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1.
The occurrence and geographic distribution of powdery mildew on cucurbits was studied in the Czech Republic, Austria, France, Germany, Great Britain, Italy, Slovakia, Slovenia, Spain, Switzerland, the Netherlands, as well as in Turkey and Israel, during the period 1995–2000. In total, 599 leaf samples with powdery mildew symptoms were collected from cucurbits from 166 locations within the Czech Republic; an additional 69 samples were collected from 42 locations elsewhere. Two powdery mildews (Golovinomyces cichoracearum and Podosphaera xanthii) were identified. The host range included the cultivated cucurbits (Cucumis melo, C. sativus, Cucurbita pepo, C. maxima, and Citrullus lanatus) and several other species. P. xanthii was the only powdery mildew pathogen found in Spain, Israel, and Turkey. P. xanthii and G. cichoracearum were detected in the other surveyed countries, occurring in both mixed infections and separately. In the more northerly latitudes and higher elevations, G. cichoracearum is more often the single species. G. cichoracearum was the dominant powdery mildew species in the Czech Republic (detected in 98.8% of the locations there); P. xanthii was found as the lone species in 1.2% of locations. At 28.4% of locations, G. cichoracearum was found with P. xanthii as mixed infections. The hyperparasitic fungus Ampelomyces quisqualis was found in 30% of the samples from the Czech Republic and was also recorded in Austria, Italy, Slovenia, Switzerland, and Great Britain.  相似文献   

2.
Microcyclic conidiogenesis (MC), a process defined as the production of conidia on a spore without any, or only a minimal, involvement of hyphal growth, has recently been reported in a little known powdery mildew species, Oidium longipes. To investigate whether this was an isolated case or it is a more general phenomenon in powdery mildew fungi, germinating conidia of eight species of the Erysiphales were examined using light microscopy. The following species were included in this work: Erysiphe necator on grapevine, Blumeria graminis f. sp. hordei on barley, Podosphaera xanthii on cucumber, Erysiphe sp. on Ligustrum vulgare, O. longipes on Petunia x grandiflora, O. neolycopersici on tomato, Golovinomyces cichoracearum on Rudbeckia laciniata and Sawadaea sp. on Acer negundo. In all these species, up to 4% of the germinated conidia exhibited MC. Moreover, when colonies of E. necator and O. neolycopersici, on detached grapevine and tomato leaves, respectively, were treated with a conidial suspension of Ampelomyces, the intracellular pycnidia of these mycoparasites appeared in microcyclic conidiophores. This represents a yet undescribed method of accelerating asexual reproduction in this mycoparasite. In the life cycle of powdery mildews, the importance of MC is still not clear but it should be taken into consideration when conidial germination is studied on the host surface for purposes such as epidemiology or species identification.  相似文献   

3.
Podosphaera xanthii and Golovinomyces orontii are the causal agents of cucurbit powdery mildew. The effect of temperature on conidial germination, infection and sporulation was studied under controlled conditions. Conidia were inoculated on cucumber leaf discs, and incubated at six constant temperatures (from 10 to 35 °C in 5 °C steps) for 3 to 72 h to evaluate conidial germination and infection, and for 6–15 days to evaluate sporulation intensity. Germination took place at all tested temperatures, but was close to zero at 35 °C. The longest germ tubes measured in this experiment were 141.74 μm for the secondary germ tube of Pxanthii at 20 °C after 48 h of incubation, and 67.92 μm for G. orontii for the primary germ tube at 20 °C after 48 h of incubation. The optimal temperatures for conidial germination, infection and sporulation were 24.4, 25.7 and 22.3 °C, respectively, for P. xanthii, and 17.9, 17.3 and 14.9 °C, respectively, for G. orontii. Equations were developed to describe conidial germination with a coefficient of determination (R2) of 0.85 and 0.90 for P. xanthii and Gorontii, respectively. Infection equations resulted in R2 of 0.94 and 0.93 for Pxanthii and Gorontii, respectively; and for sporulation, R2 of 0.75 and 0.76 for P. xanthii and G. orontii respectively, as a function of temperature. These results can be used to develop models for the risk of cucurbit powdery mildew under field conditions.  相似文献   

