Observations of apothecia of Tapesia yallundae and the cultural phenotypes of their progeny

Apothecia of Tapesia yallundae, the perfect state of Pseudocercosporella herpotrichoides, were found in March 1989 on wheat and barley stubble at nine of 37 naturally infested sites and also at one inoculated site in Germany. Apothecia were found on about 1% of stems collected from the naturally infested sites and 5% of stems from the inoculated site, and were first found in the second week of March, coinciding with the onset of an early spring. Apothecia were not found on either green plants or second-year stubble. In total, 143 single-spore isolates were obtained from the apothecia and described according to a range of morphological characteristics. The majority were ascribed to P. herpotrichoides var. herpotrichoides. This is the first isolation of this variety from T. yallundae apothecia in Germany. Even single-conidial isolates from one apothecium showed much variation, some being like var. herpotrichoides, some like var. acuformis and some intermediate.     

Species and Mating-Type Distribution of Tapesia yallundae and T. acuformis and Occurrence of Apothecia in the U.S. Pacific Northwest

ABSTRACT Eyespot of wheat is caused by the discomycete fungi Tapesia yallundae and T. acuformis. T. yallundae is considered the most important causal agent of the disease in this region but no apothecia of either species have been found in the U.S. Pacific Northwest (PNW). Two compatible isolates of T. yallundae from the PNW were used to inoculate a field plot in the fall of 1998 and apothecia developed in the spring and fall of 2000 on standing wheat stubble. In the spring of 2000, wheat stubble from eight naturally infected fields was examined for the presence of apothecia of T. yallundae and T. acuformis. Apothecia of T. acuformis were found in two fields but no apothecia of T. yallundae were found. This is the first report of apothecia of the eyespot pathogens occurring in the PNW. Species and mating-type distribution of T. yallundae and T. acuformis in the PNW were determined from 817 isolates collected from diseased wheat over 3 years at spatial scales ranging from within fields to across states. In all, 460 isolates were identified as T. yallundae and 357 isolates were identified as T. acuformis with MAT1-1/MAT1-2 ratios not significantly different from 1:1 based on chi(2) tests at most scales tested. The apparent increase in frequency of T. acuformis from previous surveys may indicate a shift in the predominant species causing eyespot. The occurrence of apothecia under field conditions, along with the widespread distribution of mating types of both species, suggests that sexual reproduction may be occurring in both species.     

Occurrence of Tapesia yallundae apothecia on field and laboratory-inoculated material and evidence for recombination between isolates

Tapesia yallundae was observed in early July on wheat straws, which had been inoculated with isolates of Pseudocercosporella herpotrichoides (W-type) 9 months previously and placed in plots of winter wheat, to act as inoculum in a resistance screen. A perfect state was not observed on straws inoculated with R-type isolates in the same experiment. One hundred and thirty-four single-ascospore isolates were obtained and observations on morphology in culture, pathogenicity to wheat and rye seedlings, resistance to benomyl and the pattern of restriction fragment length polymorphisms demonstrated that all isolates were W-type, although some grew at half the expected rate on agar, and at least two isolates were involved in the production of apothecia. Apothecia were produced in vitro on straws inoculated with single isolates of P. herpotrichoides only when they were incubated in contact with other straws similarly inoculated with certain other isolates. No apothecia were produced on straws with only isolate present, suggesting the need for at least two strains, possibly representing different mating types, for apothecium formation. Apothecia of T. yallundae were produced in vitro on straw pieces which had been inoculated individually with W-type isolates, 14-16 weeks after straws of different isolates were mixed. Evidence for genetic recombination between isolates was obtained when single-ascopore colonies were analysed for resistance to benomyl in culture and for restriction-length polymorphisms (RFLPs).     

First record of Tapesia yallundae as the teleomorph of Pseudocercosporella herpotrichoides var. acuformis, and its occurrence in the field in the Federal Republic of Germany

Tapesia yallundae is reported for the first time in the Federal Republic of Germany. Apothecia were found on wheat and rye stubble at two sites in the north of the country and one in the centre, in April 1988. Isolates of Pseudocercosporella herpotrichoides var. acuformis were obtained from germinating single ascospores, identified on the basis of colony morphology, mode of conidial production and pathogenicity to wheat and rye.     

