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Fusarium Head Blight (fusarium + head_blight)
Selected AbstractsOccurrence and Distribution of Microdochium and Fusarium Species Isolated from Durum Wheat in Northern Tunisia and Detection of Mycotoxins in Naturally Infested GrainJOURNAL OF PHYTOPATHOLOGY, Issue 9 2009Lobna Gargouri Kammoun Abstract An outbreak of Fusarium Head Blight of durum wheat occurred in 2004 being localized in sub-humid and higher semi-arid region of Northern Tunisia. A mycological survey carried out throughout these regions, revealed that 78% of the prospected fields were infested. Results of the morphological and molecular identification, showed that the most common species isolated from diseased wheat spikes was Microdochium nivale var. nivale (63.5%), followed by Fusarium culmorum (26%), F. pseudograminearum (9%) and F. avenaceum (1.5%). To evaluate mycotoxin content of naturally infected grain, the amounts of trichothecene mycotoxin deoxynivalenol (DON) in harvested grain from 45 fields were quantified by RIDASCREEN DON Enzyme Immunoassay Kit (ELISA). This study showed that the infection levels in freshly harvested grain were very low and the maximum deoxynivalenol (DON) level of the positive samples was 53 ppb. This is the first report on the natural occurrence of DON in naturally infected wheat grain sampled from Northern Tunisia. [source] Ultrastructural and Immunocytochemical Studies on Effects of Barley Yellow Dwarf Virus , Infection on Fusarium Head Blight, Caused by Fusarium graminearum, in Wheat PlantsJOURNAL OF PHYTOPATHOLOGY, Issue 1 2006Y. Liu Abstract The interactions between barley yellow dwarf virus (BYDV) and Fusarium head blight (FHB), caused by Fusarium graminearum, were studied in the two winter wheat cultivars (cvs.), Agent (susceptible to FHB) and Petrus (moderately resistant to FHB), using ultrastructural and immunocytochemical methods. Infections of wheat plants of both cvs. by BYDV increased susceptibility to FHB. BYDV infection caused numerous cytological changes in lemma tissue of both cvs. such as formation of vesicles in the cytoplasm, degradation of fine structures of chloroplasts of both cvs. and accumulation of large starch grains in the chloroplasts. Electron microscopical studies showed that the development of F. graminearum on spike surfaces was not affected in BYDV-infected plants. After penetration and intercellular growth in lemma tissue, defence responses to Fusarium infections were markedly reduced in BYDV-diseased plants compared to the tissue of virus-free plants. At sites of contact of fungal cells with host tissue, depositions of cell wall material were distinctly less pronounced than in tissues of virus-free plants of cv. Petrus. Detection of , -1,3-glucanases and chitinases in lemma tissue of cv. Agent revealed no appreciably increased accumulation of both defence enzymes in F. graminearum -infected virus-free and BYDV-infected tissues compared to the non-infected control tissue. On the other hand, in cv. Petrus, infection with F. graminearum induced a markedly enhanced activity of both enzymes 3 days after inoculation. The increase of both enzyme activities was less pronounced in BYDV-infected plants than in tissue exclusively infected with F. graminearum. Cytological studies suggest that in contrast to the susceptible cv. Agent postinfectional defence responses may play still an important role in the resistance of the moderately resistant cv. Petrus to FHB. [source] Studies on Symptom Development, Phenolic Compounds and Morphological Defence Responses in Wheat Cultivars Differing in Resistance to Fusarium Head BlightJOURNAL OF PHYTOPATHOLOGY, Issue 4-5 2002E. SIRANIDOU Abstracts Development of Fusarium head blight, caused by Fusarium culmorum, was evaluated in seven winter wheat cultivars and one spring wheat cultivar in an outdoor pot experiment. Of the cultivars studied, only the spring wheat cultivar Frontana exhibited resistance against initial infection by the fungus. Significantly higher amounts of free phenolic compounds were found in the glumes, lemmas and paleas of Frontana prior to and at all sampling times after inoculation, in comparison to the winter wheat cultivar Agent. Furthermore, the amount of p -coumaric acid increased significantly in the glumes, lemmas and paleas of the cultivar Frontana 2 days after inoculation compared to that in uninoculated spike tissues. However, the amounts of ferulic acid between inoculated and healthy plant tissues did not differ. Ultrastructural studies indicated more pronounced structural defence responses, such as cell