The effects of nematode resistant and susceptible spring oat cultivars and aldicarb on the cereal cyst nematode Heterodera avenae and yields in contrasting soil types

The effects of the cereal cyst-nematode, Heterodera auenae Woll. on resistant and susceptible oat cultivars, with and without aldicarb treatment, were compared on a clay-with-flints soil at Rothamsted and a loamy sand at Woburn. At both sites, when H. auenae was extremely scarce, yields were not further enhanced by aldicarb. At Rothamsted aldicarb increased yields by 48–72% when H. auenae averaged 10 eggs/g soil. At Woburn, aldicarb increased yields of both susceptible and resistant varieties by 80–90% with 20 eggs/g. The resistant varieties conferred yield benefits in the following oat crop equal to the residual effects of aldicarb applied before the previous crop, demonstrating that H. auenae was wholly responsible for the yield losses. Nematode resistant oats suffered as much or more damage from root invasion by H. auenae juveniles as the susceptible varieties but the resulting decrease in nematode numbers led to considerable yield improvements in the following year. At Woburn in 1977, when formalin was an added treatment, fewer females were infected by parasitic fungi and post-crop egg numbers were greater.     

Genetic variation in natural populations of the cereal cyst nematode (Heterodera avenae Woll.) submitted to resistant and susceptible cultivars of cereals

The purpose of the present work was to study the genetic characteristics of cereal cyst nematode (Heterodera avenae) populations re-established after the long-term use of resistant oat cultivars in field conditions. Population features were analyzed through fitness components and variation in enzymatic polymorphism (esterase and malate dehydrogenase loci). The longest (6 year) use of the same resistance genes (oat cv Panema) at high frequency (Rotation IB) led to the selection of a resistance-breaking pathotype and to a decrease in viability which suggested either a founder effect or a lower reproductive potential for the new pathotype. Analysis of esterase allelic frequencies led to the conclusions that: (1) the genetic constitution of this pathotype was different from the reference population maintained on the susceptible host (oat cv Peniarth), and (2) that the esterase locus may develop a disequilibrium linkage with loci involved in virulence. Random mating was recorded at the whole-field level but not always at the single-plant level, suggesting that cultivation practices such as annual ploughing could play a major role in homogenizing subpopulations developed in the vicinity of a plant. These phenomena showed that the long-term use of highly effective resistance could provok marked genetic modifications in populations. These risks should be taken into account when deciding strategies for optimal use of resistance genes in nematode management programs.

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Root responses to cereal cyst nematode (Heterodera avenae) in hosts with different resistance genes

The development of cereal cyst nematode (CCN; Heterodera avenae ) induced syncytia in the host roots of infected resistant bread wheat ( Triticum aestivum cv. AUS10894), diploid wheat ( Aegilops tauschii ), barley ( Hordeum vulgare cv. Chebec and cv. Galleon) and in the susceptible wheat cv. Meering and barley cv. Clipper were studied over a period of 13 d. The resistance to CCN in these cereal plants is conferred by the resistance genes Cre1 in the wheat cv. AUS10894, Cre3 in A. tauschii , Ha2 in barley cv. Chebec and Ha4 in barley cv. Galleon. Anatomical observations were made on the development of the syncytia in CCN-infected wheat and barley roots, which carry each of these four sources of resistance genes. Accelerated development of the syncytia in resistant plants, especially in the barley cultivars, was observed. The sites of syncytia development in susceptible wheat and barley were also closely associated with the vascular tissues in the stele, but less so in the resistant plants. The syncytia in the infected susceptible wheat and barley were also metabolically active at day 13. By contrast, the syncytia of resistant wheat plants carrying the Cre1 or Cre3 genes remained extensively vacuolated and less metabolically active. In barley plants with the Ha2 or Ha4 genes, the syncytia appeared non-functional and in early stages of degeneration by day 13 after inoculation.     

Effect of urea and certain NPK fertilizers on the cereal cyst nematode (Heterodera avenae) on wheat

Two outdoor pot experiments were conducted in two consecutive years under outdoor conditions during the wheat growing season in Saudi Arabia to determine the effects of urea and certain compound fertilizers (NPK), compared to the effects of the nematicide fenamiphos on the cereal cyst nematode (CCN), Heterodera avenae, and wheat growth. The results showed that all of the treatments, except the fertilizer diammonium phosphate (DAP), reduced the number of nematode cysts/root system and increased (P ⩽ 0.05) the dry weight of nematode-infected wheat plants. Fenamiphos and urea resulted in the best control, followed by the NPK fertilizers. The combined application of urea and fenamiphos resulted in the most significant effect in decreasing (P ⩽ 0.05) the number of cysts/root system and increasing (P ⩽ 0.05) the growth of nematode-infected wheat plants.     

