Multiphasic analysis of xanthomonads causing bacterial spot disease on tomato and pepper in the Caribbean and central america: evidence for common lineages within and between countries

ABSTRACT Four hundred thirty-three xanthomonad strains isolated from tomato or pepper plants from 32 different fields in four Caribbean and Central American countries were screened for the ability to hydrolyze starch and sodium polypectate and for resistance to copper and streptomycin. Of these, 95 representative strains were further characterized by various phnetic tests, and 63 of these strains were then analyzed by genomic fingerprinting. Most of the strains (>90%) were tolerant to copper. However, there was much more variability in sensitivity to streptomycin. All strains in Guadeloupe and 93% of the strains in Barbados were sensitive to streptomycin. The majority of strains were typical Xanthomonas campestris pv. vesicatoria group A strains. In Barbados, however, a unique group of strains was identified that was serologically similar to group A strains but was amylolytic. These strains were designated A1. The occurrence of X. campestris pv. vesicatoria group B strains in Central America was found to be limited to two fields in Costa Rica and one in Guatemala. No group B strains were identified in the Caribbean, in contrast to common occurrence in the central United States and in South America. T3 strains were not found in this study, despite the recent increase of such strains in Florida and Mexico. Unique strains from Costa Rica belonging to the X. gardneri group were identified. Little linkage was found among phenotypic and rep-polymerase chain reaction (rep-PCR) genomic fingerprinting profiles of the pathogens except at the species/pathovar level; strains displaying virtually identical fingerprint profiles were found to correspond to distinct races and vice versa. The rep-PCR genomic fingerprinting analyses suggest that certain lineages may have evolved or predominated in specific regions or specific countries.     

辣椒、番茄细菌性疮痂病及生理小种鉴定

 近3年,从北京、山西、内蒙、新疆和云南等地的辣椒和番茄病株上分离到19个菌株,经致病性测定和细菌学鉴定,确定这19个菌株为甘蓝黑腐黄单胞菌疮痂致病变种(Xanthomonas campestris pv.vesicatoria(Doidge) Dye,1978)。供试19个菌株在国内首次采用国际标准鉴别寄主进行了生理小种鉴定。其中,3个菌株为番茄小种1(XcvT race1),仅存在于北京地区,其它16个菌株均属于辣椒-番茄小种3(XcvPT race3),分布广,为我国优势小种。     

Development of bacterial spot on near-isogenic lines of bell pepper carrying gene pyramids composed of defeated major resistance genes

ABSTRACT Disease severity caused by races 1 through 6 of Xanthomonas campestris pv. vesicatoria on eight near-isogenic lines (isolines) of Early Calwonder (ECW) with three major resistance genes (Bs1, Bs2, and Bs3) in different combinations was evaluated in the greenhouse and field. Strains representing races 1, 3, 4, and 6 caused similar high levels of disease severity, followed by races 2 and 5 on susceptible ECW. Race 3 caused severe disease on all isolines lacking resistance gene Bs2. Race 4, which defeats Bs1 and Bs2, caused less disease on isoline ECW-12R (carries Bs1 + Bs2), than on isolines ECW, ECW-10R (carries Bs1), and ECW-20R (carries Bs2). Similar results were obtained with race 4 strains in field studies conducted during 1997 and 1998. In greenhouse studies, race 6, which defeats all three major genes, caused less disease on isoline ECW-13R (carries Bs1 + Bs3) and ECW-123R (carries Bs1 + Bs2 + Bs3) than on isolines ECW, ECW-10R, ECW-20R, and ECW-30R (carries Bs3), but not on ECW-23R (carries Bs2 + Bs3). In greenhouse studies with commercial hybrids, strains of races 4 and 6 caused less disease on Boynton Bell (carries Bs1 + Bs2) than on Camelot (carries no known resistance genes), King Arthur (carries Bs1), and X3R Camelot (carries Bs2). Race 6 caused less disease on hybrid R6015 (carries Bs1 + Bs2 + Bs3) and Sentinel (carries Bs1 + Bs3) than on Camelot. Residual effects were not as evident in field studies with race 6 strains. Defeated major resistance genes deployed in specific gene combinations (i.e., gene pyramids) were associated with less area under the disease progress curve than when genes were deployed individually in isolines of ECW or commercial hybrids. Successful management of bacterial spot of pepper is achieved incrementally by integrating multiple tactics. Although there is evidence of residual effects from defeated genes, these effects alone likely will not provide acceptable bacterial spot control in commercial production fields. However, when combined with sanitation practices and a judicious spray program, pyramids of defeated resistance genes may aid in reducing the risk of major losses due to bacterial spot.     

