Surveys of diseases of spring barley in England and Wales, 1976–1980

Samples from 200–300 randomly selected spring barley crops were taken annually at growth stage 73–77 (milky ripe) from 1976 to 1980. The number of samples from each region was proportional to the area of barley grown in each region. The percentage of the area of the top two leaves affected by diseases was recorded. Mildew (Erysiphe graminis) was the most widespread and severe disease recorded. Brown rust (Puccinia hordei) and rhynchosporium (Rhyn-chosporium secalis) occurred frequently but at relatively low levels. Yellow rust {Puccinia striiformis) and septoria (Septoria nodorum) were seen on less than 50% of the samples in most years, and halo spot (Selenophoma donacis) and net blotch (Pyrenophora teres) were rarely recorded. There was an association between the severity of rhynchosporium and the number of rain days in May and June. The highest levels of brown rust occurred in the south and east and rhynchosporium was more common in Wales and the south-west than in the east, but there were no differences in the regional distribution of other diseases. Cultivar resistance, sowing date, previous cropping and fungicide usage were all found to be associated with altered disease levels. The proportion of crops treated with a foliar fungicidal spray rose from 26% in 1976 to 47% in 1980. The use of tridemorph declined but that of triadimefon increased reaching 29% of crops treated by 1980. The use of ethirimol as a seed treatment declined from 16% of crops grown from treated seed in 1976 to 7% in 1980. Estimated yield losses between 1976 and 1980 varied between 4% and 9% due to mildew, between 0.3% and 0.8% due to brown rust and between 0.2% and 0.5% due to rhynchosporium.     

Surveys of diseases of winter wheat in England and Wales, 1976–1988

Samples from 300 – 400 randomly selected winter wheat crops were taken annually at growth stage 73 – 75 from 1976 to 1988 with the exception of 1983 and 1984. The number of samples from each region was proportional to the area of wheat grown in each region. The percentage of the area of the top two leaves affected by diseases, the severity of ear and stem base diseases and, in 6 years, the severity of take-all were recorded. Septoria tritici and Septoria nodorum were, on average, the most severe of the foliar diseases and eyespot (Pseudocercosporella herpotrichoides) was the most severe of the stem base diseases. Regional differences in levels of S. tritici, brown rust, sharp eyespot and nodal fusarium were significant. Cultivar resistance affected disease severity, and previous cropping patterns particularly affected take-all and eyespot. Eyespot and sharp eyespot were less severe in late- than in early-sown crops. The percentage of crops treated with a fungicidal spray increased from 14% in 1976 to over 90% between 1983 and 1985. Use of benzimidazole fungicides applied at growth stage 31 declined, while use of morpholines from flag leaf emergence onwards increased between 1985 and 1988.     

Fungal Communities and Disease Symptoms on Stem Bases of Wheat and Barley and Effects of Seed Treatments Containing Fluquinconazole and Prochloraz

Three successive crops of winter wheat or barley were grown as second, third and fourth cereals. Communities of fungi on shoot bases, identified after isolation on agar media, were more diverse (determined by number of taxa identified) on wheat than on barley, and their diversity increased from year to year. Diversity was not affected by seed treatments containing fluquinconazole or prochloraz. Eyespot (caused by Tapesia spp.) and brown foot rot (caused by Fusarium spp. or Microdochium nivale ) increased from year to year. Eyespot, brown foot rot (after the first year) and sharp eyespot (which remained infrequent), assessed in summer (June), affected wheat more than barley. Eyespot severity was increased slightly on barley by treatments containing fluquinconazole, formulated with or without prochloraz, in the second year (third cereal), when it was also decreased slightly on wheat by fluquinconazole plus prochloraz, except in plots where the treatment had been applied for two successive years. The increases or decreases in eyespot in the second year were accompanied by, respectively, decreases or increases in the frequency of Idriella bolleyi where fluquinconazole was applied alone. Although the eyespot pathogen Tapesia yallundae (but not Tapesia acuformis ) is sensitive to fluquinconazole in vitro , seed treatment, applied principally to control take-all disease, is likely to have only a small effect against eyespot (or other stem-base diseases), and then only on wheat and when formulated with prochloraz.     

