Integrated Control of Heterodera avenae1

Cereal cyst nematode is a major pathogen in southern Australia costing $40 to $80 million in lost production each year. Our research has shown that three chemicals applied with the seed in the drill row, viz. Counter (terbufos) granules, Vydate (oxamyl) as a seed dressing and Nemadi (ethylene dibromide), reduced Heterodera avenae damage and gave yield increases which are economical in the Australian wheat farming system. A plant assay of soil has been developed to assess potential damage by H. avenae before employing chemical control. Wheat yields 2 years after growing H. avenae-resistant Festiguay wheat were 0.4 to 1 t/ha higher than after other cultivars. Rotations with legumes alternating with wheat reduced damage from H. avenae. Wheat sown without cultivation (minimum tillage) resulted in less root damage from H. avenae and higher yields than when sown into cultivated soil.     

The Decline of Heterodera avenae Populations1

Populations of the cereal cyst nematode fail to multiply in many soils in Europe and farmers are able to grow susceptible crops intensively on infested land. Similar numbers of females develop on roots in summer in soils where numbers of the nematode increase or decline. Two fungi, Nematophthora gynophila and Verticillium chlamydosporium parasitize females on roots, prevent cyst formation, decrease fecundity, and limit nematode numbers. Formalin (38 % formaldehyde) soil drenches at 3000 1/ha reduce the activity of these fungi and populations of the nematode increase, N. gynophila is an Oomycete which infects by motile zoospores whose activity is decreased when summer rainfall is light whereas V. chlamydosporium is much less affected by soil moisture. The decline of Heterodera avenae populations is associated with large numbers of spores in soil. At present it is not possible to control the nematode by introducing these fungi into soils where they are few or absent.     

Characterization of Heterodera avenae in Spain1

Several types of Heterodera spp. belonging to the ≪ avenae ≫ group have been found in Spain on cereal crops and wild Gramineae. The morphological differences between them relate to the colour of the cysts, presence or absence and consistency of the subcrystalline layer, presence or absence and shape of the ≪ underbridge ≫ in the vulvar cone, length of the 2nd-stage larvae and shape of their stylet knobs. The value of these characters in the diagnosis of these species is discussed together with the identity of the types.     

Classification of Plants Resistant to Heterodera avenae1

Differentiation of barley plants resistant to Heterodera avenae is simple when they are completely without cysts and the number of cysts is high on a susceptible control cultivar. Problems with classification are discussed.     

Studies on the Australian Pathotype of Heterodera avenae1

Australian studies on populations of Heterodera avenae have been conducted in Victoria, Western Australia. South Australia, and New South Wales. Only one pathotype has been identified so far, and it is distinct from those recorded elsewhere. Few recognised sources of resistance in barley and oats are useable in Australian breeding programmes. The first commercially acceptable cultivars of wheat, barley, and oats resistant to cereal cyst nematode will be released in Victoria within the next year or two. Their resistances are derived from spring wheat(AUS 10894), Marocaine 079 (CI 8334), and Arena sterilis (Cc 4658) respectively.     

The Effect of Environment on Survival and Hatching of Heterodera avenae1

Eggs within cysts of Heterodera avenae are able to survive long periods of desiccation and can be held for several years at 5oC when stored at low relative humidity. While this provides an excellent means of storing the nematode for use in laboratory experiments, field experiments show that desiccated eggs are readily dispersed by wind. Tolerance to desiccation is similar in free and encysted eggs, which hatch at the same rate. When eggs are desiccated, water is withdrawn from the larva within the egg shell. If cysts are exposed to conditions favouring eclosion, subsequent desiccation reduces egg viability and larval emergence. The importance of defining pre-test conditions in survival studies is emphasized, and differences in the physiological state of eggs may explain different observations in Australian and European studies.     

Variations in Cereal Yield Losses associated with Heterodera avenae in England and Wales1

Many field experiments in England and Wales during the past 20 years have measured cereal yield losses and established regressions of yield on numbers of Heterodera avenae. Yield benefits from soil sterilants were greatest where most H avenae was present. Isogenic selections of barley and oats resistant and susceptible to H. avenae were extensively grown to assess the losses caused by this nematode alone. In some experiments these losses were identical with those measured by nematicide use, but in others (notably where broad spectrum biocides were used) losses were greater than those attributable to H. avenae and could be explained only by the known incidence of accompanying pathogens. Some yield loss may be due to migratory nematodes, e.g. Pratylenchus spp. occurring with H. avenae. Barley and wheat yields in the last 30 years have increased by 63 and 84 % due to improvements in cultivars and farming practice. Smaller percentage loss in a high yielding crop can be as costly as large percentage loss in poor crops grown on dry light soils. Many factors - soil type, rainfall, nutrients, other pathogens etc. - can greatly modify crop responses to similar popuiation levels of H. avenae.     