Bacterial spot of capsicum and tomato in Yugoslavia

The causal agent of bacterial spot of capsicum and tomato grown in different regions in Yugoslavia was investigated. Isolations were made from diseased material collected in recent years. The biochemical and physiological characteristics of isolated bacteria were studied by standard bacteriological tests. The race of the pathogen was determined on differential cultivars of capsicum and tomato. The causal agent of the disease was identified according to the concepts of the time as Xanthomonas campestris pv. vesicatoria. Strains isolated from diseased capsicum were non-pectolytic and non-amylolytic, and did not infect tomato plants. According to the reaction of capsicum cv. Early Calwonder and its isogenic lines, these strains belonged to'pepper races'1 and 3 of X. vesicatoria. Tomato strains showed pectolytic and amylolytic activity and were not pathogenic to capsicum. Accordingly, the capsicum strains could now be considered to be X. axonopodis pv. vesicatoria and the tomato strains X. vesicatoria.     

Detection and identification of seed-borne pathogenic bacteria of imported tomato seeds in Egypt

Imported tomato seed lots of different cultivars were assayed for the presence of seed-borne bacterial pathogens. The liquid assay method was used for detection of the bacteria, and seed extracts were plated on different semi-selective media. Pseudomonas corrugata and Xanthomonas campestris pv. vesicatoria were detected in 14.7% and 12% of the seed samples tested respectively. These pathogens were identified by means of biochemical, physiological and pathogenicity tests as well as the Biolog GN Microplate System for X. campestris pv. vesicatoria. Both P. corrugata and X. campestris pv. vesicatoria were more easily identified on Tween B and CKTM media than on other media. This is the first report of the occurrence of these important pathogens on tomato seeds in Egypt.     

Temperature Sensitivity of the Hypersensitive Response of Bell Pepper to Xanthomonas axonopodis pv. vesicatoria

ABSTRACT When bacterial spot-resistant pepper plants carrying resistance gene Bs2 and infiltrated with incompatible strains of Xanthomonas axonopodis pv. vesicatoria carrying a functional avrBs2 gene (races P1 and P3) were incubated at 32 degrees C, they exhibited an electrolyte leakage and bacterial multiplication pattern in planta similar to that obtained with a compatible strain (race P4) carrying a nonfunctional avrBs2 gene. They also developed disease-like symptoms. Pretreatment of incompatible bacteria at 32 degrees C before infiltration caused a delay in electrolyte leakage less pronounced than that caused by exposing plants to 32 degrees C. Also, plants had to be exposed to 32 degrees C for an hour prior to inoculation to increase symptom expression. These data suggest that the Bs2 gene is temperature sensitive. In other experiments, the avrBs1-Bs1 interaction appeared to be the most heat tolerant and thus the least likely to revert to compatible, whereas the avrBs3-Bs3 interaction had an intermediate sensitivity to elevated temperatures.     

Identification of Isolates that Cause a Leaf Spot Disease of Brassicas as Xanthomonas campestris pv. raphani and Pathogenic and Genetic Comparison with Related Pathovars