Interaction between Rhynchosporium secalis and Pyrenophora teres in the Field and Identification of Genotypes with Double Resistance in a Doubled-haploid Barley Population

Net blotch [Pyrenophora teres (Died.) Drechsl.] and scald [Rhynchosporium secalis (Oudem.) J.J. Davis] are the two most important foliar diseases of barley (Hordeum vulgare L.) in Tunisia. The use of cultivars with double resistance is the most effective method in controlling both diseases. A doubled‐haploid barley population derived from Tunisian cultivars was evaluated to both net blotch and scald during two growing seasons in the field. Mass disease index (MDI), area under the disease progress curve (AUDPC) and apparent infection type (r) were used to assess disease reaction. MDI of net blotch and scald reached up to 65% and 90% respectively. Least significant difference (LSD) test and comparison of the reaction of the doubled haploid (DH) lines to the overall population mean value were efficient in identifying lines with double resistance to both diseases. From the 59 DH lines screened, lines 21, 33, 37, 46 and 47 showed the best level of adult plant resistance to both diseases and may be used in a breeding program for diseases resistance. Interactions between R. secalis and P. teres were investigated at the level of the whole plant under variable epidemic conditions. Under low epidemic conditions, net blotch and scald developments were usually independent, but positively associated for tolerant lines for both diseases. Under high epidemic conditions, competition effects were obtained for susceptible and resistant genotypes. This competition seems to be an exploitation competition that is associated with decreasing resource availability as it occurs only with high levels of infestation or/and when susceptible lines are infected. This study demonstrates the variability of net blotch and scald interaction with epiphytotic conditions and group of resistance.     

A survey of foliar diseases of spring barley in England and Wales in 1967

A total of 5,250 acres of spring barley was sampled in proportion to the barley acreage in each of eight regions. The percentage leaf area ‘affected’ by each disease was recorded for the first (flag) and second leaves at a growth stage when the grain was milky ripe. The operation was automated by punching the data directly on to computer paper tape and a programme was written to analyse the results. Mildew was found to be the most important disease, causing an average loss in yield of approximately 18 %, followed in descending order by brown rust 3%, leaf blotch 1 %, yellow rust 1 %, halo spot 1 %. The total loss due to foliar diseases was estimated to be 20–25% of the national barley yield. Diseases were more severe in the southern than northern regions, e.g. mildew severity was three times higher because of the greater popularity of mildew-susceptible varieties, and the prevalence of weather more conducive to mildew development. Disease severity was not related to previous cropping, and in general the February-sown crops had more mildew than the April-sown crops.     

Eyespot fine structure in Eudorina illinoiensis

Electron microscopy of the eyespot of the coenobic yolvocacean Eudorina illinoiensis (Kofoid) Pascher shows that besides the well-documented anterior-posterior decrease in size, congruity of structure is involved in eyespot differentiation.

In the anterior cells the eyespot lies adjacent to the plasmalemma with the thylakoid underlayers parallel to it, while in the posterior cells eyespot placement and orientation may vary considerably. Deep-sited eyespots of congruent organisation probably represent the mother-cell eyespot; discongruent ones are typical of posterior cells, but the smallest ones may be random associations of osmiophilic globules.

Eyespot structure is discussed in relation to photoreception and phototaxis.     

Eyespot of wheat and barley in Scotland in 1944

In August 1944, eyespot ( Cercosporella herpotrichoides Fron.) was found in ninety out of 121 autumn-sown wheat crops distributed over twelve Scottish counties. It was abundant enough in forty fields to be likely to harm subsequent crops, and was causing obvious loss in eleven crops. The disease was found in seventeen out of eighteen spring-sown barley crops, more than half the straws being infected in seven of them. About 4 % of the wheat inspected was lodged mostly by eyespot and about 38 % of the barley mostly non-parasitically. Eyespot increased with the frequency of wheat and barley in the two preceding years, but a few infected crops occurred on fields where no wheat or barley had been grown for many years. In Scotland, where the atmosphere is more humid, eyespot tends to increase more rapidly than in similar rotations in England; lesions are found higher up the straw and the disease is much more prevalent on spring-sown barley. The long rotations practised in Scotland prevent more extensive damage by eyespot.     