4.
The present study aimed to explore the possibility of using the type I trichomes of tomato (Solanum lycopersicum) to monitor the infection processes of powdery mildews by microscopy. Individual trichome cells of two tomato genotypes were inoculated with pathogenic and non-pathogenic powdery mildew species, Pseudoidium neolycopersici, Erysiphe trifoliorum and Podosphaera xanthii. On the trichome cells of the tomato cultivar Moneymaker, hyphae of the pathogenic Pseudoidium neolycopersici (isolates KTP-03 and KTP-04) grew vigorously; whereas hyphal growth of the non-pathogenic Erysiphe trifoliorum and Podosphaera xanthii ceased after appressorium formation, which was associated with papilla formation and hypersensitive cell death, respectively. Similar infection processes of the tested powdery mildews were seen in Moneymaker epidermal cells. Therefore, tomato trichomes are suitable for analysing, at individual cell level, the infection processes of different pathotypes of powdery mildews and for observing the cytological responses of plants by non-pathogenic powdery mildews. On the other hand, it was observed that both isolates KTP-03 and KTP-04 failed to produce conidiophores on the hyphae elongating on Moneymaker trichomes. Similarly, no conidiophores were produced on the hyphae elongating on trichomes of Solanum peruvianum LA2172, which is resistant to KTP-03 and susceptible to KTP-04. Interestingly, delayed cell death occurred in LA2172 epidermal cells, which were attacked by KTP-03 hyphae elongating from trichomes and conidiophores were formed on new hyphae growing from the leaf epidermal cells. Thus, leaf trichomes and epidermal cells of the wild tomato species LA2172 reacted differently to the avirulent isolate KTP-03.  相似文献   

5.
To clarify relationships between powdery mildew resistance of cucumber and hyphal developmental state of the pathogen, haustorial formation and the hyphal branching frequency were compared among cucumber varieties that differ in resistance levels to powdery mildew pathogen Podosphaera xanthii. Cotyledons of four cultivars were inoculated with P. xanthii. By 2 days after inoculation, secondary haustoria had developed from the first hyphal cell that formed beside the conidium in susceptible cultivars. The fungus on susceptible cultivars also tended to have hyphal branches just after the hyphal cell producing haustoria.  相似文献   

6.
Powdery mildew is one of the most common and severe diseases of cucurbits, causing heavy yield losses in all growing areas when not successfully controlled. Two different fungal species, Podosphaera xanthii and Golovinomyces orontii, are generally recognized as causal agents. The results of monitoring carried out in 2016 and 2018 confirmed that P. xanthii is the exclusive pathogen causing cucurbit powdery mildew (CPM) in southern Italy. P. xanthii is a bipolar heterothallic fungus; a PCR-based method for distinguishing MAT1-1 and MAT1-2 idiomorphs was applied for assessing mating type distribution in fungal populations present on cucurbits in different sites. The idiomorph MAT1-2 was prevalent over the MAT1-1 in 2016, whereas they were approximately in a 1:1 ratio in 2018; this finding corroborated the hypothesis that the MAT1-1 idiomorph was more recently introduced in the area. Cyflufenamid-resistant isolates were widespread in commercial greenhouses and field plantings even though use of this fungicide had been drastically reduced by the farmers 1 year before the monitoring due to the effectiveness losses observed in CPM control. Occurrence of cyflufenamid resistance and its impact on efficacy were evaluated in a field trial comparing different fungicide spray schedules. Cyflufenamid-resistant isolates were detected even at the first appearance of symptoms on leaves, increasing over time. Isolates resistant to cyflufenamid showed a resistance factor as high as 900. Generally, P. xanthii was better controlled when cyflufenamid was used in integrated strategies rather than in spray schedules based on the exclusive use of the fungicide.  相似文献   

7.
Abstract

Among various organisms attacking skeleton weed, Chondrilla juncea, in Europe, a rust fungus, Puccinia chondrillina, and two powdery mildews, Erysiphe cichoracearum and Leveillula taurica f. sp. chondrillae were found to be the most damaging. P. chondrillina was shown to be specific to Chondrilla and has recently been introduced into Australia where skeleton weed is important in wheat cultivation. It has already spread throughout the weed infestations and is beginning to reduce plant populations. Encouraged by the result of the Chondrilla rust, the possible use of Puccinia xanthii for the control of Xanthium strumarium and X. spinosum has also been studied. Current research on the possible use of phytopathogens in the biological control of some other weeds is reviewed.  相似文献   