Occurrence of Tapesia yallundae, teleomorph of Pseudocercosporella herpotrichoides, on unharvested wheat culms in England

Apothecia of Tapesia yallundae are reported for the first time in the UK. They were observed on decaying stems and leaf sheaths of unharvested wheat near Bristol in January 1989. Isolates from single ascospores produced cultures typical of the W-type form of the anamorph, Pseudocercosporella herpotrichoides . Pathogenicity on wheat was confirmed by inoculation and re-isolation from glasshouse-grown wheat seedlings.     

Incidence of apothecia of Tapesia yallundae at setaside sites in England and sensitivity of the ascospore offspring to the fungicides benomyl and prochloraz

A total of 45 field sites in England were surveyed once for the presence of apothecia of Tapesia yallundae from 1992 to 1994. Apothecia were found at 21 locations and were mainly present on less than 3% of stems. However, apothecia were found on 1532% of stems at four sites. Analysis of the growth characteristics of ascospore isolates from seven sites showed that most produced colonies characteristic of the W-type of T. yallundae , with only one site yielding the R-type, Most ascospore isolates were resistant to the fungicide benomyl and effectively all remained sensitive to prochloraz. The results of the study are discussed in relation to the fungicidal control and epidemiology of T. yallundae , and the risk of spread of disease from set-aside sites.     

A species of Unguicularia on oilseed rape, and its importance in studies of the epidemiology of light leaf spot (Pyrenopeziza brassicae)

Apothecia of a species of Unguicularia were found on fallen leaves of oilseed rape from November 1989 to April 1990, and from January to July 1991. On the basis of the limited literature on this genus, these apothecia were tentatively identified as Unguicularia cfr. raripila, a species not previously reported in the UK, or on oilseed rape. Although saprophytic, it is of significance in oilseed rape crops, as its ascospores are similar in size and shape to those of the important pathogen Pyrenopeziza brassicae, the cause of light leaf spot. The ascospores of both species can be dispersed either as single spores or in groups, although those of U. cfr. raripila are more often dispersed in groups. Due to confusion in distinguishing the ascospores of these two species, it is likely that studies of the epidemiology of light leaf spot using spore samplers have overestimated the numbers of ascospores of P. brassicae dispersed, and hence their potential contribution to epidemics. Apothecia of U. cfr. raripila were abundant in the spring of both 1990 and 1991, but those of P. brassicae were much more rare. The species of Unguicularia is described.     

Isolation of R-type progeny of Tapesia yallundae from apothecia on wheat stubble in England

Apothecia of Tapesia yallundae were collected from a set-aside straw stubble site in Lincolnshire in March 1993. Single ascospore isolates were obtained which produced colonies with morphologies and growth rates characteristic of the R-pathotype of Pseudocercosporella herpotrichoides. Isolates were confirmed to be R-type by the use of randomly amplified polymorphic DNA (RAPD) markers. These observations confirm that T. yallundae is the teleomorph of the R-type of P. herpotrichoides and represent the first detection of the sexual stage of the R-type in the UK.     

Changes in populations of the eyespot fungi Tapesia yallundae and T. acuformis under different fungicide regimes in successive crops of winter wheat, 1984–2000

Effects of regular treatments with the fungicides carbendazim and prochloraz applied to whole plots divided into subplots with different initial population mixtures of carbendazim-sensitive or carbendazim-resistant Tapesia yallundae or T. acuformis were studied in successive crops of winter wheat from 1984/85 to 1999/2000. In unsprayed and carbendazim-sprayed whole plots, a stable coexistence of about 50% each of T. yallundae and T. acuformis developed within five seasons, but in whole plots sprayed with prochloraz or prochloraz plus carbendazim, the proportion of T. acuformis increased to > 80%. A discrete time difference equation model was derived from knowledge of the biology of eyespot and competition theory to describe the population changes. The model was fitted to the data from treatments where coexistence occurred [subplots in unsprayed (1985–92) and carbendazim-sprayed (1985–89) whole plots], using nonlinear least squares regression. The optimized value of the resource overlap coefficient was small, suggesting niche differences between the two species. Populations were nearly 100% carbendazim-resistant in carbendazim-sprayed whole plots by July 1985 (one season) and in whole plots sprayed with prochloraz plus carbendazim by July 1986 (two seasons). In prochloraz-sprayed whole plots, the proportion of carbendazim-resistant isolates decreased more rapidly than in unsprayed whole plots in the 1980s, but by July 1992 a shift in populations in unsprayed and prochloraz-sprayed whole plots towards predominantly carbendazim-resistant strains had occurred.     

