index. Root diameter distribution was the same for all crops examined and independent of soil depth. Fine roots (<0.4 mm in diameter) constituted around 50% of total root length. Using a zone of influence of 1.00, 0.75 and 0.50 cm around the centre of each root, a minimal RLD for sufficient soil exploration (75%) was estimated. Depth at which that minimal RLD was exceeded was linearly related to total root length (km m^?2) and to above‐ground crop biomass, enabling estimations being made of the potential hatching efficacy as related to measurable properties of S. sisymbriifolium crops. The proposed approach to derive potential hatching effects from crop properties needs further validation; particularly, the distance of influence of root exudates is a critical factor.     

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.