8.
In 2002, a powdery mildew with catenate conidia lacking fibrosin bodies was found on cucumber in a greenhouse in Kanagawa Prefecture, Japan. Morphological observation revealed that the fungus belongs to Oidium subgenus Reticuloidium, anamorph of the genus Golovinomyces. Molecular phylogenetic analyses of the nucleotide sequences of the rDNA ITS regions and D1/D2 domains of the 28S rDNA indicated that the fungus belongs to the clade of G. orontii with other Golovinomyces fungi from a wide range of host plants, suggesting that the fungus was newly transported from abroad. Because there has been no prior report of cucumber powdery mildew caused by Reticuloidium, further research on the physiology, epidemiology, control and resistant cucumber varieties is required.  相似文献   

9.
BACKGROUND: Physcion is a key active ingredient of the ethanol extract from roots of Chinese rhubarb (Rheum officinale Baill.) that has been commercialised in China for controlling powdery mildews. The biological mechanism of action of physcion against the barley powdery mildew pathogen was studied using bioassay and microarray methods. RESULTS: Bioassay indicated that physcion did not directly affect conidial germination of Blumeria graminis Speer f. sp. hordei Marchal, but significantly inhibited conidial germination in vivo. Challenge inoculation indicated that physcion induced localised resistance rather than systemic resistance against powdery mildew. Gene expression profiling of physcion‐treated barley leaves detected four upregulated and five downregulated genes (ratio ≥ 2.0 and P‐value < 0.05) by using an Affymetrix Barley GeneChip. The five upregulated probe sequences blasted to the same barley leaf‐specific thionin gene, with significant changes varying from 4.26 to 19.91‐fold. All downregulated genes were defence‐related, linked to peroxidase, oxalate oxidase, bsi1 protein and a pathogenesis‐related protein. These changes varied from ? 2.34 to ? 2.96. Quantitative real‐time PCR data confirmed that physcion enhanced the gene expression of leaf‐specific thionin of barley. CONCLUSION: Results indicated that physcion controls powdery mildew mainly through changing the expression of defence‐related genes, and especially enhancing expression of leaf‐specific thionin in barley leaves. Copyright © 2010 Society of Chemical Industry  相似文献   

10.
Since 2003, Torenia fournieri plants grown for experimental purposes were repeatedly infected by powdery mildew in a laboratory in Hungary. Based on morphological characteristics, the pathogen belonged to the mitosporic genus Oidium subgen. Reticuloidium, the anamorph stage of Golovinomyces. The rDNA ITS sequence was identical to that of two other powdery mildew fungi, infecting Arabidopsis and Veronica, respectively, in different parts of the world. According to a previous phylogenetic analysis of ITS and 28S rDNA sequences, those two powdery mildews belong to a recently evolved group of Golovinomyces characterized by multiple host range expansions during their evolution. Both the ITS sequence and the morphological data indicate that the powdery mildew anamorph infecting Torenia also belongs to this group. It is likely that the powdery mildew infections of the experimental T. fournieri plants, native to south-east Asia, were the result of a very recent host range expansion of a polyphagous Golovinomyces because (i) T. fournieri is absent from our region, except as an experimental plant grown in the laboratory, (ii) the powdery mildew fungus infecting this exotic plant belongs to a group of Golovinomyces where host range expansion is a frequent evolutionary scenario, (iii) cross-inoculation tests showed that this pathogen is also able to infect other plant species, notably A. thaliana and tobacco, and (iv) no Golovinomyces species are known to infect T. fournieri anywhere in the world. Although host range expansion has often been proposed as a common evolutionary process in the Erysiphales, and also in other biotrophic plant pathogens, this has not been clearly demonstrated in any case studies so far. To our knowledge, this is the first convincing case of a host range expansion event in the Erysiphales.  相似文献   