ABSTRACT Twenty-five Xanthomonas isolates, including some isolates received as either X. campestris pv. armoraciae or pv. raphani, caused discrete leaf spot symptoms when spray-inoculated onto at least one Brassica oleracea cultivar. Twelve of these isolates and four other Xanthomonas isolates were spray- and pin-inoculated onto 21 different plant species/cultivars including horseradish (Armoracia rusticana), radish (Raphanus sativus), and tomato (Lycopersicon esculentum). The remaining 13 leaf spot isolates were spray-inoculated onto a subset of 10 plant species/cultivars. The leaf spot isolates were very aggressive on several Brassica spp., radish, and tomato causing leaf spots and dark sunken lesions on the middle vein, petiole, and stem. Based on the differential reactions of several Brassica spp. and radish cultivars, the leaf spot isolates were divided into three races, with races 1 and 3 predominating. A differential series was established to determine the race-type of isolates and a gene-for-gene model based on the interaction of two avirulence genes in the pathogen races and two matching resistance genes in the differential hosts is proposed. Repetitive-DNA polymerase chain reaction-based fingerprinting was used to assess the genetic diversity of the leaf spot isolates and isolates of closely related Xanthomonas pathovars. Although there was variability within each race, the leaf spot isolates were clustered separately from the X. campestris pv. campestris isolates. We propose that X. campestris isolates that cause a nonvascular leaf spot disease on Brassica spp. should be identified as pv. raphani and not pv. armoraciae. Race-type strains and a neopathotype strain for X. campestris pv. raphani are proposed.     

Incidence and pathogenicity of races and isolates of Verticillium dahliae in Crete, southern Greece

Classification of 32 Verticillium dahliae isolates originating from 19 plant species in eight different botanical families to races and determination of host range pathogenicity were carried out. The physiological races of isolates were identified using the two differential tomato cultivars ??Belladonna?? (susceptible to both races 1 and 2 of V. dahliae) and ??Ace 55VF?? (resistant to race 1, susceptible to race 2 of V. dahliae). Among these isolates, 14 were race 2 (43.8%), 12 race 1 (37.5%) and six nonpathogenic (18.7%) on tomato. The host range pathogenicity of isolates was determined using four differential hosts (eggplant, turnip, tomato (Ve ^? ) and sweet pepper). Among isolates, five were pathogenic to both eggplant and turnip (15.6%), 21 to eggplant, turnip and tomato (65.6%), five to eggplant, turnip, tomato and sweet pepper (15.6%) and one was pathogenic to eggplant, turnip and sweet pepper (3.2%). The pathogenicity of isolates on the aforementioned five hosts was investigated on the basis of external symptoms and by calculating the relative areas under disease progress curves (relative AUDPC). Results showed that eggplant was the most susceptible, followed by turnip and tomato cv. Belladonna, while sweet pepper and tomato cv. Ace 55VF were less susceptible to all the isolates used. The pathogenicity of isolates varied from highly to mildly virulent on eggplant and turnip while on Belladonna, Ace 55VF and sweet pepper it varied from highly virulent to nonpathogenic. Belladonna exhibited a similar level of susceptibility to races 1 and 2 of V. dahliae, but was more susceptible than Ace 55VF to race 2. Interestingly, the isolates originating from eggplant were clearly more virulent than those originating from tomato and black nightshade on all solanaceous plants tested.     

Mutation in the avrBs1 Avirulence Gene of Xanthomonas campestris pv. vesicatoria Influences Survival of the Bacterium in Soil and Detached Leaf Tissue

ABSTRACT The role of the avrBs1 avirulence gene of Xanthomonas campestris pv. vesicatoria in survival of the bacterium was investigated by testing two strains that differ in the structure and function of the gene in 37 different soil types of Barbados and detached leaf tissue of four pepper genotypes. One strain carried a mutation in the avrBs1 gene and lacked avirulence activity, while the other expressed wild-type avrBs1 activity. In 30 to 32 soil types and all leaf tissue tested, the mutant strain persisted longer and more abundantly than the wild-type strain over a 2- to 6-week period. During this time, the mutant strain generally replaced the wild-type strain completely in soil initially infested with a mixture of equal amounts of each strain and by a factor of 6.7 in similarly infested pepper leaves. Nine selected soil factors, namely pH, clay and cation content, and percent nitrogen, carbonates, carbon, K(+), Na(+), and Ca(2+) did not affect bacterial survival significantly.     