The Status of Heterodera avenae on Cereals in New Zealand1

The cereal cyst nematode (Heterodera avenae) was discovered in New Zealand in 1975 on wheat and oats. It was not at that time causing appreciable economic losses in yield, presumably because usual management practice incorporated a good crop rotation with lucerne. However, now that the growing of cereals has become more profitable, farmers are tending to grow consecutive cereal crops and more damage from H. avenae is being experienced. On one farm, up to 9 % of an oat crop was lost due to this nematode. The biotype has not yet been determined.     

青海省小麦孢囊线虫病发生和分布特点

小麦孢囊线虫病是青海省小麦青稞产区的主要病害之一。2007-2010年,采用随机取样方法对青海省小麦孢囊线虫病的发生、分布情况进行了调查。调查结果表明,小麦孢囊线虫病主要分布在青海省西宁、海东、海北、黄南、玉树、海西州6个地区,在不同海拔、不同生态区均有分布,但不同生态区以及同一生态区不同地块间发生量差异较大。垂直分布调查结果表明,同一地块中禾谷孢囊线虫主要分布在20 cm以上的土层中。     

甘肃省小麦禾谷孢囊线虫的发生及分布

为了解甘肃省小麦孢囊线虫病发生程度和分布情况,2009-2012年于小麦抽穗至灌浆期,采用Z字形取样法,取根际0~20cm土壤,用漂浮法分离孢囊,统计100g土样的孢囊数量,用形态学方法鉴定线虫。从甘肃省12个地区50个县区,134个乡镇共采集到463个小麦田间土样。结果表明,孢囊平均检出率为47.9%,其中冬麦区孢囊平均检出率为61.5%,春麦区平均检出率为34.2%,冬麦和春麦混播区为52.3%,冬麦、春麦及冬春麦混播区平均孢囊数分别为3.5、6.8和3.9个/100g土样。其中平凉市灵台县和兰州市永登县孢囊线虫发生最为严重,个别地块每100g土样中孢囊数超过40个。根据孢囊形态学特征的观察,将甘肃省采集到的孢囊线虫鉴定为禾谷孢囊线虫(Heterodera avenae)。研究结果可为有效防治甘肃省麦类孢囊线虫病提供依据。     

燕麦孢囊线虫在田间的水平和垂直分布

为明确燕麦孢囊线虫在田间的水平和垂直分布,分别在小麦收获后玉米播种前和小麦播种前土壤翻耕后进行水平和垂直取样调查。调查结果表明,燕麦孢囊线虫在田间水平分布主要为聚集分布,翻耕对其水平分布影响不大; 小麦收获后翻耕前燕麦孢囊线虫在田间垂直分布主要分布于5～10cm处,占取样深度土层孢囊总数量的35.8%,翻耕后燕麦孢囊线虫的垂直分布主要在0～15cm的土层发生变化,0～10cm的孢囊数量下降而10～15cm的孢囊数量增加。在0～20cm土壤中,翻耕前与翻耕后孢囊数量分别占孢囊总数量的89.4%和88.6%。     

禾谷孢囊线虫与不同小麦根系的互作表型特征

为了明确高抗品种‘华麦1号’的抗禾谷孢囊线虫机制,以Pluronic F-127胶体为介质,比较了禾谷孢囊线虫2龄幼虫(J2)侵入根系前对抗病品种‘华麦1号’与感病品种‘豫麦34’和‘矮抗58’的根尖趋性差异,并采用室内人工接种法观察了线虫侵入3个品种后的发育进程。结果表明,J2对3个品种的根尖均表现明显的趋性,对‘矮抗58’的趋性最强,而对‘华麦1号’的最弱,接种4h和6h时‘华麦1号’与‘矮抗58’根尖吸引的线虫总量差异显著(P0.05);组织染色观察到J2对3个品种的根系均有一定数量的侵入,但高抗品种‘华麦1号’根系侵入的幼虫量和后期形成的白雌虫量均显著低于感病品种‘豫麦34’和‘矮抗58’。结果证实,‘华麦1号’的抗性机制主要表现为减少线虫的有效侵入量、抑制侵入后的线虫生长发育。     