wall appositions, in the infected lemma tissue of the resistant cultivar Frontana compared with the susceptible Agent 3 days after inoculation. Immunogold labelling of lignin revealed no differences in the density of gold particles on the cell walls of healthy lemma tissue in the two cultivars. However, density of particles increased in the infected lemma tissue of the resistant cultivar Frontana on cell walls adjacent to the fungal cells, 3 days after inoculation. The susceptible cultivar Agent showed little or no response to the infection. Phenolic compounds appear to play a role in the resistance of the cultivar Frontana to F. culmorum. [source] Osmotic shock tolerance and membrane fluidity of cold-adapted Cryptococcus flavescens OH 182.9, previously reported as C. nodaensis, a biocontrol agent of Fusarium head blightFEMS YEAST RESEARCH, Issue 3 2007Christopher A. Dunlap Abstract Cryptococcus flavescens (previously reported as C. nodaensis), a biological control agent of Fusarium head blight, has been previously shown to have improved desiccation tolerance after cold adaptation. The goal of the current study was to determine the effect of cold adaptation on the physicochemical properties of C. flavescens that may be responsible for its improved desiccation tolerance. The results show that cold adaptation improves liquid hyperosmotic shock tolerance and alters the temperature dependence of osmotic shock tolerance. Fluorescence anisotropy was used to characterize differences in the membrane fluidity of C. flavescens with and without cold adaptation. Force curves from atomic force microscopy showed a significant increase in the cell wall spring constant after cold adaptation. Cold adaptation of C. flavescens during culturing was shown to produce smaller cells and produced a trend towards higher CFU yields. These results suggest that cold adaptation significantly alters the membrane properties of C. flavescens and may be an effective method of improving the desiccation tolerance of microorganisms. In addition, we provide information on the correct naming of the isolate as C. flavescens. [source] Ultrastructural and Immunocytochemical Studies on Effects of Barley Yellow Dwarf Virus , Infection on Fusarium Head Blight, Caused by Fusarium graminearum, in Wheat PlantsJOURNAL OF PHYTOPATHOLOGY, Issue 1 2006Y. Liu Abstract The interactions between barley yellow dwarf virus (BYDV) and Fusarium head blight (FHB), caused by Fusarium graminearum, were studied in the two winter wheat cultivars (cvs.), Agent (susceptible to FHB) and Petrus (moderately resistant to FHB), using ultrastructural and immunocytochemical methods. Infections of wheat plants of both cvs. by BYDV increased susceptibility to FHB. BYDV infection caused numerous cytological changes in lemma tissue of both cvs. such as formation of vesicles in the cytoplasm, degradation of fine structures of chloroplasts of both cvs. and accumulation of large starch grains in the chloroplasts. Electron microscopical studies showed that the development of F. graminearum on spike surfaces was not affected in BYDV-infected plants. After penetration and intercellular growth in lemma tissue, defence responses to Fusarium infections were markedly reduced in BYDV-diseased plants compared to the tissue of virus-free plants. At sites of contact of fungal cells with host tissue, depositions of cell wall material were distinctly less pronounced than in tissues of virus-free plants of cv. Petrus. Detection of , -1,3-glucanases and chitinases in lemma tissue of cv. Agent revealed no appreciably increased accumulation of both defence enzymes in F. graminearum -infected virus-free and BYDV-infected tissues compared to the non-infected control tissue. On the other hand, in cv. Petrus, infection with F. graminearum induced a markedly enhanced activity of both enzymes 3 days after inoculation. The increase of both enzyme activities was less pronounced in BYDV-infected plants than in tissue exclusively infected with F. graminearum. Cytological studies suggest that in contrast to the susceptible cv. Agent postinfectional defence responses may play still an important role in the resistance of the moderately resistant cv. Petrus to FHB. [source] Studies on Symptom Development, Phenolic Compounds and Morphological Defence Responses in Wheat Cultivars Differing in Resistance to Fusarium Head BlightJOURNAL OF PHYTOPATHOLOGY, Issue 4-5 2002E. SIRANIDOU Abstracts Development of Fusarium head blight, caused by Fusarium culmorum, was evaluated in seven winter wheat cultivars and one spring wheat cultivar in an outdoor pot experiment. Of