Eyespot membranes in newly released zoospores of the green algaChlorosarcinopsis gelatinosa (Chlorosarcinales) and their fate during zoospore settlement

Summary Eyespot membranes in zoospores of the green algaChlorosarcinopsis gelatinosa were studied with the freeze-fracture technique. The PF of the plasmalemma overlying the eyespot lipid globules contains significantly greater numbers of intramembraneous particles (IMP; 8,200 IMP/m²) compared to other areas of the plasmalemma (2,100 IMP/m²). In the eyespot area the EF of the plasmalemma reveals no IMP, but regularly arranged depressions corresponding to the PF particles. Sizes of PF particles are not significantly different between the eyespot area and other areas of the plasmalemma. Zoospore settlement starts approximately two hours after release and involves in sequence, rounding up of the cells, retraction of the flagella and secretion of a cell wall. Eyespot membrane specializations on the PF of the plasmalemma disappear during flagellar retraction and before cell wall secretion.The functional significance of eyespot membrane specializations is discussed in accordance with the view that these membranes are engaged in photoreception and primary sensory transduction relating to green algal phototaxis.     

Progress in Wheat Resistance to Spot Blotch in Bangladesh

Spot blotch, caused by Cochliobolus sativus, is considered one of the most destructive diseases of wheat (Triticum aestivum) in the warm areas of South Asia. Over the past 20 years, wheat breeding efforts in the region have improved spot blotch resistance in susceptible commercial cultivars. This study assessed resistance and spot blotch‐induced yield losses in newly released wheat cultivars developed in Bangladesh since the release of the landmark wheat variety ‘Kanchan’. Replicated field studies were conducted during the 2003 and 2004 wheat seasons at two sites: a farmer's field and a research station in a warm region of Bangladesh where spot blotch has been a serious problem. Spot blotch affected 60% of the crop and caused yield losses of from 2% to 22%. Disease severity and disease‐induced grain yield reductions were less in wheat genotypes developed since 1983, with a corresponding trend towards higher yield in newly developed varieties. The level of resistance to spot blotch in the new cultivars and advanced breeding lines represents considerable progress in breeding for resistance over the past two decades.     

EFFECTS OF CULTURAL TREATMENTS ON WHEAT AND ON THE INCIDENCE OF EYESPOT, LODGING, TAKE-ALL AND WEEDS. Field Experiments 1945–8

An experiment was made on the fourth, fifth and sixth successive crops of winter wheat to determine the effects of various treatments on the troubles which result from close cereal cropping. Eyespot and lodging were prevalent in the first year (1946); weeds in the second; eyespot, lodging, take-all and weeds in the third.
Spraying with H₂SO₄ reduced the incidence of eyespot, lodging and weeds, and increased yield of grain on plots which received sulphate of ammonia (by 2.7, 2.2 and 10.0 cwt./acre in successive years).
Sulphate of ammonia increased the incidence at harvest of eyespot and lodging, reduced take-all and consistently increased yield of straw. Eyespot and lodging reduced the effect of the fertilizer on yield of grain, take-all increased it.
Increase in seed rate increased the incidence of severe eyespot and of take-all; it increased lodging except when plants were dwarfed by take-all.
Weight of straw and percentage straws with severe eyespot lesions independently affected lodging, together accounting for 51% of the variance in percentage area lodged at harvest and 64 % of that lodged 33 days earlier.
Mean yields of grain on untreated plots sown with 3.3 1/2 bushels seed/acre fell from 26.0 to 22.5 to 11.7 cwt./acre in successive years, whereas yields of 28.4, 29.9 and 29.1 cwt./acre were obtained on sprayed plots sown with 1 1/2.2 bushels seed/acre which received 4 cwt./acre sulphate of ammonia, showing that high yields were maintained when eyespot, lodging, take-all and weeds were controlled.
By 1948 yields of grain on unsprayed plots had fallen to the level of those on similarly manured plots on the continuous wheat experiment on Broadbalk field. Spraying increased grain by amounts similar to those resulting from one year's fallow on Broadbalk; but fallow had its greatest effects on plots with low nitrogen, spraying on those with high nitrogen.     