Development of two species-specific primer sets to detect the cereal cyst nematodes Heterodera avenae and Heterodera filipjevi

Twelve Heterodera species are of major economic significance in wheat and barley. Of these, H. avenae, H. filipjevi and H. latipons are among the most important ones, and sometimes coexist. The identification of Heterodera species using morphological characteristics is time consuming, requires specialized skill and can be imprecise, especially when they occur mixed in field populations. Molecular techniques can provide a more accurate way for nematode identification. This study reports the results of experiments targeting the mitochondrial cytochrome oxidase subunit 1 (COI) gene to develop species-specific primers that could be used for the identification of H. avenae and H. filipjevi. The COI gene of 9 Heterodera spp. and Punctodera punctata was partially sequenced and the resultant sequences were aligned to find unique sites suitable for the design of primers. The alignment showed variability between H. avenae, H. filipjevi and other Heterodera species. Two sets of species-specific primers were identified for the identification of both species and the conditions for their use in PCR were optimised. The specificity of the designed primers was checked by comparison with one population of P. punctata and populations of 14 other Heterodera species, nine populations of H. avenae and 10 populations of H. filipjevi originating from different countries. To test the sensitivity, the PCR was run with DNA extracted from five second-stage juveniles (J2) of H. avenae or five J2 of H. filipjevi mixed with DNA extracted from varying numbers of J2 of H. latipons. It was possible to detect as few as five J2 of H. avenae or H. filipjevi among 100 J2 of H. latipons. The two primers sets allow the detection of H. avenae and H. filipjevi where they occur in mixed populations with other Heterodera spp.     

Heredity of Heterodera avenae Resistance Originating from two Barley Cultivars and one Spring Wheat Cultivar1

Two spring barley cultivars Bajo Aragon-1-1 and Martin 403-2 from the cyst nematode test assortment were each crossed with a susceptible, a pathotype-11 -resistant, and a pathotype-11 and 12-resistant cultivar, as well as with each other. A spring wheat cultivar, AUS 10894, from the cyst nematode test assortment was crossed with a susceptible and a resistant cultivar. From these crosses F! and F₂ single plants were tested against pathotype 12 and/or pathotype 11 of Heterodera avenae in Denmark. Some plants of the spring barley cultivar Bajo Aragon-1-1 have two resistance genes, which probably are both dominant, but it is not out of the question that one might be recessive. Each gene is inherited independently of the other or with low linkage frequency. Neither gene is the same as or allelic to the resistance gene in the cultivar Ortolan, but one gene is allelic to or the same as the 191 gene in the cultivar Siri. The spring barley cv. Martin 403-2 has one dominant resistance gene which is not allelic to or the same as the resistance gene in Ortolan but is the same as or allelic to the 191 gene in the cultivar Siri. It is possible that there is one more gene, which might be recessive against pathotype 11. The spring wheat cv. AUS 10894 has one dominant resistance gene which might be the same as or allelic to the Loros gene.     

Resistance of Cereals to Heterodera avenae: Methods of Investigation, Sources and Inheritance of Resistance1

Techniques used in cereal breeding programmes for selecting for resistance to cereal cyst nematodes are described. Routine screening is done in field nurseries and in pots of infested soil in glasshouse and controlled environments. Other investigations of resistance use a laboratory technique which permits identification of the penetration sites of individual nematodes. Erosion of resistance, as indicated by 1) the development of only one or two females or 2) the occasional development of more females on plants with resistance genes, is discussed in relation to variation in nematode virulence and to environmental, especially temperature effects. Identification of tolerance is more difficult unless the differences are great, or the experimentation carefully controlled and replicated. Direct selection for tolerance in segregating generations is not presently practicable. Indirectly selecting for mechanisms of tolerance may eventually prove possible but has not yet been exploited. Available sources of resistance are described in relation to genetic studies and complementary studies of nematode pathotype and host range. There are three distinct resistances in barley which may have been responsible for selecting present pathotypes. Two, Rha1 and Rha2, are useful in breeding and linkage and allelism at these loci are described. In oats, as in wheat, only one locus has been used by breeders. There are probably additional loci in some of the other resistance sources but polyploidy of the host and different virulence gene frequencies in unselected nematode populations make it more difficult to identify genetic relationships in oats and wheat.     