the cultivars studied, only the spring wheat cultivar Frontana exhibited resistance against initial infection by the fungus. Significantly higher amounts of free phenolic compounds were found in the glumes, lemmas and paleas of Frontana prior to and at all sampling times after inoculation, in comparison to the winter wheat cultivar Agent. Furthermore, the amount of p -coumaric acid increased significantly in the glumes, lemmas and paleas of the cultivar Frontana 2 days after inoculation compared to that in uninoculated spike tissues. However, the amounts of ferulic acid between inoculated and healthy plant tissues did not differ. Ultrastructural studies indicated more pronounced structural defence responses, such as cell wall appositions, in the infected lemma tissue of the resistant cultivar Frontana compared with the susceptible Agent 3 days after inoculation. Immunogold labelling of lignin revealed no differences in the density of gold particles on the cell walls of healthy lemma tissue in the two cultivars. However, density of particles increased in the infected lemma tissue of the resistant cultivar Frontana on cell walls adjacent to the fungal cells, 3 days after inoculation. The susceptible cultivar Agent showed little or no response to the infection. Phenolic compounds appear to play a role in the resistance of the cultivar Frontana to F. culmorum. [source] Solid substrate bioassay to evaluate impact of Trichoderma on trichothecene mycotoxin production by Fusarium speciesJOURNAL OF THE SCIENCE OF FOOD AND AGRICULTURE, Issue 3 2008Maciej Busko Abstract BACKGROUND:Fusarium head blight of wheat is a destructive disease in various wheat-growing regions and leads to significant yield losses for farmers and to contamination of cereal grains with mycotoxins, mainly deoxynivalenol and its derivatives. Toxigenic Fusarium species sporulate on cereal crop residues and produce significant inoculum for epidemics. Reduction of mycotoxin production and pathogen sporulation may be influenced by saprophytic fungal antagonists. RESULTS:Trichoderma isolates were examined in dual culture bioassays on rice with two isolates of Fusarium graminearum Schwabe and two isolates of Fusarium culmorum (W.G. Smith) Saccardo, belonging to three different chemotypes. Production of five trichothecene mycotoxins, deoxynivalenol (DON), 3-acetyl-deoxynivalenol (3AcDON), 15-acetyl-deoxynivalenol (15AcDON), nivalenol (NIV) and fusarenone X (FUS), was reduced by over 95%. Two Trichoderma isolates partially reduced the amounts of four metabolites when inoculated on autoclaved cultures of the same four Fusarium isolates. However, in the case of the 15AcDON chemotype of F. culmorum culture the content of DON increased and that of 15AcDON decreased. Isolates of Trichoderma varied in their ability to inhibit production of the five trichothecene mycotoxins by Fusarium. Susceptibility of the four Fusarium isolates to antagonistic activity of the same Trichoderma isolate differed significantly. CONCLUSION: Selected non-toxigenic Trichoderma isolates proved to be useful biocontrol agents against toxigenic Fusarium pathogens of wheat, significantly reducing production of the trichothecene mycotoxins DON, NIV and their acetylated derivatives. Copyright © 2007 Society of Chemical Industry [source] Involvement of trichothecenes in fusarioses of wheat, barley and maize evaluated by gene disruption of the trichodiene synthase (Tri5) gene in three field isolates of different chemotype and virulenceMOLECULAR PLANT PATHOLOGY, Issue 6 2006FRANK J. MAIER SUMMARY Fusarium graminearum is the main causative agent of Fusarium head blight on small grain cereals and of ear rot on maize. The disease leads to dramatic yield losses and to an accumulation of mycotoxins. The most dominant F. graminearum mycotoxins are the trichothecenes, with deoxynivalenol and nivalenol being the most prevalent derivatives. To investigate the involvement of trichothecenes in the virulence of the pathogen, the gene coding for the initial enzyme of the trichothecene pathway was disrupted in three field isolates, differing in chemotype and in virulence. From each isolate three individual disruption mutants were tested for their virulence on wheat, barley and maize. Despite the different initial virulence of the three wild-type progenitor strains on wheat, all disruption mutants caused disease symptoms on the inoculated spikelet, but the symptoms did not spread into other spikelets. On barley, the trichothecene deficient mutants showed no significant difference compared to the wild-type strains: all were equally aggressive. On