Development of apothecia ofTapesia yallundae in contrasting populations selected by fungicides

Apothecia of the eyespot fungus,Tapesia yallundae, were found on 0–18% of straws in plots of wheat stubble in February–March 1994. The fungicides carbendazim, prochloraz or carbendazim plus prochloraz had been applied repeatedly to the same plots in each of the previous 9 years in which successive wheat crops had been grown. The factors most strongly correlated with the incidence of apothecia were the incidence and severity of eyespot in the preceding wheat crop and the frequency of carbendazim-resistant W-type fungus in populations recovered from that wheat crop. Plots treated with carbendazim, which had previously had more disease and more resistance to carbendazim in the pathogen population relative to untreated plots, therefore yielded most apothecia. Plots treated with prochloraz, which had selected for predominantly R-type fungus and decreased eyespot, yielded few apothecia. Single-ascospore isolates were all of the W-type and were more frequently carbendazim-sensitive than expected, except those from plots treated only with carbendazim. None showed decreased sensitivity to prochloraz. The implications of applying fungicides regularly for controlling eyespot on the capability of the eyespot fungus for genetic variation through sexual reproduction are discussed.     

The effect of winter weather conditions on the ability of pseudothecia of Leptosphaeria maculans and L. biglobosa to release ascospores

Leptosphaeria maculans and L. biglobosa are damaging pathogens of oilseed rape. The infection of plants occurs predominantly in early autumn or spring by spores produced in pseudothecia. The aim of this study was to investigate whether pseudothecia formed in the autumn are still viable in the spring and to what extend they are destroyed by winter frosts. The studies presented here demonstrated that winter frosts can render pseudothecia unable to release spores. Nevertheless, ascospores present in pseudothecia unable to discharge ascospores, were fully capable of germination, regardless of the incubation temperature. No significant differences were found between the studied Leptosphaeria species in their response to frost. A multiple regression equation has been elaborated to forecast the ability of pseudothecia to release ascospores, based on winter temperatures. Considerable correlation was found between the ascospore release in the autumn and the ability of pseudothecia to release ascospores over the winter period and the subsequent symptoms of stem canker before harvest. We have demonstrated that the potential and the survival of inoculum can have a large impact on the success of the pathogen. This may be particularly important in the light of forecasted climate change. Higher winter temperatures may increase the ability of pseudothecia to release ascospores and the discharge of ascospores of L. maculans and L. biglobosa into the air, and cause early plant infections. This in turn will increase the number of infected plants, the disease incidence at harvest, and reduce the yield of oilseed rape.     

The production and release of ascospores of Pyrenopeziza brassicae on oilseed rape

The results of observation for three seasons in oilseed rape crops of the production and release of ascopores of Pyrenopeziza brassicae , the cause of light leaf spot disease of brassicas, are presented. Large apothecia (1–2 mm in diameter) take at least 3 weeks to develop on leaf petioles after infected leaves die, but small apothecia (50–200 um) may form on leaf lamellae after about 15 days. Apothecia were found on all 12 oilseed rape varieties examined. Spore trapping experiments in infected crops show that ascospore release is associated with rain, but most ascospores are released after rainfall when the crop debris bearing apothecia are wet. Ascospores can be released for up to 5 days after rain. Laboratory measurements show that apothecia can continue to release ascospores for up to 3 weeks even when they are subjected to wet and dry cycles. The consequences of the widespread occurrence of the sexual stage of P. brassicae are discussed and possible cycles of disease and spore production in oilseed crops are suggested.     

