Accumulation of Pathogenesis-Related Proteins in the Apoplast of a Susceptible Cultivar of Apple (Malus domestica cv. Elstar) after Infection by Venturia inaequalis and Constitutive Expression of PR Genes in the Resistant Cultivar Remo

Leaves of apple (Malus domestica cv. Elstar) were infected with a cloned isolate of the apple scab Venturia inaequalis. The intercellular washing fluid (IWF) of these plants was collected and the variation in the composition of proteins in the IWF was analysed by SDS-PAGE and two-dimensional gel electrophoresis during and after the infection with V. inaequalis, the causal agent of apple scab. The subsequent analysis of induced proteins by electron spray ionization quadrupole time of flight mass spectroscopy revealed the presence of -1,3-glucanase, chitinase, thaumatin-like protein and a cysteine-like protease in M. domestica leaves infected by V. inaequalis. These results were confirmed by immunoblotting with antibodies against some of these proteins. Moreover, a non-specific lipid transfer protein was identified in uninfected leaves: the amount declined to a non-detectable level within the first week after infection by V. inaequalis. The analysis of the IWF of M. domestica cv. Remo, bearing resistances to apple scab, powdery mildew and fire blight, showed a protein pattern comparable to that of the IWF from V. inaequalis infected leaves from cultivar Elstar indicating the constitutive production at least of some of the pathogenesis-related proteins in the resistant cultivar.     

The ubiquity of non-specific eliciting activity in intercellular washing fluids from rust-infected wheat leaves

Intercellular washing fluids (IWF) were harvested from a range of wheat cultivars infected with either the leaf rust fungus ( Puccinia recondite f.sp. tritici ) or the stem rust fungus ( Puccinia graminis f.sp. tritici ). These fluids were then tested for their ability to elicit symptoms of chlorosis and necrosis in 14 wheat cultivars. Eliciting activity was found in all samples tested, irrespective of both the host cultivar infected by the rust fungus and the genotype of the infecting pathogen isolate. Healthy test cultivars either responded to all IWF preparations or were responsive to none of them. Investigations into the nature of this non-specific eliciting activity revealed that active molecules are periodate sensitive, negatively charged at neutral pH, and associated with a high molecular weight fraction.     

Race-specific elicitors from the Peronospora parasitica / Arabidopsis thaliana pathosystem

Intercellular washing fluids (IWFs) were isolated from compatible interactions ofPeronospora parasitica with Arabidopsis. IWFs were filter-sterilized then infiltrated into leaves of resistant and susceptible Arabidopsis genotypes. P. parasitica isolates WELA (virulent on Landsberg erecta (La-er) and Weiningen (Wei-0) but avirulent on Columbia (Col-0) and RLD) and NOCO (virulent on Col-0 and RLD but avirulent on La-er) were used. A “reciprocal check” pattern of necrosis-induction was observed in which IWFs from a compatible interaction caused necrosis in leaves of the Arabidopsis genotypes resistant to the “parent” fungal isolate but not in those genotypes susceptible to it. Analysis of variance showed that the interaction between fungal isolate and necrosis in host genotypes was highly significant. The necrosis-inducing activity was significantly correlated with a high degree of asexual sporulation in the compatible interaction from which the IWF was isolated. However, the necrosis-inducing activity was not correlated with the protein content of the IWF.     

Mitochondrial haplotype analysis as a tool for differentiating isolates of Verticillium dahliae