11.
Apple shoots and aerial parts of 13 other plant species infected with powdery mildews during the previous season were collected in late winter and early spring between 1998 and 2003 at a total of 34 sample sites in Hungary. Samples were examined for the presence of overwintering structures of Ampelomyces, common mycoparasites of powdery mildews. Pycnidia and resting hyphae resembling those of Ampelomyces were found on six plant species, including apple. Their viability and subsequent mycoparasitic activity of the hyphae emerging from the overwintered fungal structures were studied in vitro to determine whether they can serve as sources of primary inocula of Ampelomyces in the spring. Overwintered pycnidia of Ampelomyces collected in the spring, and produced in both the ascomata and the conidiophores of powdery mildews during the previous season, initiated the life cycle of these mycoparasites when placed close to fresh powdery mildew colonies in vitro. Similarly, thick-walled resting hyphae, found in the dried powdery mildew mycelia which covered the overwintered aerial parts of the host plants, also germinated and gave rise to new intracellular pycnidia of Ampelomyces when powdery mildew colonies were inoculated with them in vitro. On apple trees, Ampelomyces mycoparasites overwintered as resting hyphae in the dried powdery mildew mycelia covering the shoots and in the parasitized ascomata of Podosphaera leucotricha on the bark and the scales of the buds. Approximately 31% of the field samples collected from apple trees in spring between 1998 and 2003 contained overwintered structures of Ampelomyces. Artificial bursting of apple buds in the laboratory showed that both P. leucotricha and Ampelomyces start their life cycle during or soon after bud burst, but Ampelomyces can only slowly follow the spread of its mycohost on infected leaves. Most probably, the mycoparasites did not overwinter in the dormant hyphae of P. leucotricha in the buds, but only on the bark and the bud scales, as their hyphae were not found in the young hyphae of apple powdery mildew that appeared on the leaf tissues during bud burst. This study demonstrated that Ampelomyces mycoparasites can survive the winter in the field as pycnidia and as resting hyphae in the dried mycelia of their mycohosts.  相似文献   

12.
Podosphaera macularis, the causal agent of hop powdery mildew, is known to produce chasmothecia (formerly cleistothecia) in eastern North America and Europe. Ascocarps have not yet been reported from the Pacific Northwestern region of North America. Reasons for the apparent absence of chasmothecia in the Pacific Northwest were unknown. This study established that Pmacularis is heterothallic and ascocarp ontogeny, maturation, dehiscence and ascospore infection proceed similarly to other powdery mildew fungi. Genome sequencing of a MAT1‐1 isolate revealed the structure of the MAT1 locus and presence of MAT1‐1‐3, demonstrating further similarities to other powdery mildew fungi. PCR assays with primers designed from conserved domains of the MAT1 idiomorphs were developed to characterize the frequency of idiomorphs in populations of P. macularis. Amongst 317 samples of P. macularis collected during 2012 and 2013 from the Pacific Northwest only the MAT1‐1 idiomorph was found. In contrast, among 56 samples from the eastern United States and Europe, MAT1‐1 and MAT1‐2 idiomorphs were detected at equivalent frequencies. At temperatures representative of late season conditions in the Pacific Northwest, chasmothecia formed readily when a Pacific Northwest MAT1‐1 isolate was paired with a MAT1‐2 isolate collected from outside the region. Although these findings do not encompass all climatic, geographic or temporal barriers that could inhibit the formation of chasmothecia, the current absence of the ascigerious stage of Pmacularis in the Pacific Northwest could be explained by the absence of the MAT1‐2 mating type idiomorph.  相似文献   

13.
D-pinitol is an effective agent for controlling powdery mildew (Podosphaera xanthii) in cucumber. In this study, we determined the mechanisms of D-pinitol in controlling powdery mildew in cucumber plants. We compared P. xanthii development on cucumber leaf surface treated with D-pinitol or water (2 mg ml−1) at different time points after inoculation. The germinating conidia, hyphae, and conidiophores of the pathogen were severely damaged by D-pinitol at any time of application tested. The highest level of suppression of fungal development was obtained at 3 days after inoculation. The contents of chlorophyll, total phenolics, flavonoid, and gallic acid and its derivatives (GAD); the activities of phenylalanine ammonialyase (PAL), polyphenoloxidase (PPO), peroxidase (POX), and superoxide dismutase (SOD); and the expression of the genes encoding for PR-1, peroxidase (POX), lipoxygenase (LOX1), chitinase (Chit1) were higher in the cucumber leaves treated with D-pinitol and inoculated than in the leaves either treated with D-pinitol or inoculated with the pathogen. These results suggest that D-pinitol triggers several plant defense responses in cucumber leading to pathogen suppression and resistance to powdery mildew.  相似文献   

