J. P. Thompson

Root-lesion nematodes (Pratylenchus thornei and P. neglectus): a review of recent progress in managing a significant pest of grain crops in northern Australia

Two species of root-lesion nematode (predominantly Pratylenchus thornei but also P. neglectus) are widespread pathogens of wheat and other crops in Australia’s northern grain belt, a subtropical region with deep, fertile clay soils and a summer-dominant rainfall pattern. Losses in grain yield from P. thornei can be as high as 70% for intolerant wheat cultivars. This review focuses on research which has led to the development of effective integrated management programs for these nematodes. It highlights the importance of correct identification in managing Pratylenchus species, reviews the plant breeding work done in developing tolerant and resistant cultivars, outlines the methods used to screen for tolerance and resistance, and discusses how planned crop sequencing with tolerant and partially resistant wheat cultivars, together with crops such as sorghum, sunflower, millets and canaryseed, can be used to reduce nematode populations and limit crop damage. The declining levels of soil organic matter in cropped soils are also discussed with bl]Reference to their effect on soil health and biological suppression of root-lesion nematodes.

QTL mapping of multiple foliar disease and root-lesion nematode resistances in wheat

A genetic linkage map, based on a cross between the synthetic hexaploid CPI133872 and the bread wheat cultivar Janz, was established using 111 F1-derived doubled haploid lines. The population was phenotyped in multiple years and/or locations for seven disease resistance traits, namely, Septoria tritici blotch (Mycosphaeralla graminicola), yellow leaf spot also known as tan spot (Pyrenophora tritici-repentis), stripe rust (Puccinia striiformis f. sp. tritici), leaf rust (Puccinia triticina), stem rust (Puccinia graminis f. sp. tritici) and two species of root-lesion nematode (Pratylenchyus thornei and P. neglectus). The DH population was also scored for coleoptile colour and the presence of the seedling leaf rust resistance gene Lr24. Implementation of a multiple-QTL model identified a tightly linked cluster of foliar disease resistance QTL in chromosome 3DL. Major QTL each for resistance to Septoria tritici blotch and yellow leaf spot were contributed by the synthetic hexaploid parent CPI133872 and linked in repulsion with the coincident Lr24/Sr24 locus carried by parent Janz. This is the first report of linked QTL for Septoria tritici blotch and yellow leaf spot contributed by the same parent. Additional QTL for yellow leaf spot were detected in 5AS and 5BL. Consistent QTL for stripe rust resistance were identified in chromosomes 1BL, 4BL and 7DS, with the QTL in 7DS corresponding to the Yr18/Lr34 region. Three major QTL for P. thornei resistance (2BS, 6DS, 6DL) and two for P. neglectus resistance (2BS, 6DS) were detected. The recombinants combining resistance to Septoria tritici blotch, yellow leaf spot, rust diseases and root-lesion nematodes from parents CPI133872 and Janz constitute valuable germplasm for the transfer of multiple disease resistance into new wheat cultivars.

Progress in breeding wheat for tolerance and resistance to root-lesion nematode (Pratylenchus thornei)

Root-lesion nematode (Pratylenchus thornei) is a serious pathogen of wheat and other crops and is estimated to cost the Australian wheat industry

Identification of quantitative trait loci for resistance to two species of root-lesion nematode (Pratylenchus thornei and P. neglectus) in wheat

36M per year in lost production. Use of tolerant and resistant wheat cultivars is the most important component of integrated control of this nematode. A targeted approach has been adopted to develop such cultivars for the northern grain belt of Australia. Methods are described that have been used successfully to screen for tolerance in the field and resistance in the glasshouse. Backcross programs used to incorporate superior tolerance into locally adapted cultivars have doubled tolerance levels, which translates into double the grain yield on severely infested sites. Resistance from the bread wheat line GS5Oa backcrossed into locally adapted cultivars has reduced the nematode multiplication rate more than tenfold in advanced lines. New sources of resistance have been located in Aegilops tauschii, in durum wheat and in synthetic hexaploid wheats derived from them. Other new sources have been identified in landrace wheats and wheat cultivars from the Middle East and North Africa. A concerted effort is required to exploit these new found resistances and knowledge to accelerate the production of suitable cultivars for farmers’ use.