The ability to monitor mitochondrial background in Verticillium dahliae may provide an additional tool for population studies and monitoring clonal populations. Published mitochondrial genome sequences of V. dahliae (DQ351941) were used to design primers for amplification of spacer regions for assessment of mitochondrial haplotype differences among isolates. Five regions were examined (5,229 bp, or 19% of the total genome size) for 30 isolates representing a range in vegetative compatibility group (VCG), host, and geographic origin. Observed differences among isolates were due to single nucleotide polymorphisms, different numbers of bases in specific homopolymeric regions, and copies of subrepeated sequences. When the differences observed for each locus were totaled there were 28 total groupings; when the results of each locus for individual isolates were combined there were 15 mitochondrial haplotypes. Some of the observed groupings correlated with VCG. For example, five VCG-1A and VCG-1B isolates from California, Spain, and Greece had identical haplotypes; however, this was not observed for VCG-2 or VCG-4 isolates. While some VCG-2 isolates also were identical and fell into a single haplotype, five haplotypes were found for this group (five other haplotypes were observed for other isolates that had not been characterized to VCG but grouped with VCG-2 isolates in the phylogenetic analysis). Likewise, five VCG-4 isolates fell into four mitochondrial haplotypes, one of which was identical to the largest VCG-2 grouping. A heterokaryon self-incompatible isolate that was reported in the literature to cluster with VCG-2 isolates by amplified fragment length polymorphism analysis was identical with VCG-1 isolates for four of the five loci, but was intermediate between VCG-1 and VCG-2 in the haplotype analysis. Phylogenetic analysis with these regions revealed the mitochondrial background of VCG-1 and VCG2-B to be monophyletic but VCG-2A and VCG-4 could not be separated. The results obtained indicate that there is variation in mitochondrial haplotypes and this type of analysis may be a useful for characterization of isolates. While data from all five regions was used for the haplotype separation in this study, depending on the VCG or the level of variability observed within a population it is possible to use fewer loci.     

Components of partial resistance to Puccinia striiformis in barley

Substantial differences in the rates of mycelial growth and sporulation of different isolates of Puccinia striiformis were observed when seedling and upper leaves of five barley cultivars with susceptible reaction types were inoculated. The highest rate of mycelial growth was usually on cv. Sultan, followed by Proctor, Astrix, Senta and Zephyr in that order, but more spores were produced on Senta and Astrix than on Proctor, Zephyr or seedlings of Sultan. Host-specific differences between isolates were slight, but some isolates were more aggressive than others, with generally higher mycelial growth and sporulation rates and shorter latent periods. The effect of temperature on sporulation varied little between cultivars or isolates, but there was some evidence of high temperature resistance in Senta and Zephyr. Latent periods were longer on flag leaves than on seedling leaves, and spore production per unit area of leaf infected was lower on the flag leaf of all cultivars except Sultan. Low sporulation rates seemed due to adult plant resistance in Astrix and Senta, and overall resistance, more strongly expressed in adult plants than in seedlings, in Proctor and Zephyr. Sultan exhibited slight seedling resistance, but lacked the adult plant resistance of other cultivars.     

Relative Contribution of Seed-Transmitted Inoculum to Foliar Populations of Phaeosphaeria nodorum

ABSTRACT A marked-isolate, release-recapture experiment was conducted to assess the relative contributions of seed-transmitted (released isolates) versus all other inocula to foliar and grain populations of Phaeosphaeria nodorum in winter wheat rotated with nonsusceptible crops in New York and Georgia, United States. Seed infected with two distinct groups of marked isolates of P. nodorum containing rare alleles (identified by amplified fragment length polymorphisms [AFLPs]) and balanced for mating type were planted in experimental field plots in two locations in each state. Recapture was done by isolating P. nodorum from leaves showing necrotic lesions at spring tillering and flowering stages, and mature grains from spikes showing glume blotch. Isolates from these samples were genotyped by AFLPs and categorized as released or nonreleased to infer sources of inoculum. Both infected seed and other sources of the pathogen contributed significant primary inocula to populations recovered from leaves and harvested grain. Seed-transmitted genotypes accounted for a total of 57% of all isolates recovered from inoculated plots, with a range of 15 to 90% of the populations of P. nodorum collected over the season in individual, inoculated plots at the four locations. Plants in the noninoculated control plots also became diseased and 95% or more of the isolates recovered from these plots were nonreleased genotypes. Although other potential sources of P. nodorum within and adjacent to experimental plots were not ruled out, nonreleased genotypes likely were derived from immigrant ascospores potentially from sources at a considerable distance from the plots. Our results suggest that, although reduction of seedborne inoculum of P. nodorum may delay foliar epidemics, this strategy by itself is unlikely to result in high levels of control in eastern North America because of the additional contribution from alternative sources of inoculum.     

苹果叶片应答斑点落叶病菌胁迫的蛋白质组学分析

In order to elucidate key proteins related to disease-resistance in apple leaves,the total proteins were extracted from control and infected apple leaves, and separated by two-dimensional electrophoresis. The differential expressed proteins were then identified by MALDI-TOF-TOF/MS.In total, 25 differential expressed proteins were detected by Image Master Software. After tryptic digestion, MALDI-TOF-TOF/MS analysis and database searching, 20 protein spots were finally identified, including 11 functional proteins related to photosynthesis, energy metabolism, stress and defense responses. The pathogenesis-related proteins involved in defense responses such as APX, GPX and Mal d1 were differentially expressed in apple leaves. It indicates that these proteins may play a key role in the resistance to A. alternata apple pathotype.     