Growth of seedlings of Agropyron repens (L.) Beauv. and Agrostis gigantea Roth in cereal crops

Experiments were undertaken to define the conditions under which seeds and seedlings of Agropyron repens and Agrostis gigantea may start infestations in cereal crops. When seedlings were planted early or late in spring wheat and spring barley, most growth of shoots and rhizomes was produced by Agropyron planted early in wheat. Late planting halved the amount of shoot growth and severely inhibited rhizome formation. In winter wheat given a moderate or zero amount of nitrogen fertilizer in spring, growth of the weed seedlings was slow. Rhizomes were not produced during the time the crop was growing but only after harvest. Agrostis made more growth than Agropyron in most treatments throughout most of the experiment. Late planting decreased growth more than in the spring cereals. Nitrogen fertilizer, although it had little effect on the amount of growth made by winter wheat, increased the growth of the early-planted seedlings but decreased that of the late-planted ones of both weed species. When planted into plots given nitrogen, more seedlings of both species died after late than after early planting. Clearly, the amount of growth and rhizome produced by seedlings of these two species will depend on the type of cereal, the time of emergence of the seedlings in relation to the cereal, and on other factors affecting the relative vigour of cereal and weed. Evidently, where the weed seedlings emerge early in weakly or moderately competitive cereal crops or when growth is unchecked in the cereal stubble, seedlings could give rise to infestations.     

DIRECT-DRILLING OF CEREALS AFTER USE OF PARAQUAT

Summary. Cereal crops were grown at Jealott's Hill in 1962 and 1963 without cultivations, following the use, before drilling, of paraquat to kill unwanted vegetation. This investigation was extended with a series of trials at outside centres. Three trials with winter wheat were drilled in the autumn of 1963, two into pasture and one into stubble. Three barley trials were drilled into stubble in spring 1964.
All the autumn trials and one spring trial compared direct-drilling with normal cultivations at two levels of nitrogen. The other two spring trials compared rates and times of application of paraquat without any plough comparison.
Individual trials showed significant yield differences between direct-drilling and ploughing, but there was no consistent advantage in favour of either technique.
Spraying in advance of drilling generally resulted in similar yields to spraying and drilling on the same day. Winter wheat drilled into pastures showed some retardation of spring growth when spraying and drilling had been carried out on the same day.
When averaged over times of application, yields after spraying paraquat at 2 lb/ac were significantly higher than after 1 lb/ac.
In the trials where two levels of nitrogen were included crops responded to an increase from 60 to 120 units nitrogen/ac as top dressing for winter wheat and from 60 to 90 units nitrogen/ac combine-drilled for spring barley; the response was not always significant.
Semis direct de céréales après application de paraquat     

西北黄土区主要作物茬口对土壤理化性质和冬油菜农艺性状的影响

为了探讨不同茬口对土壤肥力及后茬油菜产量的影响,以休闲田为对照,研究了西北黄土区冬油菜、马铃薯、玉米、冬小麦、春小麦、蚕豆等6种主要农作物茬口土壤耕作层肥力效应及其对后茬白菜型冬油菜产量和农艺性状的影响,结果表明：（1）不同作物茬口土壤养分含量总体表现为越冬作物>休闲田>春播作物,除固氮作物蚕豆茬口碱解氮含量较高外,冬油菜茬口土壤营养各项指标、土壤微生物种群结构及土壤物理性状均显著优于其它作物茬口,其有机质、全氮、全磷、全钾、碱解氮、速效磷、速效钾含量分别比休闲田增加32.6%、12.1%,5.9%、7.2%、11.6%、99.8%、44.2%,其次为蚕豆、冬小麦茬口和休闲田,玉米和春小麦茬口肥力状况较差;与休闲田相比,冬油菜、冬小麦和蚕豆茬口耕层（0~20 cm）土壤容重分别降低10.14%、5.80%和5.80%,而冬油菜茬口土壤总孔隙度、毛管孔隙度和田间持水量分别提高7.33%、4.12%和5.65%,马铃薯、玉米、春小麦茬口均有不同程度下降,冬小麦与蚕豆茬口无明显差异;（2）前茬对白菜型冬油菜农艺性状的影响主要表现在株高、根量、全株角果数、角果粒数及千粒重等指标,其中蚕豆茬口白菜型冬油菜比休闲田增产4.50%,冬小麦茬口与休闲田相当,而马铃薯、玉米、春小麦、冬油菜茬口分别较休闲田减产11.05%、15.04%、16.27%、21.14%。白菜型冬油菜的最佳前茬是豆类作物,休闲田和冬小麦茬口次之,但白菜型冬油菜连作可能会产生自毒作用或病害加重,从而造成显著减产。     