maize, mutants derived from the NIV-producing strain caused less disease than their wild-type progenitor strain, while mutants derived from DON-producing strains caused the same level of disease as their progenitor strains. These data demonstrate that trichothecenes influence the virulence of F. graminearum in a highly complex manner, which is strongly host as well as moderately chemotype specific. [source] Mapping of QTL for resistance against Fusarium head blight in the winter wheat population Pelikan//Bussard/Ning8026PLANT BREEDING, Issue 1 2009J. Häberle Abstract We report on the identification of FHB (Fusarium head blight) resistance quantitative trait loci (QTL) of the donor ,G93010' (Bussard/Ning8026) in the background of elite breeding material adapted to the central European climate. With a multiple interval mapping method, two major resistance QTL were identified. Qfhs.lfl-7BS/5BL and Qfhs.lfl-6BS reduced FHB severity individually by 30% and 24%. The combination of both QTL decreased disease severity most effectively by about one half. Qfhs.lfl-6BS is most likely identical to Fhb2, thus, the effectiveness of Fhb2 in central European breeding material has been validated. Qfhs.lfl-7BS/5BL overlapped with QTL for plant height and heading date. Nevertheless, the selection of lines combining a good FHB resistance level with an acceptable plant height was possible. As the donors of the QTL have probably not yet been utilized in European breeding material, we identified well-adapted lines of the mapping population as valuable donors for marker-assisted breeding programmes. [source] Effect of the Rht-D1 dwarfing locus on Fusarium head blight rating in three segregating populations of winter wheatPLANT BREEDING, Issue 4 2008H.-H. Voss Abstract Fusarium head blight (FHB) is one of the major fungal diseases in wheat throughout the world. To control FHB severity, breeding genetically resistant varieties is thought to be the most promising strategy. In wheat breeding programmes, short cultivars predominantly carrying the Norin 10 derived semi-dwarfing allele Rht-D1b (Rht2) are preferred worldwide because of higher achievable grain yields and lower risk of lodging. This study was conducted to determine the influence of different alleles at the Rht-D1 locus on FHB reaction. Three winter wheat populations were produced by crossing rather susceptible varieties ,Biscay', ,Pirat' and ,Rubens' carrying mutant-type allele Rht-D1b with the more resistant varieties ,Apache', ,Romanus' and ,History' containing the Rht-D1a wild-type allele (rht2). The 190, 216 and 103 progeny of the F4 -derived populations were assayed for the presence of Rht-D1a or Rht-D1b, plant height, and mean FHB rating after spray inoculation at flowering time with a highly aggressive isolate of Fusarium culmorum. Comparably, high mean FHB severities ranging from 28% to 49% for all population × environment combinations were achieved, with significant genotypic variation for FHB rating and plant height within all populations. Both traits were negatively correlated with r ranging from ,0.48 to ,0.61 in the complete populations. However, within the subpopulations homozygous for one or other height allele these correlations decreased considerably. The Rht-D1b semi-dwarfing allele resulted in 7,18% shorter plants, depending on the population, but a considerably increased FHB reaction of 22,53%. Nevertheless, significant genotypic variance for FHB resistance remained in all tested Rht-D1b subpopulations indicating that selection for moderately FHB resistant genotypes within agronomically beneficial Rht-D1b genotypes is still feasible. [source] A reciprocal backcross monosomic analysis of the scab resistant spring wheat (Triticum aestivum L.) cultivar, ,Frontana'PLANT BREEDING, Issue 3 2007W. A. Berzonsky Abstract Fusarium head blight (FHB) reduces grain yield of spring wheat and results in deoxynivalenol (DON) production. The resistant spring wheat ,Frontana' could be used to diversify and pyramid resistance. A backcross reciprocal monosomic analysis was conducted to identify ,Frontana' chromosomes with resistance. Disomic lines, one set containing critical chromosomes from ,Frontana' and the other containing chromosomes from susceptible ,Chris', were spray-inoculated and evaluated in separate greenhouse studies (GH-1 and GH-2). Measurements were disease severity at 7 and 14 days, visually diseased kernels, kernels/g and DON content. In GH-2, reciprocal chromosome lines 5B for visually diseased kernels, lines 7B for kernels/g, and lines 4B and 6A for DON content were significantly different. Lines