Biological, serological, and molecular differences among isolates of potato a potyvirus

ABSTRACT Sequences of the coat protein (CP) and 3'-end nontranslated region (3'NTR) of 13 isolates and the helper component proteinase (HC) of nine isolates of potato A potyvirus (PVA) were determined and compared with the eight previously determined PVA CP and 3'NTR sequences and one HC sequence. CP amino acid (aa), 3'NTR nucleotide, and HC aa sequence identities were 92.9, 93.4, and 94.8%, respectively. Sequence data, serological tests, and the necrotic local lesions induced in the leaves of the potato hybrid 'A6' confirmed that tamarillo mosaic virus is a strain of PVA. The aa substitutions A6T and G7S in the CP N-terminus were correlated with loss of aphid transmissibility. Development of necrotic lesions or nonnecrotic symptoms in the systemically infected leaves or lack of systemic spread in potato cv. King Edward were used to place the PVA isolates into four strain groups, but this grouping was not correlated with any differences in CP, HC, or 3'NTR. Recognition of CP by three monoclonal antibodies was used to place the PVA isolates into three groups different from the four groups above. The epitopes of two mono-clonal antibodies were mapped by site-directed mutagenesis to the same lysine residue at the CP aa 34.     

Sequences of the internal transcribed spacers of the ribosomal RNA genes and relationships between isolates of Verticillium alboatrum and V. dahliae

The sequences of the internal transcribed spacers (ITS) 1 and 2 of the rRNA genes of 38 Verticillium alboatrum and V. dahliae isolates have been determined. The isolates represented RFLP groups of both species, V. dahliae vegetative compatibility groups and pathotypes, and V. alboatrum 'group 2'(the majority of the V. alboatrum isolates studied were in'group 1'). The ITS sequences of a single V. tricorpus isolate were also determined. RFLP groups L and NL of V. atboatrum were distinct, with a maximum of three nucleotide differences between any isolate. Most haploid V. dahliae isolates were identical and separated from most L/NL V. alboatrum isolates by differences at five or six positions. A few haploid V. dahtiae isolates, not forming any obvious grouping, differed from the majority, each at a single position. Diploid isolates of V. dahtiae were identical but only one nucleotide difference separated them from some RFLP group L V. alboatrum isolates. At least six differences separated the diploid isolates of V. dahliae from the haploid ones. The'group 2' V. atboatrum isolates were more distantly related to'group 1'isolates (at least 17 positions different) than was V. tricorpus. The possibility of defining specific primers for use in PCR to discriminate species and subspecific groups is discussed.     

Cellulase in the Host–parasite System Phaseolus vulgaris (L.)–Uromyces appendiculatus [Pers.] Link

Activity of cellulase and xylanase in the intercellular washing fluid (IWF) of bean plants (Phaseolus vulgaris, cultivar Fori) was monitored during infection with bean rust (Uromyces appendiculatus). In infected plants, cellulase activity could be detected at 2 days after inoculation and reached its maximum between 7 and 8 days after inoculation. The enzyme activity was not detected in healthy controls. The cellulase had a pH optimum at pH 5.5 and a temperature optimum at 30°C. Complete inactivation of cellulase occurred after heating to 50°C for 30min. In non-denaturing polyacrylamide gradient gels, the enzyme exhibited four bands (molecular masses approximately 70, 95, 120, 170kDa). After isoelectric focusing, eight cellulase isoforms with pI values pI 4.6–4.8; 5–5.1; 5.4; 5.5; 5.9; 6; 6.5; 7 appeared. Two dimensional electrophoresis yielded 13 cellulase isoforms. Unlike cellulase, low levels of xylanase were detected in healthy controls. The activity of this hydrolase did not increase due to rust infection.     