The identification of candidate genes associated with Pch2 eyespot resistance in wheat using cDNA-AFLP

Eyespot is a fungal disease of the stem base of cereal crops and causes lodging and the premature ripening of grain. Wheat cultivar Cappelle Desprez contains a highly durable eyespot resistance gene, Pch2 on the long arm of chromosome 7A. A cDNA-amplified fragment length polymorphism (AFLP) platform was used to identify genes differentially expressed between the eyespot susceptible variety Chinese Spring (CS) and the CS chromosome substitution line Cappelle Desprez 7A (CS/CD7A) which contains Pch2. Induced and constitutive gene expression was examined to compare differences between non-infected and plants infected with Oculimacula acuformis. Only 34 of approximately 4,700 cDNA-AFLP fragments were differentially expressed between CS and CS/CD7A. Clones were obtained for 29 fragments, of which four had homology to proteins involved with plant defence responses. Fourteen clones mapped to chromosome 7A and three of these mapped in the region of Pch2 making them putative candidates for involvement in eyespot resistance. Of particular importance are two fragments; 4CD7A8 and 19CD7A4, which have homology to an Oryza sativa putative callose synthase protein and a putative cereal cyst nematode NBS-LRR disease resistance protein (RCCN) respectively. Differential expression associated with Pch2 was examined by semi-quantitative RT-PCR. Of those genes tested, only four were differentially expressed at 14 days post inoculation. We therefore suggest that a majority of the differences in the cDNA-AFLP profiles are due to allelic polymorphisms between CS and CD alleles rather than differences in expression. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Strategies for control of eyespot (Pseudocercosporella herpotrichoides) in UK winter wheat and winter barley

This review considers strategies for control of eyespot (Pseudocercosporella herpotrichoides) in winter wheat and winter barley in the UK. Varietal resistance to eyespot may take the form of direct resistance to the growth of the pathogen in the stem base or of indirect tolerance to eyespot through resistance to lodging. The French variety Cappelle-Desprez has been a source of resistance for most UK wheat varieties and a new source of resistance in wild goat grass has been discovered recently. Use of fungicides for control of eyespot increased rapidly after the introduction of the highly effective MBC fungicides in the 1970s, but has decreased recently because UK populations of P. herpotrichoides are now predominantly resistant to MBC fungicides and alternative fungicides are more expensive. Historically, cultural methods, especially crop rotation, have been important in the control of eyespot. The importance of cultural control of eyespot is now increasing because more non-cereal break crops are being grown and fungicide use is declining.     

AB-QTL analysis in spring barley. I. Detection of resistance genes against powdery mildew, leaf rust and scald introgressed from wild barley

The objective of this study was to map new resistance genes against powdery mildew (Blumeria graminis f. sp. hordei L.), leaf rust (Puccinia hordei L.) and scald [Rhynchosporium secalis (Oud.) J. Davis] in the advanced backcross doubled haploid (BC₂DH) population S42 derived from a cross between the spring barley cultivar Scarlett and the wild barley accession ISR42-8 (Hordeum vulgare ssp. spontaneum). Using field data of disease severity recorded in eight environments under natural infestation and genotype data of 98 SSR loci, we detected nine QTL for powdery mildew, six QTL for leaf rust resistance and three QTL for scald resistance. The presence of the exotic QTL alleles reduced disease symptoms by a maximum of 51.5, 37.6 and 16.5% for powdery mildew, leaf rust and scald, respectively. Some of the detected QTL may correspond to previously identified qualitative (i.e. Mla) and to quantitative resistance genes. Others may be newly identified resistance genes. For the majority of resistance QTL (61.0%) the wild barley contributed the favourable allele demonstrating the usefulness of wild barley in the quest for resistant cultivars.     

Different Assessment Techniques of Leaf Blotch (Rhynchosporium secalis (Oud.) J. Davis) in Winter Barley after Artificial Inoculation

Seven winter barley cultivars expressing different field resistance against leaf blotch (Rhynchosporium secalis) and 60 doubled haploid (DH) lines descending from and F₁ hybrid resistant x susceptible were used to develop a test for the quantitative determination of disease severity. Differences in the level of resistance could be measured very precisely after artificial infection of seedlings at the 4-leaf-stage and subseqent incubation at 7 C. The protein of six isolates was usedf, or antiserum production which was subsequently supplied for serological testing (Enzyme-Linked-Immunosorbent-Assay) of infected leaves. The results of visible scoring of the seedlings was significantly correlated with measuring with the ELISA reader. The, results of the seedling test also corresponded to field scores after natural infection.     