Barley segregates resistant and susceptible to the cereal cyst-nematode (Heterodera avenae Woll.)

In the first year of this experiment both nematode-resistant and susceptible segregates of spring barley were equally invaded by Heterodera avenae larvae, but few females developed in the roots of the resistant plants and numbers of eggs in the soil were fewer after the resistant crop. In the second year, a commercial, nematode-susceptible barley crop yielded significantly more grain following the resistant segregate. The resistant segregate, derived from Hordeum pallidum var 191, suppresses the development of Heterodera avenae pathotypes 1 and 2 [British nomenclature] and should be useful in areas where these pathotypes predominate.     

A cereal cyst nematode (Heterodera avenae Woll.) resistance gene transferred from Aegilops triuncialis to hexaploid wheat

 The cereal cyst nematode (Heterodera avenae) is an important root parasite of common wheat. A high level of resistance was transferred to wheat from Aegilops triuncialis (TR lines) using the cross [(T. turgidum×Ae. triuncialis)×T. aestivum]. Low fertility (3–5 viable kernels per plant) was observed during the process but the surviving hybrid plants were highly vigorous. To obtain stable resistant lines further crosses to T. aestivum were performed. The resistance in TR lines seems to be transferred from the C genome of Ae. triuncialis (genomes CCUU). Ae. triuncialis was highly resistant to the two Spanish populations of H. avenae tested, as well as to four French races and two Swedish populations. The histological analysis showed a hypersensitive reaction in the roots of a resistant TR line inoculated with the Ha71 pathotype of H. avenae, whereas well-formed syncytia were observed in the roots of the susceptible control. Resistance to the H. avenae Ha71 pathotype seemed to be inherited as determined by a single dominant factor in the crosses between resistant TR lines and susceptible cultivars. Received: 11 November 1997 / Accepted: 9 December 1997     

Influence of Heterodera avenae on winter wheat in France: experiments with resistant and susceptible varieties

A succession of oat crops resistant or susceptible to Heterodera avenue produced plots lightly or moderately infested with the pest with little risk of interaction with other soil pathogens in succeeding cereal crops. Using these plots, the effects of the nematode on growth and yield of winter wheat were studied. The nematode affected the crop throughout its development, with a marked effect on yield. The effects of H. avenue are similar to those of water stress and are accentuated to a greater or lesser extent by climatic conditions. In the field the efficacy and durability of resistant genes incorporated in wheat and oats were checked. After four-years' cultivation of oats (cv. Panema), no resistance-breaking pathotypes were detected. Some advanced wheat lines with nematode resistance were also tolerant of nematode attack.     

Cloning and characterization of two neuropeptide genes from cereal cyst nematode, Heterodera avanae from India

The cereal cyst nematode, Heterodera avenae (Wollenweber, 1924) is one of the most important plant parasitic nematodes of cereals. It is an obligate sedentary endo parasite causing considerable crop losses in wheat, barley and oats worldwide. FMRFamide-like peptides (FLPs) play critical role as neurotransmitters or neuromodulators in the nervous system and proposed as one of the important targets for the plant parasitic nematode management. Therefore, for the first time we have cloned and characterized two neuropeptide genes (flp-12 and flp-16) from the cDNA library of feeding female of H. avenae. Sequence analysis of FLPs revealed that both the neuropeptides are closely related with the parasitic as well as free-living nematodes. The flp-12 contains putative 22 residue long signal peptide at N-terminal suggesting its association with extra-cellular functions, while flp-16 does not contain signal peptide. Besides this, we have found highly conserved motif KFEFIRF in flp-12 and RFGK motif in flp-16. These two flp genes could be interesting and potential targets for functional validation to explore their utility for designing management strategies.     

Analysis of expressed sequence tags from roots of resistant soybean infected by the soybean cyst nematode.