Occurrence of root-lesion nematodes (Pratylenchus thornei and P. neglectus) and stunt nematode (Merlinius brevidens) in the northern grain region of Australia

Pratylenchus thornei and P. neglectus are two species of root-lesion nematode that cause substantial yield losses in wheat. No commercially available wheat variety has resistance to both species. A doubled-haploid population developed from a cross between the synthetic hexaploid wheat line CPI133872 and the bread wheat Janz was used to locate and tag quantitative trait loci (QTLs) associated with resistance to both P. thornei and P. neglectus. Wheat plants were inoculated with both species of nematode in independent replicated glasshouse trials repeated over 2 years. Known locations of wheat microsatellite markers were used to construct a framework map. After an initial single-marker analysis to detect marker-trait linkages, chromosome regions associated with putative QTLs were targetted with microsatellite markers to increase map density in the chromosome regions of interest. In total, 148 wheat microsatellite markers and 21 amplified fragment length polymorphism markers were mapped. The codominant microsatellite marker Xbarc183 on the distal end of chromosome 6DS was allelic for resistance to both P. thornei and P. neglectus. The QTL were designated QRlnt.lrc-6D.1 and QRlnn.lrc-6D.1, for the 2 traits, respectively. The allele inherited from CPI133872 explained 22.0-24.2% of the phenotypic variation for P. thornei resistance, and the allele inherited from Janz accounted for 11.3-14.0% of the phenotypic variation for P. neglectus resistance. Composite interval mapping identified markers that flank a second major QTL on chromosome 6DL (QRlnt.lrc-6D.2) that explained 8.3-13.4% of the phenotypic variation for P. thornei resistance. An additional major QTL associated with P. neglectus resistance was detected on chromosome 4DS (QRlnn.lrc-4D.1) and explained a further 10.3-15.4% of the phenotypic variation. The identification and tagging of nematode resistance genes with molecular markers will allow appropriate allele combinations to be selected, which will aid the successful breeding of wheat with dual nematode resistance.

Mapping quantitative trait loci for resistance to Pratylenchus thornei from synthetic hexaploid wheat in the International Triticeae Mapping Initiative (ITMI) population

Pratylenchus thornei, P. neglectus and Merlinius brevidens were respectively present in soil from 67, 32 and 73% of 795 wheat fields sampled in the northern grain region of Australia. Population densities of P. thornei (31% of samples with >2000 nematodes/kg soil) were higher than those of P. neglectus (5%) or M. brevidens (11%). The mean proportion of plant-parasitic to total nematodes was 0.40. The incidence of P. thornei and P. neglectus increased significantly with the number of years that fields had been cropped since clearing natural vegetation. All three species occurred in soil samples across a wide range of texture and pH. Maximal incidence of P. thornei was in heavier textured soils (48% clay, 37% sand) than P. neglectus (39% clay,45%sand). The incidence of M. brevidens increased linearly with increasing clay and decreasing sand contents. M. brevidens occurred in the pH range 6.0–9.5 and the Pratylenchus spp. in the range 6.5–9.5, with maximal incidence of P. neglectus at pH 8.2, P. thornei 8.4 and M. brevidens 8.8. Population densities of P. neglectus and especially non-parasitic nematodes were higher in the 0–15cm soil layer than 15–30 cm. Overall the soil conditions of the northern grain region appear more favourable for P. thornei and M. brevidens than for P. neglectus.

Resistance to root-lesion nematodes (Pratylenchus thornei and P. neglectus) in synthetic hexaploid wheats and their durum and Aegilops tauschii parents

Root-lesion nematode (Pratylenchus thornei) is a serious pathogen of wheat in many countries. The International Triticeae Mapping Initiative (ITMI) population of recombinant inbred lines (RILs) was assessed for resistance to P. thornei to determine the chromosome locations of the resistance genes. The ITMI population is derived from a cross between the resistant synthetic hexaploid wheat W-7984 and a susceptible bread wheat cultivar Opata 85. Two years of phenotypic data for resistance to P. thornei were obtained in replicated glasshouse trials. Quantitative trait locus (QTL) analysis was performed using available segregation and map data for 114 RILs. A QTL on chromosome 6DS showed consistent effects for reduced nematode numbers (partial resistance) across years and accounted for 11% and 23% of the phenotypic variation. A second QTL for P. thornei resistance on chromosome 2BS accounted for an additional 19% and 5%. Restriction fragment length polymorphism (RFLP) and simple sequence repeat (SSR) markers associated with the QTLs are physically located in regions rich in major genes at the distal ends of the short chromosome arms of 6D and 2B. SSR markers with potential for marker-assisted selection of P. thornei resistance effective in different genetic backgrounds have been identified.

Effects of vesicular-arbuscular mycorrhizae on growth and phosphorus and zinc nutrition of peanut (Arachis hypogaea L.) in an Oxisol from subtropical Australia