Identification and Origin of Xanthomonas campestris pv. campestris Races and Related Pathovars

ABSTRACT One hundred sixty-four isolates of Xanthomonas campestris pv. campestris and other X. campestris pathovars known to infect cruciferous hosts (X. campestris pvs. aberrans, raphani, armoraciae, and incanae) were inoculated onto a differential series of Brassica spp. to determine both pathogenicity to brassicas and race. Of these, 144 isolates were identified as X. campestris pv. campestris and grouped into six races, with races 1 (62%) and 4 (32%) being predominant. Other races were rare. The remaining 20 isolates from brassicas and other cruciferous hosts were either nonpathogenic or very weakly pathogenic on the differential series and could not be race-typed. Five of these isolates, from the ornamental crucifers wallflower (Cheiranthus cheiri), stock (Matthiola incana) and candytuft (Iberis sp.), showed clear evidence of pathovar-like specificity to the hosts of origin. A gene-for-gene model based on the interaction of four avirulence genes in X. campestris pv. campestris races and four matching resistance genes in the differential hosts is proposed. Knowledge of the race structure and worldwide distribution of races is fundamental to the search for sources of resistance and for the establishment of successful resistance breeding programs.     

Characterization and PCR-based Typing of Xanthomonas campestris pv. vesicatoria from Peppers and Tomatoes in Serbia

During the last two decades bacterial strains associated with necrotic leaf spots of pepper and tomato fruit spots were collected in Serbia. Twenty-eight strains isolated from pepper and six from tomato were characterized. A study of their physiological and pathological characteristics, and fatty acid composition analysis revealed that all of the strains belong to Xanthomonas campestris pv. vesicatoria. Being non-amylolytic and non-pectolytic, pathogenic on pepper but not on tomato, containing lower amounts of fatty acid 15 : 0 ante–iso, the pepper strains were designated as members of the A group of X. campestris pv. vesicatoria. However, the tomato strains hydrolyzed starch and pectate, caused compatible reactions on tomato but not on pepper, had higher percent of 15 : 0 ante–iso fatty acid, and were classified into B phenotypic group and identified as X. vesicatoria. PCR primers were developed which amplified conserved DNA regions related to the hrp genes of different strains of X. campestris pv. vesicatoria associated with pepper and tomato. Restriction analysis of the PCR product resulted in different patterns and enabled grouping of the strains into four groups. When xanthomonads isolated from pepper and tomato in Serbia were analyzed, they clustered into two groups corresponding to the grouping based on their physiological and pathological characteristics. According to the reaction of pepper and tomato differential varieties, the strains from pepper belong to races P7 and P8 and tomato strains belong to the race T2. All strains were sensitive to copper and streptomycin. Advantages and disadvantages of various bacterial spot management practices are discussed.     

Evidence for the Preemptive Nature of Tomato Race 3 of Xanthomonas campestris pv. vesicatoria in Florida

ABSTRACT Until recently, tomato race 1 (T1) of Xanthomonas campestris pv. vesicatoria was the only race causing bacterial spot of tomato in Florida. In 1991, tomato race 3 (T3) was first identified in 3 of 13 tomato production fields surveyed. By 1994, T3 was observed in 21 of 28 fields and was the only race identified in 14 fields. In field studies, tomato genotypes with resistance to either T1 or T3 or susceptibility to both were co-inoculated with strains of both races. Lesions on 10 plants in each of three replications for each genotype were sampled three times during the experiment; bacterial isolations were made from each lesion, and tomato race identifications were made for each strain. At the third sampling date, T3 was isolated from 97% of the lesions on the susceptible genotype Walter and the T1-resistant genotype Hawaii 7998, while T3 was isolated from 23% of the lesions and T1 from the remaining 77% on the T3-resistant genotypes PI 128216 and PI 126932. In surface population studies done in growth rooms, suspensions of T1 and T3 were applied alone and in combination to the leaf surfaces of susceptible and resistant genotypes. T1 populations were reduced more than 10-fold when applied in combination with T3, compared with populations that developed when T1 was applied alone. T3 populations were not affected when applied in combination with a T1 strain. In greenhouse studies with the T3-resistant genotype Hawaii 7981, disease was significantly reduced in plants inoculated with T3 in combination with T1, compared with plants inoculated with T1 alone. These results clearly demonstrate the competitive nature of T3 in the presence of T1 and help explain the emergence of T3 as a prevalent race in Florida.     