The soybean cyst nematode (SCN) Heterodera glycines is the most devastating pest of soybean in the U.S.A. The resistance response elicited by SCN in soybean is complex, and genes involved in the response to a large extent are unknown and not well characterized. We constructed cDNA libraries made from mRNA extracted from roots of the resistant soybean Glycine max L. Merr. 'Peking' at 12 h, 2 to 4 days, and 6 to 8 days post inoculation with the soybean cyst nematode, population NL1-RHp, similar to race 3. Expressed sequence tag analysis of the libraries provides rapid discovery of genes involved in the response of soybean to the nematode. A total of 3454 cDNA clones were examined from the three libraries, of which 25 cDNAs were derived from nematode RNA. The levels of certain stress-induced genes such as SAM22 and glutathione S-transferase (GST8) were elevated in the SCN-infected roots relative to uninoculated roots. Early defense response genes, particularly ascorbate peroxidase and lipoxygenase, were abundant in the 12-h library. By 6-8 days, the expression of most of those genes was not as abundant, whereas genes coding for unknown proteins and stress-induced proteins continued to be highly expressed. These ESTs and associated information will be useful to scientists examining gene and protein interactions between nematodes and plants.     

The invasion of root tips of cereals by the cereal cyst nematode Heterodera avenue

Newly germinated seedlings of susceptible cultivars of oats, wheat, barley and rye were inoculated with second-stage juveniles of Heterodera avenae in pots of sand. Subsequent examination showed oat root tips to be more commonly invaded, and by a greater range of nematode numbers than the other cereals. A comparison of oats and barley showed that lower nematode numbers in barley were not due to a higher emigration from barley; invasion, establishment and emigration by nematodes all being greater in oats. Second-stage juveniles were more likely to migrate prior to establishment in barley than in oats.     

The effect of Heterodera avenae, on the root system of susceptible and resistant oat seedlings

The effects of invasion by the cereal cyst nematode, Heterodera avenae, on the root growth of susceptible and resistant oat seedlings were similar. Nematode invasion directly inhibited root extension and lateral root production, which was dependent upon the developmental status of the invaded root tissue and the nematode density cm-¹ root. Both responses are caused by mechanical damage due to nematode invasion leading to decreased top growth in the host. Reduced root development, decreased efficiency of water and nutrient uptake in invaded plants being indirect effects of nematode invasion.     

Development and reproductive potential of the cereal aphid Sitobion avenae on resistant wheat lines (Triticum monococcum)

The effect of wheat resistance in lines of Triticum monococcum L., on the reproductive performances of the cereal aphid (Sitobion avenae F.) was investigated. Aphids were reared from birth to adult moult either on resistant or susceptible wheat lines, and transferred as apterae to both host genotypes. The influence of these transfers on the subsequent adult weight, gonad status and reproductive performances was evaluated. Aphids transferred from resistant to susceptible plants proved able to compensate for their poor nymphal growth, mainly through additional embryo growth and an increase in the number of matured embryos within the first 10 days of their adult life. Most aphids transferred from susceptible to resistant plants died within the first week following the transfer. Their most advanced embryos matured and were born, but subsequent embryo growth was quickly reduced. The reproductive strategies adopted by S. avenae when facing plant resistance, and the hypothesis of a resistant mechanism based on a poor nutritional state of the resistant plants are discussed.     

Impact of Cre1, Cre8 and Cre3 genes on cereal cyst nematode resistance in wheat

The cereal cyst nematode (CCN; Heterodera avenae), a root disease of cereal crops, is a major economic constraint in many wheat (Triticum aestivum)-growing areas of the world. The objective of this study was to assess the impact of the Cre1, Cre8 and Cre3 genes on CCN resistance. A population of 92 doubled-haploid (DH) lines derived from a cross between wheat cvs. Frame and Silverstar as well as 1,851 wheat breeding lines were screened for CCN resistance at the Primary Industries Research Victoria (PIRVic). A second population of 9,470 wheat breeding lines was screened at the South Australian Research and Development Institute (SARDI). Cre3 had the largest impact on reducing the number of female cysts, followed by Cre1 and Cre8. There was no significant difference in number of cysts between DH lines with or without the Cre8 marker, suggesting that the marker is not perfectly linked to Cre8. The estimated heritabilities were 0.32 in the DH population, 0.48 in the PIRVic data set and 0.32 in the SARDI data set, which confirm that this is a trait of low heritability. The repeatability of CCN resistance improved with an increase in the number of plants assessed per line—up to ten. However, 85–88% of the improvement was achieved with the assessments of the first five plants.     