Susceptibility to purple blotch (Alternaria porri) in garlic (Allium sativum)

Experiments were conducted to study the progress of purple blotch disease of garlic caused by Alternuria porri in the field, to determine the relationship between garlic leaf age and susceptibility to Alternaria porri, and also to assess loss in bulb characters due to purple blotch of garlic. Per cent disease severity and number of purple blotch lesions on four garlic genotypes of known susceptibility, Sel-10 (highly susceptible), G-41 (highly susceptible), IC-49382 (moderately susceptible) and IC-49373 (moderate to less susceptible) were monitored from bulb formation to bulb maturity at weekly intervals. Lesions appeared early on highly susceptible cultivars, Sel-10 and G-41. Rapid progress of disease development was noticed during the last 3 wk before bulb maturity. Peak severity at the maturity of the crop was significantly higher on highly susceptible genotypes. No definite correlation could be established between number of lesions and disease severity. A logistic curve was fitted to predict the disease progress on different weeks before bulb maturity. Levels of leaf tissue found damaged by A. porri at weekly intervals from bulb initiation to bulb maturity were significantly lower on younger leaves than on older leaves. Leaves that emerged 7 wk before bulb maturity required more than a 5 wk period to reach 50% leaf damage, whereas leaves emerging 2, 3 and 4 wk before bulb maturity exceeded 50% leaf damage within a 2–3 wk period. Individual garlic leaves became more susceptible to purple blotch as they aged and emerging leaves were more susceptible the closer they emerged to bulb maturity. Per cent loss in bulb weight and bulb volume was found to be significantly higher on highly susceptible genotypes. No significant reduction in number of cloves/bulb was observed. We propose 4 wk before bulb maturity as the action threshold for initiation of fungicidal application to prevent damaging levels of disease.     

Response of mulberry accessions to major foliar diseases in temperate conditions of Jammu and Kashmir,India

Thirty-three indigenous and 24 exotic mulberry accessions belonging to five Morus spp. originated from seven countries distributed in temperate and tropical climates were observed for their response to two major foliar diseases during 2010 and 2011 under temperate conditions of Jammu and Kashmir, India. Leaf spot and powdery mildew severity (Percent Disease Index (PDI)) ranged from 0.00 to 74.90% and 0.00 to 59.85%, respectively. Indigenous and exotic accessions responded similarly to leaf spot, but varied too much to powdery mildew. Irrespective of origin, response in ascending order of PDI for leaf spot is M. multicaulis, M. indica, M. alba, M. kayayama and M. bombycis and for powdery mildew is M. multicaulis, M. kayayama, M. bombycis M. alba and M. indica. Among indigenous accessions, Brentul Kashmir offered highest resistance to both the diseases. Nadigam offered maximum resistance only to leaf spot followed by Himachal local; while Chinarpati followed by Mysore local only for powdery mildew. Among exotic accessions, Ichinose offered maximum resistance to both the diseases followed by Kokusou-21 and Tagowase.     

Effectiveness of plant extracts to control purple blotch and Stemphylium blight diseases of onion (Allium cepa L.) in Assiut,Egypt

The present investigation aimed to isolate the causative agents of onion purple blotch and Stemphylium blight diseases and evaluate the efficacy of certain plant extracts against the two identified pathogens, in vitro and under greenhouse condition. Fourteen isolates of S. vesicarium and two isolates of Alternaria porri were tested for pathogenicity. The results indicated that all isolates were able to produce the symptoms of Stemphylium blight and onion purple blotch diseases with different degrees of severity ranging from 10.42 to 81.25%. A. porri No. 6022 caused the highest disease severity (81.25%), while S. vesicarium No. 6003 was the best one out of the tested 14 isolates (37.5%). Antifungal activity of some aqueous plant extracts (Azadirachta indica, Cydonia oblonga, Datura stramonium, Eucalyptus globulus, Foeniculum vulgare, Ocimum basilicum, Rosmarinus officinalis and Salix mucronata) was assayed in vitro by dry weight technique. The data indicated that there were significant differences between these extracts in their effect on fungal growth of A. porri and S. vesicarium and the best were A. indica and D. stramonium. Under greenhouse conditions, application of the aqueous extract of A. indica either before or after 48?h A. porri inoculation produced the highest reduction in disease severity comprising 70 and 74.7%, respectively. On the other hand, the highest percentage of disease reduction before and after 48?h S. vesicarium inoculation was produced by Ridomil gold plus reached to 84.4 and 95.8% respectively, followed by the aqueous extract of A. indica (74.1 and 89.7, respectively). According to our results, it can be concluded that plant extracts of A. indica and D. stramonium can be used for the biocontrol of purple blotch and Stemphylium blight diseases instead of fungicides to minimise the risks and hazards of using toxic fungicides.     