14.
During the past years, nrDNA ITS sequences have supported the identification of many powdery mildew fungi because comprehensive analyses showed that differences in these sequences have always correlated with the delimitation of different species and formae speciales of the Erysiphales. Published data, obtained using direct sequencing of the PCR products, suggested that even one to five nucleotide differences in the ITS sequences delimit different, albeit closely related, species, and/or indicate differences in host range patterns. Here we show that such differences in the ITS sequences can be detected even in a single sample of a powdery mildew fungus. We sequenced the ITS region in 17 samples, representing six powdery mildew species, both directly and after cloning the PCR products. Among these, samples of O. longipes exhibited two or three, samples of O. neolycopersici three or four, those of an Oidium sp. from Chelidonium majus up to seven, and a sample of another Oidium sp. from Passiflora caerulea two different ITS types determined after cloning. No ITS nucleotide polymorphisms were found in samples of O. lycopersici and Erysiphe aquilegiae. This suggests that some powdery mildew taxa are more variable at the ITS level than others. Thus, although the ITS sequences determined by direct sequencing represent robust data useful in delimitation and phylogenetic analysis of distinct species of the Erysiphales, these need to be used with precaution, and preferably determined after cloning, especially when dealing with closely related taxa at species and sub-species levels. With this method a hitherto undetected genetic diversity of powdery mildews can be revealed.  相似文献   

15.
A strain of Podosphaera clandestina has been highlighted as a priority pest threat to the Australian cherry industry. Australia currently has no records of powdery mildew on cherry (Prunus avium). P. clandestina is reported to cause disease on a range of Rosaceae genera including Crataegus and Prunus; in Australia, P. clandestina has only been recorded on Crataegus. A recent species revision identified Podosphaera cerasi on P. avium as a separate species from P. clandestina. Therefore, a revision of which powdery mildew species is present in Australia on Crataegus is required to inform Australian plant biosecurity. Reference collection specimens from the Victorian Plant Pathology Herbarium (VPRI) recorded as Podosphaera spp. collected between 1889 to 2008 on cherry and three other host plant genera from Australia and overseas were sampled for DNA extraction and next-generation sequencing (NGS). Sequence data from preserved specimens were successfully mapped to internal transcribed spacer (ITS) sequences of P. clandestina in the strict sense, P. cerasi, and Podosphaera prunicola, and chloroplast matK sequences were used to identify plant hosts. Australian specimens on Crataegus hosts were P. clandestina in the strict sense and specimens on Prunus from the USA were identified as P. cerasi and P. prunicola. The outcome of this study confirmed the powdery mildew on Australian Crataegus specimens to be P. clandestina and none of the cherry powdery mildews (Podosphaera pruni-avium, P. cerasi, or P. prunicola) are present on Australian specimens in the VPRI collection, which suggests they are not present in Australia.  相似文献   

16.
This study examines the effects of a vegetable fungicide on sugar beet powdery mildew (Erysiphe betae) and cucumber powdery mildew (Erysiphe cichoracearum). The formulations consisting of a dispersion of Brassicaceae meal in vegetable or mineral oils on infected leaves of sugar beet, reared in the greenhouse, and of musk melons cultivated under plastic tunnels, were tested in comparison to each oil taken separately. Both formulations containing Brassicaceae meals, caused 94% of conidia to be distorted while for the untreated group only 2% were distorted. Furthermore, the leaf area infected by E. betae was 56% for untreated plants and 2.7 and 9.9% respectively, for plants treated with meal containing mineral and vegetable oil. Vegetable oil considered separately or with Brassicaceae meals showed no phytotoxicity, while the formulations based on mineral oil showed a significantly lower fresh and dry weight on tomato plants. The low level or absence of phytotoxicity of plants treated with vegetable oil formulations suggests that to improve the efficacy of powdery mildew control, they could be used mixed with sulphur. The efficiency of the vegetable formulations in the powdery mildew control observed during these trials encourages further investigation on other parasitic fungi and foliar pathogens.  相似文献   