The global importance of the cereal cyst nematode (Heterodera spp.) on wheat and international approaches to its control

The Cereal Cyst Nematodes (CCNs) are a group of several closely related species which have been documented to cause economic yield loss on rainfed wheat production systems in several part of the world including North Africa, West Asia, China, India, Australia, America and several countries in Europe. The most commonly reported species is Heterodera avenae, however there are at least two other species H. filipjevi and H. latipons are implicated. It is well appreciated that plants under water and nutrient stress suffer greater yield loss. Control of CCNs requires maintaining nematode populations below economic thresholds. Chemicals are not environmentally sustainable or economic and the major emphasis on control has been with host genetic resistance applied with other integrated pest managent options. Unfortunately due to the number of species and pathotype variation genetic control of Cereal Cyst Nematode with plant resistance is complex. Turkey is one of the top ten wheat producers in the world and has identified these nematode as a major biotic constraint in their rainfed wheat systems. In 2001 a new joint intiative was established between CIMMYT International, the Turkish Ministry of Agriculture and (Ukurova University in Adana to understand i) the distribution of cereal nematodes on wheat; ii) assess the economic importance and improve our understanding of the population dynamics iii) culture, screen and assess known sources of resistance and identify new sources to both groups of nematodes; iv) integrate new sources of resistance into bread wheat cultivars for Turkey and International germplasm using conventional and molecular tools; v) investigate other integrated control options such as rotation and different wheat management strategies and finally vi) capacity build scientists to work in this important area. Some highlights of this work will be presented and the newly formed ICCNI - International Cereal Cyst Nematode Initative introduced.     

Identification of putative parasitism genes expressed in the esophageal gland cells of the soybean cyst nematode Heterodera glycines

Cloning parasitism genes encoding secretory proteins expressed in the esophageal gland cells is the key to understanding the molecular basis of nematode parasitism of plants. Suppression subtractive hybridization (SSH) with the microaspirated contents from Heterodera glycines esophageal gland cells and intestinal region was used to isolate genes expressed preferentially in the gland cells of parasitic stages. Twenty-three unique cDNA sequences from a SSH cDNA library were identified and hybridized to the genomic DNA of H. glycines in Southern blots. Full-length cDNAs of 21 clones were obtained by screening a gland-cell long-distance polymerase chain reaction cDNA library. Deduced proteins of 10 clones were preceded by a signal peptide for secretion, and PSORT II computer analysis predicted eight proteins as extracellular, one as nuclear, and one as plasmalemma localized. In situ hybridization showed that four of the predicted extracellular clones were expressed specifically in the dorsal gland cell, one in the subventral gland cells, and three in the intestine in H. glycines. The predicted nuclear clone and the plasmalemma-localized clone were expressed in the subventral gland cells and the dorsal gland cell, respectively. SSH is an efficient method for cloning putative parasitism genes encoding esophageal gland cell secretory proteins that may have a role in H. glycines parasitism of soybean.     

Invasion and development of the cereal cyst-nematode Heterodera avenae in roots of barley infected by the take-all fungus Gaeumannomyces graminis

Observations on the invasion and development of Heterodera avenae in relation to take-all infection of barley roots were made in three pot experiments. Post-cropping H. avenae populations were lower on fungus-infected plants, as a result of the production of fewer, smaller and less fecund cysts. The reduction in cyst numbers was associated with reduced larval invasion of take-all infected roots and with the effects of the unfavourable environment of these roots on female development, possibly through its effects on sex determination and survival of the larvae.     

Differentially expressed genes in resistant and susceptible Bombyx mori strains infected with a densonucleosis virus

We investigated variations in the gene expression of Bombyx mori following infection with a densonucleosis virus (BmDNV-Z). Two B. mori near-isogenic lines, Jingsong and Jingsong.nsd-Z.NIL, which are highly susceptible and completely resistant to BmDNV-Z, respectively, were used in this study. The infection profiles of BmDNV-Z in the midguts of the B. mori Jingsong and Jingsong.nsd-Z.NIL larvae revealed that the virus invaded the midguts of both of these strains. However, its proliferation was notably inhibited in the midgut of the resistant strain. By using the suppression subtractive hybridization method, three cDNA libraries were constructed to compare BmDNV-Z responsive gene expression between the two silkworm lines. In total, 151 differentially expressed genes were obtained. Real-time qPCR analysis confirmed that 11 genes were significantly up-regulated in the midgut of the Jingsong.nsd-Z.NIL strain following BmDNV-Z infection. Our results imply that these up-regulated genes might be involved in B. mori immune responses against BmDNV infection.