with ,Frontana' chromosomes 3A, 6A and 4D reduced visually diseased kernels, kernels/g and DON content in both studies, while ,Frontana' chromosomes 2A, 2B, 4B and 7B increased susceptibility, as indicated by an increase in these same measurements in both studies. Genes carried on ,Frontana' chromosomes 3A, 6A and 4D could be useful for diversifying and pyramiding sources of FHB resistance. [source] Fusarium head blight resistance derived from Lophopyrum elongatum chromosome 7E and its augmentation with Fhb1 in wheatPLANT BREEDING, Issue 5 2006X. Shen Abstract The objective of this study was to assess the effectiveness of Fusarium head blight (FHB) resistance derived from wheatgrass Lophopyrum elongatum chromosome 7E and to determine whether this resistance can augment resistance in combination with other FHB resistance quantitative trait loci (QTL) or genes in wheat. The ,Chinese Spring',Lophopyrum elongatum disomic substitution line 7E(7B) was crossed to three wheat lines: ,Ning 7840', L3, and L4. F2 populations were evaluated for type II resistance with the single-floret inoculation method in the greenhouse. Simple sequence repeat markers associated with Fhb1 in ,Ning 7840' and L4 and markers located on chromosome 7E were genotyped in each population. Marker,trait association was analysed with one-way or two-way analysis of variance. The research showed that, in the three populations, the average number of diseased spikelets (NDS) in plants with chromosome 7E is 1.2, 3.1 and 3.2, vs. NDS of 3.3, 7.2 and 9.1 in plants without 7E, a reduction in NDS of 2.1, 4.1 and 5.9 in the respective populations. The QTL on 7E and the Fhb1 gene augment disease resistance when combined. The effect of the QTL on 7E was greater than that on 3BS in this experiment. Data also suggest that the FHB resistance gene derived from L. elongatum is located on the long arm of 7E. [source] Development of molecular markers for crown rot resistance in wheat: mapping of QTLs for seedling resistance in a ,2-49' × ,Janz' populationPLANT BREEDING, Issue 6 2005B. C. Y. Collard Abstract Crown rot, caused by Fusarium pseudograminearum, is an important disease of wheat in Australia and elsewhere. In order to identify molecular markers associated with partial seedling resistance to this disease, bulked segregant analysis and quantitative trait loci (QTL) mapping approaches were undertaken using a population of 145 doubled haploid lines constructed from ,2-49' (partially resistant) × ,Janz' (susceptible) parents. Phenotypic data indicated that the trait is quantitatively inherited. The largest QTLs were located on chromosomes 1D and 1A, and explained 21% and 9% of the phenotypic variance, respectively. Using the best markers associated with five QTLs identified by composite interval mapping, the combined effect of the QTLs explained 40.6% of the phenotypic variance. All resistance alleles were inherited from ,2-49' with the exception of a QTL on 2B, which was inherited from ,Janz'. A minor QTL on 4B was loosely linked (19.8 cM) to the Rht1 locus in repulsion. None of the QTLs identified in this study were located in the same region as resistance QTLs identified in other populations segregating for Fusarium head blight, caused by Fusarium graminearum. [source] QTL analysis of resistance to Fusarium head blight in wheat using a ,Wangshuibai'-derived populationPLANT BREEDING, Issue 4 2005M. Mardi Abstract Fusarium head blight (FHB) is a devastating disease that reduces the yield, quality and economic value of wheat. For quantitative trait loci (QTL) analysis of resistance to FHB, F3 plants and F3:5 lines, derived from a ,Wangshuibai' (resistant)/,Seri82'(susceptible) cross, were spray inoculated during 2001 and 2002, respectively. Artificial inoculation was carried out under field conditions. Of 420 markers, 258 amplified fragment length polymorphism and 39 simple sequence repeat (SSR) markers were mapped and yielded 44 linkage groups covering a total genetic distance of 2554 cM. QTL analysis was based on the constructed linkage map and area under the disease progress curve. The analyses revealed a QTL in the map interval Xgwm533-Xs18/m12 on chromosome 3BS accounting for up to 17% of the phenotypic variation. In addition, a QTL was detected in the map interval Xgwm539-Xs15/m24 on chromosome 2DL explaining up to 11% of the phenotypic variation. The QTL alleles originated from ,Wangshuibai' and were tagged with SSR markers. Using these SSR markers would facilitate marker-assisted selection to improve FHB resistance in wheat. [source] Molecular cytogenetic analysis of a durum wheat ×Thinopyrum distichum hybrid used as a new source of resistance to Fusarium head blight in the greenhousePLANT BREEDING, Issue 5 2001Q. Chen Abstract Fusarium head blight (FHB, scab), caused by Fusarium graminearum Schwabe, is a serious and damaging disease of wheat. Although some hexaploid wheat lines express a good level of resistance to FHB, the resistance available in hexaploid wheat has not yet been transferred to durum wheat. A germplasm collection of Triticum durum× alien hybrid lines was tested as a potential source of resistance to FHB under controlled conditions. Their FHB reaction was evaluated in three tests against conidial suspensions of three strains of F. graminearum at the flowering stage. Two T. durum×Thinopyrum distichum hybrid lines, ,AFR4' and ,AFR5,, expressed a significantly higher level of resistance to the spread of FHB than other durum-alien hybrid lines and a resistant common wheat line ,Nyu-Bay'. Genomic in situ hybridization using total genomic DNA from alien grass species demonstrated that ,AFR5' had 13 or 14 alien genome chromosomes plus 27 or 28 wheat chromosomes, while ,AFR4' had 22 alien genome and 28 wheat chromosomes. All of the alien chromosomes present in these two lines belonged to the J genome. ,AFR5' is likely to be more useful as a source of FHB resistance than ,AFR4' because of its relatively normal meiotic behaviour, high fertility and fewer number of alien chromosomes. ,AFR5' shows good potential as a source for transferring FHB resistance gene into wheat. The development of T. durum addition lines carrying resistance genes from ,AFR5' is underway. [source] Chromosomal location of Fusarium head blight resistance genes and analysis of the relationship between resistance to head blight and brown foot rotPLANT BREEDING, Issue 1 2000A. Mentewab Abstract In order to identify chromosomes involved in resistance to Fusarium head blight, a set of 21 substitution lines of Triticum macha (resistant) chromosomes into ,Hobbit''sib'(susceptible) were evaluated in trials over 2 years. For the first year's trial, all plants were inoculated on the same day with a conidial suspension of F. culmorum. For the second trial, individual plants were inoculated precisely at mid anthesis of each plant over a period of 2 weeks. The disease level was assessed by visual scoring, relative ear weight and F. culmorumn -specfic quantitative polymerase chain reaction. The results showed that T. macha chromosomes 1B, 4A and 7A conferred good overall resistance, suggesting that they carry important genes for resistance. In two additional trials, T. macha and ,Hobbit''sib' were evaluated for resistance to brown foot rot. The results showed that T. macha was more susceptible than ,Hobbit',sib', indicating that stem base disease response is not correlated with head blight resistance in these cultivars. [source] Geographic distribution and genetic diversity of Fusarium graminearum and F. asiaticum on wheat spikes throughout ChinaPLANT PATHOLOGY, Issue 1 2008B. Qu A large number of Fusarium graminearum and F. asiaticum isolates were collected from wheat spikes from all regions in China with a history of fusarium head blight (FHB) epidemics. Isolates were analysed to investigate their genetic diversity and geographic distribution. Sequence characterized amplified region (SCAR) analyses of 437 isolates resolved both species, with 21% being F. graminearum (SCAR type 1) and 79% being F. asiaticum (SCAR type 5). AFLP profiles clearly resolved two groups, A and B, that were completely congruent with both species. However, more diversity was detected by AFLP, revealing several subgroups within each group. In many cases, even for isolates from the same district, AFLP haplotypes differed markedly. Phylogenetic analyses of multilocus DNA sequence data indicated that all isolates of SCAR type 1, AFLP group A were F. graminearum, whilst isolates of SCAR type 5, AFLP group B were F. asiaticum, demonstrating that it is an efficient method for differentiating these two species. Both species seem to have different geographic distributions within China. Fusarium graminearum was mainly obtained from wheat growing in the cooler regions where the annual average temperature was 15°C or lower. In contrast, the vast majority of F. asiaticum isolates were collected from wheat growing in the warmer regions where the annual average temperature is above 15°C and where FHB epidemics occur most frequently. This is the first report of the distribution of, and genetic diversity within, F. graminearum and F. asiaticum on wheat spikes throughout China. [source] | ||||||||||