Evidence that a Leaf-Disk Test Allows Assessment of Isolate-Specific Resistance in Brassica oleracea Crops Against Downy Mildew (Peronospora parasitica)

A rapid resistance/susceptibility test for Peronospora parasitica (downy mildew) was established by inoculating leaf-disks of four Brassica oleracea accessions. Several conditions were tested: disk disinfection or not, agar medium with or without nutrients and with 50 or 100 ppm of benzimidazole. Using disinfected disks placed on agar (no nutrient and benzimidazole at 50 or 100 ppm), the responses of leaf-disks to four isolates were similar to those obtained using the classical cotyledon test, whereas undesired contaminations occurred in all other conditions. The possible effect of the particular leaf used for obtaining the disks was also studied. In each incompatible interaction tested, disks were resistant whatever the leaf used. In compatible interactions, susceptible phenotypes were observed on disks derived from the six lowest leaves, but disks from upper leaves were resistant. The genetic basis of resistance in a F1 hybrid broccoli was assessed, by testing six isolates on an F2 population derived from this hybrid. The cotyledon test only allows inoculation of two isolates per seedling, whereas many isolates can be tested on each plant by using leaf-disks. The segregation of the resistance to each of the six isolates was analysed: two dominant genes (tightly linked) control resistance to all isolates (one to five isolates; the other to only one isolate).     

黄瓜绿斑驳花叶病毒基因组全序列分析及病害的田间调查

为明确近年来在浙江省葫芦科作物上发生的黄瓜绿斑驳花叶病毒(Cucumber green mottle mosaic virus,CGMMV)基因组特征及其发生分布情况,从浙江省及上海地区的甜瓜、西瓜和瓠瓜上采集疑似样品进行RT-PCR鉴定,通过分段扩增测序的方法拼接获得基因组全序列并进行系统进化分析,利用特异性引物扩增获得CGMMV外壳蛋白(coat protein,CP)基因序列,制备CGMMV CP抗血清进行Western-blot和Dot-ELISA检测。结果显示,来自甜瓜、西瓜和瓠瓜的3个CGMMV分离物基因组全序列均具有烟草花叶病毒属典型基因组结构特征,全部由6 423 nt构成;3个全序列间的核苷酸同源性高达99.11%~99.67%,编码的CP氨基酸同源性为100%。系统进化分析发现,CGMMV不同分离物形成2个进化相关群体,3个浙江的CGMMV分离物均位于第I组内,与已报道的中国CGMMV分离物和韩国CGMMV分离物亲缘性较高。Western-blot检测表明CGMMV CP抗血清可以与感病植株中的病毒发生特异性反应,可用于CGMMV鉴定;Dot-ELISA检测发现CGMMV在浙江省和上海市的葫芦科作物上普遍存在。     

甜瓜种子携带的镰刀菌种类鉴定及其致病性研究

 采用洗涤检验法和马铃薯葡萄糖琼脂培养基法,从甜瓜种子上分离得到23个镰刀菌分离物。采用形态观察和分子生物学方法对其进行鉴定,并研究其致病性以及对甜瓜种子发芽的影响。通过观察菌落形态和色素颜色,以及大型分生孢子、小型分生孢子和厚垣孢子形态,同时分析ITS-rDNA序列和翻译延伸因子-1α(TEF-1α)基因序列,以确定真菌分离物的分类属性。结果表明,4个分离物为尖孢镰刀菌(Fusarium oxysporum),5个分离物为层生镰刀菌(F. proliferatum),5个分离物为木贼镰刀菌(F. equiseti),9个分离物为轮枝样镰刀菌(F. verticillioides)。这4种镰刀菌均对甜瓜幼苗有致病性,其分生孢子悬浮液对甜瓜种子发芽有抑制作用。这是我国首例开展甜瓜种子携带致病性镰刀菌的研究报道。     

甜瓜种子携带的镰刀菌种类鉴定及其致病性研究

 采用洗涤检验法和马铃薯葡萄糖琼脂培养基法,从甜瓜种子上分离得到23个镰刀菌分离物。采用形态观察和分子生物学方法对其进行鉴定,并研究其致病性以及对甜瓜种子发芽的影响。通过观察菌落形态和色素颜色,以及大型分生孢子、小型分生孢子和厚垣孢子形态,同时分析ITS-rDNA序列和翻译延伸因子-1α(TEF-1α)基因序列,以确定真菌分离物的分类属性。结果表明,4个分离物为尖孢镰刀菌(Fusarium oxysporum),5个分离物为层生镰刀菌(F. proliferatum),5个分离物为木贼镰刀菌(F. equiseti),9个分离物为轮枝样镰刀菌(F. verticillioides)。这4种镰刀菌均对甜瓜幼苗有致病性,其分生孢子悬浮液对甜瓜种子发芽有抑制作用。这是我国首例开展甜瓜种子携带致病性镰刀菌的研究报道。     