17.
Since 2001, several isolates of Blumeria graminis, the causal agent of cereal powdery mildew, maintained on detached leaves at the John Innes Centre, Norwich, UK, have spontaneously become infected with an unknown filamentous fungus whose mycelia have quickly overgrown the powdery mildew colonies and destroyed them completely. A total of five isolates of the contaminant were obtained and identified as Paecilomyces farinosus based on morphological characteristics and rDNA ITS sequence data. To determine whether these P. farinosus isolates can be considered as biocontrol agents (BCAs) of powdery mildews, we studied the interactions between P. farinosus and the following four powdery mildew species: B. graminis f.sp. hordei infecting barley, Oidium neolycopersici infecting tomato, Golovinomyces orontii infecting tobacco and Podosphaera fusca infecting cucumber. The powdery mildew colonies of all these four powdery mildew species were quickly destroyed by P. farinosus in leaf cultures but neither conidial suspensions nor cell-free culture filtrates of P. farinosus isolates could suppress the spread of powdery mildew infections on diseased barley, tomato, tobacco or cucumber plants in the greenhouse. It is concluded that P. farinosus cannot be considered as a promising BCA of powdery mildew infections although it can destroy powdery mildew colonies in detached leaf cultures and can be a menace during the maintenance of such cultures of cereal, apple, cucurbit and tomato powdery mildew isolates.  相似文献   

18.
The lengths of conidiophores in fungal colonies of the melon powdery mildew pathogen Podosphaera xanthii Pollacci KMP-6 N cultured under greenhouse (natural) conditions differed markedly from those cultured in a growth chamber. We hypothesized that light wavelength was responsible for the differences in conidiophore length. In this study, we examined the effects of light-emitting diode (LED) irradiation (purple, blue, green, orange, and red light) and white light on colony development and conidiophore formation in KMP-6 N using a stereomicroscope and a high-fidelity digital microscope. Colonies on leaves were flat under greenhouse conditions and under red LED light irradiation but were stacked under growth chamber conditions and under purple, blue, green, and orange LED light irradiation. In addition, KMP-6 N formed catenated conidia comprising six conidia per conidiophore under greenhouse conditions and red light but more than seven conidia per conidiophore under growth chamber conditions and purple, blue, green, and orange light. Furthermore, almost none of the conidia on top of the conidiophores grown under blue light were fully constricted. Therefore, these fungi could not scatter their conidia and spread infection. This is the first report of the effects of LED lights on conidiophore formation in the melon powdery mildew fungus P. xanthii. The results provide insight into the mechanisms underlying the responses of conidiophores to light of specific wavelengths and conidial scatter from conidiophores of melon powdery mildew fungi.  相似文献   

19.
Identification of the physiological races ofPodosphaera xanthii (syn.Sphaerotheca fuliginea), the causal agent of powdery mildew in cucurbits, is based upon the differing responses of various melon cultigens to the pathogen. Eight races of the pathogen have been identified to date in the USA, Africa, Europe and around the Mediterranean Sea, and four new races were reported from greenhouse melons in the major growing area of Japan. Plant responses to powdery mildew may be affected by environmental factors such as light intensity, temperature and humidity, as well as by age and nutritional status of the plants. The same factors affect the accuracy and reliability of race identification. In an attempt to overcome those obstacles, the genetic diversity ofP. xanthii was studied using molecular markers. Unfortunately, no correlation was found between DNA polymorphism and the race of the pathogen as identified by biological tests. The usefulness of race identification as a guide for the grower in selecting appropriate cultivars is limited because changes or shifts in the pathogen population are common. Such changes may be found among growing seasons, geographic regions and hosts, and also within a single greenhouse during a single season. On the other hand, race identification is important for basic research and is especially important for the commercial seed industry, which requires accuracy in declaring the type and level of resistance to powdery mildew in its products. http://www.phytoparasitica.org posting March 2, 2004. Contribution from the Agricultural Research Organization. No. 501/04.  相似文献   

20.
The activity of different types of natural compounds and of two biofungicides based on Bacillus subtilis and Ampelomyces quisqualis alone and in combination with fungicides was tested against powdery mildew of zucchini. The efficacy was compared to the activity of fungicides used alone in four experimental trials carried out in the open field and under greenhouse conditions. The Podosphaera xanthii population used throughout the work was partially resistant to azoxystrobin, whereas it was susceptible to mychlobutanil. Sulphur plus terpenes and mustard oil consistently controlled powdery mildew, followed by mychlobutanil alone or in combination with A. quisqualis. B. subtilis and A. quisqualis when tested alone were partially effective. The combination of azoxystrobin and B. subtilis only delayed the spread of the pathogen.  相似文献   

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