Root-lesion nematodes (Pratylenchus thornei Sher and Allen and P. neglectus (Rensch) Filipijev and Schuurmans Stekhoven) cause substantial yield loss to wheat crops in the northern grain region of Australia. Resistance to P. thornei for use in wheat breeding programs was sought among synthetic hexaploid wheats (2n= 6x = 42, AABBDD) produced through hybridisations of Triticum turgidum L. subsp. durum (Desf.) Husn (2n= 4x = 28, AABB) with Aegilops tauschii Coss. (2n= 2x = 14, DD). Resistance was determined for the synthetic hexaploid wheats and their durum and Ae. tauschii parents from the numbers of nematodes in the roots of plants grown for 16 weeks in pots of pasteurised soil inoculated with P. thornei. Fifty-nine (32%) of 186 accessions of synthetic hexaploid wheats had lower numbers of nematodes than Gatcher Selection 50a (GS50a), a partially resistant bread wheat. Greater frequencies of partial resistance were present in the durum parents (72% of 39 lines having lower nematode numbers than GS50a) and in the Ae. tauschii parents (55% of 53 lines). The 59 synthetic hexaploids were re-tested in a second experiment along with their parents. In a third experiment, 11 resistant synthetic hexaploid wheats and their F-1 hybrids with Janz, a susceptible bread wheat, were tested and the F(1)s were found to give nematode counts intermediate between the respective two parents. Synthetic hexaploid wheats with higher levels of resistance resulted from hybridisations where both the durum and Ae. tauschii parents were partially resistant, rather than where only one parent was partially resistant. These results suggest that resistance to P. thornei in synthetic hexaploid wheats is polygenic, with resistances located both in the D genome from Ae. tauschii and in the A and/or B genomes from durum. Five synthetic hexaploid wheats were selected for further study on the basis of (1) a high level of resistance to P. thornei of the synthetic hexaploid wheats and of both their durum and Ae. tauschii parents, (2) being representative of both Australian and CIMMYT (International Maize and Wheat Improvement Centre) durums, and (3) being representative of the morphological subspecies and varieties of Ae. tauschii. These 5 synthetic hexaploid wheats were also shown to be resistant to P. neglectus, whereas GS50a and 2 P. thornei-resistant derivatives were quite susceptible. Results of P. thornei resistance of F(1)s and F(2)s from a half diallel of these 5 synthetic hexaploid wheats, GS50a, and Janz from another study indicate polygenic additive resistance and better general combining ability for the synthetic hexaploid wheats than for GS50a. Published molecular marker studies on a doubled haploid population between the synthetic hexaploid wheat with best general combining ability (CPI133872) and Janz have shown quantitative trait loci for resistance located in all 3 genomes. Synthetic hexaploid wheats offer a convenient way of introgressing new resistances to P. thornei and P. neglectus from both durum and Ae. tauschii into commercial bread wheats.

Resistance to the root-lesion nematode Pratylenchus thornei of Iranian landrace wheat

Peanut plants (cv. Shulamit) were grown in an Oxisol soil in pots in the glasshouse to assess effects of soil sterilization and inoculation with spores of vesicular-arbuscular mycorrhizal fungi (VAMF) on the response to five rates of phosphorus (0 to 240 kg P ha−1) and two rates of zinc (0 and 10 kg Zn ha−1) fertilizers.Both P and Zn nutrition were affected by VAMF activity but the dominant role of VAMF in this soil type was in uptake of P. In the absence of VAMF there was a clear threshold level of P application (60 kg P ha−1) below which plants grew poorly, which resulted in a sigmoidal response of dry matter to applied P. The maximum response was not fully defined because dry matter production continued to increase up to 240 kg P ha−1. Tissue P concentration of non-mycorrhizal plants increased linearly with P rate and was always significantly less than that in mycorrhizal plants.Mycorrhizal plants responded without threshold to increasing P rate, attaining maximum dry matter at 120 kg P ha−1 in inoculated sterilized soil and at 30 kg P ha−1 in nonsterile soil. These differences in maximal P rates and in the greater dry matter produced in sterile soil at high P rates were attributed to the negative effects of the root-knot nematodeMeloidogyne hapla in nonsterile soil.Plant weight did not respond to zinc fertilizer but tissue Zn concentration increased with applied Zn. Tissue Zn concentration and uptake were increased by VAMF.

Resistance to the root-lesion nematode Pratylenchus thornei in wheat landraces and cultivars from the West Asia and North Africa (WANA) region.

Pratylenchus thornei is widespread throughout the wheat-growing regions in Australia and overseas and can cause yield losses of up to 70% in some intolerant cultivars. The most effective forms of management of P. thornei populations are crop rotation and plant breeding. There have been no wheat accessions identified as completely resistant to P. thornei, therefore breeding programs have used moderately resistant parents. The objective of the present research was to evaluate 274 Iranian landrace wheats for resistance to P. thornei and identify accessions with resistance superior to the current best resistance source (GS50a). Plants were grown in P. thornei inoculated soil under controlled conditions in a glasshouse pot experiment for 16 weeks. Ninety-two accessions found to be resistant or moderately so were retested in a second experiment. From combined analysis of these experiments, 34 accessions were identified as resistant with reproduction factors (final population per kg soil/initial inoculum rate per kg soil) ≤1. In total, 25 accessions were more resistant than GS50a, with AUS28470 significantly (P<0.05) more resistant. The resistant Iranian landraces identified in the present study are a valuable untapped genetic pool offering improved levels of P. thornei resistance over current parents in Australian wheat-breeding programs.