Response of spring cereal crops to soil residues of ethofumesate*

For evaluation of soil residues of ethofumseate [(±)2-ethoxy-2,3-dihydro-3,3-dimethylbenzofuran-5-yl-methylsulphonate], spring barley (Hordeum vulgare L.) and spring wheat (Triticum aestivum L.) were sown in soil planted 11 months previously with sugar-beet (Beta vulgaris L.) and previously treated with band or broad-cast applications of the herbicide at rates of 1.68-4.48 kg/ha. The height of barley and wheat was suppressed only where ethofumesate was applied broadcast and the soil was not ploughed after sugar-beet harvest. Three months after sowing, the 4.48 kg/ha rate suppressed the height of barley 18% and wheat 34%. In another experiment, barley grew normally on soil treated the previous year with pre-planting and post-emergence applications of ethofumesate where ploughing followed sugar-beet harvest. For all treatments, yields of straw and grain were not significantly lower than those of the untreated check plots. The dissipation of ethofumesate 24 weeks after application to sugar-beet was 89–100% when applied on a band compared to 85-95% when applied broadcast.     

Development of Oculimacula yallundae and O. acuformis (eyespot) lesions on stems of winter wheat in relation to thermal time in the UK

Relationships between development of eyespot, caused by Oculimacula yallundae (OY) or O. acuformis (OA) on stems of winter wheat (cv. Avalon), and thermal time (°C days after sowing) were investigated in field experiments in 1985/86, 1986/87 and 1987/88 (two experiments). In all experiments, the incidence and severity of stem eyespot (uninoculated plots, OY- and OA-inoculated plots) were linearly related to accumulated thermal time after sowing. There were ca. 600–800°C days from the time of the first sample when eyespot lesions were recorded on stems to the time when maximum eyespot incidence or severity was recorded. Relationships between stem eyespot incidence or severity and thermal time differed between seasons, with more severe eyespot in 1986/87 and 1987/88 than 1985/86. Both the severity and volume of stem lesions were initially greater in OY-inoculated plots than OA-inoculated plots in spring but differences were less by harvest in all seasons. The percentage of plants with stems colonized by OY or OA over all plots (including uninoculated) showed a consistent pattern in 1986/87 and 1987/88 (two experiments), with the percentage colonized by OY greater initially and the percentage colonized by OA gradually increasing with time towards harvest.     

麦园蜘蛛(Penthaleus sp.)及麦长腿蜘蛛(Petrobia latens(Müller))的生物学研究

1953—1955年在晋南、北京等地对麦园蜘蛛及麦长腿蜘蛛的发生规律及生活习性进行了研究。结果明确两种麦蜘蛛一年中:秋季密度最大时期为11月上、中旬。春季密度最大的时期,麦园蜘蛛在3月下旬至4月上旬;麦长腿蜘蛛在5月上旬前后。两种麦蜘蛛均以滞育卵越夏。直至10月中旬,越夏卵始孵化,孵化最盛在11月上旬,此时幼、若虫出现最多,若进行防治可基本上防止以后的猖獗。根据结果分析,估计麦园蜘蛛每年发生2—3代,麦长腿蜘蛛发生3—4代。室内饲育初步观察麦园蜘蛛一世代(从卵到产卵)历时平均约57.8天;麦长腿蜘蛛为32.1天。两种麦蜘蛛都是单性生殖。麦长腿蜘蛛的滞育卵,具多年滞育习性,初步观察有些卵能在土中生活两年。两种麦蜘蛛的分布和猖獗都受湿度的影响,主要猖獗为害地区麦长腿蜘蛛在黄河以北的旱地;麦园蜘蛛在黄淮地区的水浇地或低洼地。耕作栽培对麦蜘蛛的发生有重要影响;连作对两种麦蜘蛛发生有利;深耕能将卵翻至深土之下,可以减少发生;晚播影响麦蜘蛛孵化后的食料。因此,合理轮作、深耕和适当晚播是综合防治中的重要措施。     