Genetic differentiation in the French population of Erysiphe cichoracearum, a causal agent of powdery mildew of cucurbits

The relative incidence of Erysiphe cichoracearum and Sphaerotheca fuliginea , both agents of powdery mildew of cucurbits, was determined from 275 samples of mildewed leaves of cucurbits collected in 1994 from five regions of France. E. cichoracearum was identified in 9 to 39% of the mildewed leaf samples from four of the regions but was not detected in samples from the Mediterranean island of Corsica. The genetic structure of the French population of E. cichoracearum was examined using RFLPs of the ribosomal internal transcribed spacers amplified by PCR, random amplified polymorphic DNA (RAPD) markers, pathogenicity and mating-type tests. Forty-one isolates, including one from England, were analysed. Cluster analysis from 147 RAPD fragments using 16 primers revealed the existence of three distinct genetic lineages corresponding to three rDNA haplotypes (designated groups A, B and C). Bootstrap, genetic diversity, gametic disequilibrium and private allele analyses supported this differentiation. The genetic differentiation observed in the French population was not related to the geographical origin of the isolates. Group A isolates may be more specialized on melon as, with one exception, they were of race 1 (growth on four of the five melon cultivars tested) in comparison with group B and C isolates, which were of race 0 (growth on IranH only). Thus, the genetic differentiation observed may indicate a host-specialized subdivision within the French population of E. cichoracearum from cucurbits. Gametic disequilibrium analysis among RAPD loci and biological observations suggest that the sexual stage is of minor importance for epidemics of E. cichoracearum on cucurbits.     

Variation in Occurrence of Dematiaceous Hyphomycetes on Forage Bermudagrass over Years, Sampling Times, and Locations

ABSTRACT Leaf samples of forage bermudagrass with symptoms of infection by species of Bipolaris, Curvularia, and Exserohilum (dematiaceous hyphomycetes) were collected from three swine waste application sites in Mississippi at eight sampling times during each of 3 years. Samples were assayed for pathogens by observing sporulation on plated leaf tissue. Among 3,600 leaves assayed, eight species of the three genera were observed. Features and criteria for the practical identification of species on plated leaf samples are described. Sporulation by dematiaceous hyphomycetes was observed on 97% of leaves; a single pathogen was observed on 20% and two to five pathogens were observed on 77% of leaves. Distributions of leaves among classes with one to five pathogens per leaf, for sites within years, always differed significantly (P = 0.01) from a Poisson distribution and usually included fewer leaves than expected with four or five pathogens. Significant (P = 0.05) variation in frequencies of occurrence of pathogens among 72 samples of 50 leaves each was attributed to pathogen species, sampling times, and species-time interactions. Exserohilum rostratum, Curvularia lunata, and Bipolaris cynodontis were the most frequent pathogens across years and sites; B. spicifera and C. geniculata were intermediate; and B. hawaiiensis, B. sorokiniana, and B. stenospila were least frequent. For the five most common pathogens, significant differences in frequency among locations were commonplace. Six pathogens exhibited significant (P = 0.05) positive and negative correlations with others in overall frequencies of occurrence across years, sampling times, and sites. However, chi(2) tests of dual occurrence indicated that interactions between specific pairs of pathogens in or on leaves are not likely to be major causes for overall frequency correlations. Results indicate that dematiaceous hyphomycetes typically infect forage bermudagrass on swine waste application sites in complexes rather than as individual species; that E. rostratum, C. lunata, and B. cynodontis are the predominant pathogens; and that frequencies of pathogens often differ significantly between locations.     