Prediction of Eyespot Severity on Winter Wheat or Winter Barley Inoculated with W-type or R-type Isolates of Pseudocercosporella herpotrichoides

In crops of winter wheat (1986—88) or winter barley (1987—88) inoculated with W-type or R-type isolates of Pseudocercosporella herpotrichoides and sown on different dates (1986) or at different seed rates (1987,1988) eyespot epidemics developed in different ways. Methods of measuring eyespot incidence/severity during crop growth were compared for their ability to predict eyespot severity at grain filling. Regressions were calculated for eyespot severity score at GS 71 on earlier measurements, either at GS 30/31 (11 methods) or from GS 22 to GS 65 (3 methods). Based on measurements at GS 30/31, all the methods predicted eyespot severity at GS 71 well in plots of winter barley inoculated with W-type isolates (r, 0.83—0.97) but the accuracy of predictions in plots inoculated with R-type isolates was very variable (r, 0.09—0.71). Predictions for 1987 and 1988 were less accurate in wheat than in W-type plots of barley, but did not differ between W-type and R-type plots (r, 0.70—0.89). When the wheat data for 1986 were also included predictions were less accurate, especially in R-type plots (r, 0—0.59). Generally, it was easier to predict eyespot severity at GS 71 in W-type than in R-type plots, especially in barley and in wheat before GS 37/39. Predictions of eyespot severity at GS 71 based on measurements before GS 25 were inaccurate for both wheat and barley. After GS 25 the accuracy of the prediction was generally good in W-type plots and did not improve greatly except in wheat after GS 59. However, there was a steady improvement in the accuracy of the prediction in R-type plots of barley from GS 24 to GS 53. Assessments of eyespot incidence on stems predicted eyespot severity at GS 71 more accurately than assessments on leaf sheaths on wheat after GS 37/39, but were not as good on barley until GS 53.     

Effect of Leaf Infection by Kabatiella zeae on Stalk Rot Prevalence and Grain Yield of Maize Hybrids

Eight maize hybrids were tested for resistance to eyespot (Kabatiella zeae) and stalk rot (Fusarium spp.) under field conditions in 1993–1995. The relationship between eyespot and stalk rot and their effect on grain yield of hybrids were studied. Effectiveness of the prochloraz fungicide in controlling both diseases was evaluated. The experiments were performed in three subtrials where plants were naturally infected, inoculated with K. zeae or treated with the fungicide, Sportak 450 EC. The relationship between intensity of leaf damage by eyespot and incidence of stalk rot has been evidently confirmed. High infection of leaf by K. zeae reduced grain yield of hybrids. However, considerable differences in prevalence of disease and yield losses were noted in successive years. The intensity of eyespot depended upon environmental conditions highly variable over the years: it was enhanced by low temperature and high air humidity in July and August. Hybrids showed significant differences in susceptibility to both diseases. Mona – hybrid from the USA and two experimental hybrids from Poland were less affected by diseases than Smolimag and Hidosil (commercial hybrids from Poland). The fungicide Sportak was effective in controlling eyespot and had an indirect influence on decreasing fusarium stalk rot.     

Effects of crop debris and cultivations on the development of eyespot of wheat caused by Oculimacula spp.

Results from a series of crop sequence and single-year experiments that tested different straw and cultivation treatments in a total of 11 site–season combinations confirmed previous evidence that the development of eyespot disease in cereals can be decreased by debris in the seed bed even if that debris includes eyespot-infected stem bases, which are the principal sources of primary inoculum. Two of the experiments, which followed non-cereal break crops and tested the effects of crop debris applied after ploughing or tining on eyespot in winter wheat that was artificially inoculated with Oculimacula spp., provided convincing evidence that the effects can be attributed to the debris per se, and not to any associated husbandry practices. There were often larger effects on disease in summer than in spring or on severity than on incidence suggesting that the effects of debris cannot be explained solely by effects on inoculum or initial infection, and that debris has a disease-suppressive effect.