Population Genetic Structure of Tapesia acuformis in Washington State

ABSTRACT Eyespot of wheat is caused by Tapesia yallundae and T. acuformis. Historically, T. yallundae has been considered the more important causal agent of the disease in Washington state and consists of a large homogeneous population with a genetic structure consistent with both sexual and asexual reproduction. T. acuformis has increased significantly in Washington in the past 10 years and apothecia were found recently under natural field conditions, indicating that T. acuformis may have a more important role in eyespot of wheat than previously was thought. To determine the genetic structure of T. acuformis in Washington, 141 single conidial isolates were sampled from four subpopulations in the eastern wheat-growing region of the state. Isolates were scored for mating type and six amplified fragment length polymorphism markers. All markers segregated in a 1:1 ratio and were determined to be unlinked based on genetic analysis of 24 progeny from an in vitro cross. No significant differences in allele frequencies (0.127 < P < 0.809) were found among individual loci across the four subpopulations and over all loci based on contingency table analysis of the log-likelihood ratio statistic G(2). Likewise, no overall differences between subpopulations were detected using the population differentiation statistic theta (theta = -0.004, P = 0.537). Random mating could not be rejected within each subpopulation or for the combined data using clone-corrected data sets based on (i) 1:1 ratio of mating-type, (ii) multilocus gametic disequilibrium analyses (index of association), (iii) phylogenetic analyses (parsimony tree length permutation test), and (iv) genotypic diversity analyses. T. acuformis has a genetic structure similar to that of sympatric populations of T. yallundae in Washington, with both sexual and asexual reproduction contributing to the structuring of this species.     

A comparative assessment of potential components of partial disease resistance to <Emphasis Type="Italic">Fusarium</Emphasis> head blight using a detached leaf assay of wheat,barley and oats

The relative resistance of 15 winter barley, three winter wheat and three winter oat cultivars on the UK recommended list 2003 and two spring wheat cultivars on the Irish 2003 recommended list were evaluated using Microdochium nivale in detached leaf assays to further understand components of partial disease resistance (PDR) and Fusarium head blight (FHB) resistance across cereal species. Barley cultivars showed incubation periods comparable to, and latent periods longer than the most FHB resistant Irish and UK wheat cultivars evaluated. In addition, lesions on barley differed from those on wheat as they were not visibly chlorotic when placed over a light box until sporulation occurred, in contrast to wheat cultivars where chlorosis of the infected area occurred when lesions first developed. The pattern of delayed chlorosis of the infected leaf tissue and longer latent periods indicate that resistances are expressed in barley after the incubation period is observed, and that these temporarily arrest the development of mycelium and sporulation. Incubation periods were longer for oats compared to barley or wheat cultivars. However, oat cultivars differed from both wheat and barley in that mycelial growth was observed before obvious tissue damage was detected under macroscopic examination, indicating tolerance of infection rather than inhibition of pathogen development, and morphology of sporodochia differed, appearing less well developed and being much less abundant. Longer latent periods have previously been related to greater FHB resistance in wheat. The present results suggest the longer latent periods of barley and oat cultivars, than wheat, are likely to play a role in overall FHB resistance if under the same genetic control as PDR components expressed in the head. However the limited range of incubation and latent periods observed within barley and oat cultivars evaluated was in contrast with wheat where incubation and latent periods were shorter and more variable among genotypes. The significance of the various combinations of PDR components detected in the detached leaf assay as components of FHB resistance in each crop requires further investigation, particularly with regard to the apparent tolerance of infection in oats and necrosis in barley, after the incubation period is observed, associated with retardation of mycelial growth and sporulation.