Impact of tobacco recessive resistance gene va on biological properties of Brazilian Potato virus Y (PVY) isolates

To investigate the role of environmental conditions on the selection of virulent Potato virus Y (PVY) isolates subject to pressure from the recessive resistance gene va in tobacco, a field survey was performed in Brazil where va‐derived genotypes have been recently introduced and now represent less than one‐third of cultivated tobacco genotypes. A serological analysis of 397 leaves collected from different Brazilian tobacco‐growing areas and mainly from plants with symptoms indicated that 52·4% of samples were infected by at least one of the viral species tested. PVY was present in 72·1% of infected samples. The probability of a plant being infected with PVY was reduced in va hosts. However, the biological characterization of PVY isolates on indicator hosts showed that 20 of the 29 tested isolates were able to overcome the alleles of the va gene. Moreover, the observed biological diversity of isolates was higher in susceptible tobacco genotypes than in va‐resistant ones. Comparison of these data with the PVY diversity in French tobacco fields shows that the use of va‐derived genotypes in two environments with contrasting climatic conditions, local hosts and cultural contexts, leads to a similar outcome: the prevalence of virulent isolates. These results strongly suggest an important role of the va gene in the modification of PVY populations.     

黄龙病菌在柑橘枝条上的分布和多样性分析

 柑橘黄龙病是由“Candidatus Liberibacter asiaticus”(CLas)引起的毁灭性病害,在染病植株的叶片和果实上表现出不同症状。但目前还未见在同一植株或枝条上表现不同症状的叶片或果实中的CLas是否存在多样性的报道。本研究分析来自美国佛罗里达与中国广东两地染病柑橘样品,发现佛罗里达的CLas遗传多样性较低。以3对位于CLas短串联重复位点两端的序列为引物,对广东柑橘黄龙病病株的样品进行PCR扩增和PAGE检测,结果发现:同一植株上的CLas多样性高于同一枝条;同一病枝上显不同症状的叶片和果实中的CLas浓度分布不均匀,但成熟病果中CLas的浓度最高。以同一病枝条为一个组,PCR扩增和PAGE检测53个枝条的样品,发现不同组间CLas多样性不一:3对引物扩增条件下分别有4组(7.55%)、7组(13.21%)和11组(20.75%)样品存在组间多样性,推测同一枝条中含有不同的CLas菌株。以上结果说明了CLas在柑橘枝条上的分布规律,为监测我国CLas的多样性提供了科学依据。     

苹果茎沟病毒柑橘碎叶株系的分布与序列分析

为明确苹果茎沟病毒(apple stem grooving virus,ASGV)柑橘碎叶株系(citrus tatter leaf virus,CTLV)在中国柑橘产区的发生分布及其分子特性,于2017—2021年对我国12个柑橘主产区开展系统调查,并采用多重比对以及系统发育分析法对CTLV的全序列进行分析。结果表明,从12个柑橘主产区采集的2 012份疑似样品中检测出413份感CTLV阳性样品。除陕西省外,其余11个柑橘主产区样品中均检测出CTLV,并且柚类样品中CTLV的检出率最高,达到32.31%。对分离获得的22株CTLV毒株和已知的7株CTLV以及5株ASGV毒株进行全序列分析,发现所有34株毒株的核苷酸序列相似性为78.6%～99.8%,且CTLV序列的保守性较高。此外,与其他4株ASGV毒株相比,本研究中22株CTLV毒株与ASGV-MK毒株(GenBank登录号为MZ127820.1)具有更近的亲缘关系。推测CTLV中国毒株间的亲缘关系可能与其采样地和寄主品种有关。     

福建西番莲病毒病的发生及其病原黄瓜花叶病毒亚组鉴定

1988～1995年对福建西番莲病毒病的发生为害进行调查,发现在西番莲栽培区病毒病普遍发生,发病率通常在30％～40％,严重的达90％以上。田间症状主要表现为叶片环斑、皱缩、花叶、环斑花叶,死顶和果实木质化等。从田间病株采集40份样本,经电镜观察和采用A蛋白夹心ELISA(PAS-ELISA),用西番莲木质化病毒(PWV)、西番莲黄花叶病毒(PFYMV)、紫果西番莲花叶病毒(GMV)、烟草花叶病毒(TMV)及黄瓜花叶病毒(CMV)的抗血清进行测定,其中36个样本检测出CMV,表明福建西番莲病毒病的主要病原为CMV。与此同时,对西番莲上CMV进行亚组鉴定,采用鉴别寄主和单、多克隆抗体双夹心ELISA(DAS-ELISA)对36个CMV阳性样本进行测定,结果35个属CMV亚组Ⅰ,1个属CMV亚组Ⅱ,表明田间以CMV亚组Ⅰ分离物占绝对优势。