Experience in Denmark with Cereal Cultivars Resistant to Cereal Cyst Nematode (Heterodera avenae Woll.)1)

On the basis of investigations on the cereal cyst nematode's distribution, it is presumed that one third of all the cereal growing areas in Denmark is infested to a degree, and that yield losses of 5 % or more may be assumed. This means an annual loss of about 100 million D.Kr. By using resistant cultivars every second year, the population of Heterodera avenae Woll. can be kept at a harmless level. It may be assumed that intensive growing of nematode-resistant cereals includes a risk of promoting the occurrence of pathotypes other than the 2 known to exist in Denmark. So far, there is no indication for their existence.     

Cultural Practices and their Effects on Heterodera avenae and Grain Yields of Wheat in Victoria, Australia1

The effects of various cultural practices on cereal cyst nematode (Heterodera avenae) populations, and the subsequent yield of wheat, have been studied over many seasons in Victoria, Australia. Crop rotations which include periods of fallow, or of non-host crop reduce population levels and improve yields. The inclusion of a legume has the additional advantage of improving soil nitrogen levels. Early sown crops (April/May) are less severely damaged, and produce better yields, than late sown crops (June/July). The resowing of damaged wheat crops with either wheat or barley is not effective in improving yields, although cyst numbers are lower on resown crops. Nitrogenous fertilisers are not generally applied to cereals, and although small increases in yield are sometimes obtained, they are rarely economic. Sulphate of ammonia applications increase cyst numbers, especially when applied at seeding, but urea has only a marginal effect on cyst numbers.     

不同轮作模式下小麦禾谷孢囊线虫的发生动态和种群密度

轮作是防治小麦禾谷孢囊线虫的重要农业措施,为了明确青海省春麦区不同轮作模式对小麦禾谷孢囊线虫的控制效果,采用田间大区试验法对生产中应用的6种轮作模式进行了研究。结果表明:不同轮作模式下,小麦禾谷孢囊线虫种群密度变化差异极显著,其中小麦与马铃薯、油菜、蚕豆轮作两年或以上能有效降低小麦禾谷孢囊线虫种群密度,土壤中的孢囊量减少39.31%~84.39%,单孢虫口数量减少73.21%~95.35%,虫口密度减少83.76%~97.82%;不同作物间,小麦与马铃薯或蚕豆的轮作效果(虫口密度减少74.39%~79.37%)显著优于小麦与油菜的轮作效果(虫口密度减少67.16%)。在同一地块相同条件下,种植油菜、蚕豆、马铃薯、小麦4种作物,小麦禾谷孢囊线虫均能正常孵化,4月底土壤中的2龄幼虫(J2)量增加,5月上旬达到高峰期,5月中旬开始,土壤中的J2、孢囊量、虫口密度和单孢虫口数量均急剧下降,6月至7月份下降幅度小,趋于稳定;田间空孢囊率于5月中旬至6月中旬急剧增加,7月份趋于稳定,8月份以前,4种作物田禾谷孢囊线虫的孵化动态和种群密度变化趋势一致,8月中旬,小麦田随着新孢囊脱落到土壤中,禾谷孢囊线虫种群密度开始上升,小麦收获后土壤中的孢囊量比播种前增加28.62%,虫口密度增加41.30%;而油菜、蚕豆、马铃薯田土壤中的孢囊量比播种前减少32.27%~48.36%,虫口密度减少70.91%~81.73%,8月中旬至10月份小麦田禾谷孢囊线虫种群密度极显著高于油菜、蚕豆、马铃薯田。     

Phosphite Inhibits Development of the Nematodes Heterodera avenae and Meloidogyne marylandi in Cereals

ABSTRACT Phosphonic acid (H(3)PO(3)) solutions were applied to wheat or to bristle oat as soil drenches before inoculation with juveniles of the sedentary, endoparasitic nematodes Heterodera avenae or Meloidogyne marylandi. All the solutions, which were pH adjusted and added at levels as low as 0.63 mg of phosphite (HPO(3)(2-)) per plant, reduced the numbers of H. avenae females and M. marylandi egg masses. Phosphate (PO(4)(3-)), applied as potassium phosphate at the same concentrations, did not reduce the number of female nematodes on the wheat. Addition of phosphate to the phosphite solutions did not change the inhibitory effect of phosphite on H. avenae, but it reduced phosphite's effect on M. marylandi. Phosphite also reduced the number of H. avenae females when applied as many as 20 days after addition of nematodes. The phosphite treatment did not prevent M. marylandi juveniles from penetrating wheat roots or inducing giant cells. However, phosphite inhibited giant cell development: 14 days after inoculation, the giant cells in the phosphite-treated wheat were almost completely vacuolated, whereas those in untreated wheat contained dense cytoplasm.