Bacteriocin-Like Substances from Tomato Race 3 Strains of Xanthomonas campestris pv. vesicatoria

ABSTRACT Tomato race 3 (T3) strains of Xanthomonas campestris pv. vesicatoria are antagonistic in vitro to tomato race 1 (T1) strains of the bacterium. All T1 strains and 11 strains of other X. campestris pathovars tested were inhibited by T3 strains. Sensitivity of tomato race 2 (T2) strains was variable. No strains from other bacterial genera tested were inhibited. Cell-free filtrates from T3 strains were inhibitory to sensitive strains. The inhibitory activity of these filtrates was lost after treatment at temperatures above 80 degrees C and with selected protease enzymes. However, treatment with trypsin or DNase had no effect on their activity. Seven cosmid clones from a genomic library of a T3 strain were selected for their ability to consistently inhibit a sensitive indicator strain in plate assays. Southern hybridization analysis placed these into three bacteriocin (BCN)-producing groups designated BCN-A, BCN-B, and BCN-C. The BCN-like groups could be differentiated by variations in inhibitory spectra and levels of activity in plate assays. Mutations that inactivated expression of each BCN group individually in a wild-type T3 strain had inhibitory activity confirming that multiple BCNs are present in the T3 strain. T3 strains were inhibitory to a sensitive indicator strain in tomato leaf tissue, but this effect was observed only when T3 strains were applied in advance of the sensitive strain. BCN-A was the major BCN-like substance involved in the suppression of the sensitive indicator strain in tomato leaf tissues.     

Inheritance of Race-Specific Resistance to Xanthomonas campestris pv. campestris in Brassica Genomes

ABSTRACT The inheritance of resistance to three Xanthomonas campestris pv. campestris races was studied in crosses between resistant and susceptible lines of Brassica oleracea (C genome), B. carinata (BC genome), and B. napus (AC genome). Resistance to race 3 in the B. oleracea doubled haploid line BOH 85c and in PI 436606 was controlled by a single dominant locus (Xca3). Resistance to races 1 and 3 in the B. oleracea line Badger Inbred-16 was quantitative and recessive. Strong resistance to races 1 and 4 was controlled by a single dominant locus (Xca1) in the B. carinata line PI 199947. This resistance probably originates from the B genome. Resistance to race 4 in three B. napus lines, cv. Cobra, the rapid cycling line CrGC5, and the doubled haploid line N-o-1, was controlled by a single dominant locus (Xca4). A set of doubled haploid lines, selected from a population used previously to develop a restriction fragment length polymorphism map, was used to map this locus. Xca4 was positioned on linkage group N5 of the B. napus A genome, indicating that this resistance originated from B. rapa. Xca4 is the first major locus to be mapped that controls race-specific resistance to X. campestris pv. campestris in Brassica spp.     

Pathogenicity and RAPD analysis of Phytophthora nicotianae pathogenic to pepper in Tunisia

Nine isolates of Phtophthora nicotianae were isolated from infected pepper plants. Their pathogenicity was studied in Capsicum annuum in comparison with P. nicotianae isolates from tomato and tobacco. The pathogenicity test showed that pepper isolates of P. nicotianae are adapted to their host. Banding patterns obtained by RAPD analysis with six oligonucleotide primers revealed polymorphism that grouped the isolates independently of the plant host. The polygenic dendrogram showed that pepper isolates were more similar to tomato isolates than to tobacco isolates. The RAPD bands of 1300 and 1500 bp, detected with primers OPD-01 and OPD-10, respectively, appeared specific to the most pathogenic pepper isolates. The OPK-08-1950 seems specific to the isolates of P. nicotianae from tomato. These results suggest that host specified might occur in P. nicotianae and that may be due to interspecific hybridization events resulting in novel pathogenic behavior.     

Pathogenicity of plant and soil isolates of <Emphasis Type="Italic">Phytophthora parasitica</Emphasis> on tomato and pepper

Crown and root rot of tomato and sweet pepper can be caused by Phytophthora parasitica. In this work, 23 P. parasitica isolates from diseased pepper or tomato plants as well as 54 isolates from 23 monocrop tomato soils (from Spain and Chile) and one from a pepper soil were studied for their host–pathogen response. Results show significant host specificity for the isolates from tomato plants and tomato soils (63 of 64 isolates were unable to cause disease in pepper). None of the pepper plant/soil isolates showed pathogenicity on tomato, and only four of 14 reproduced their pathogenicity on pepper. Only one tomato isolate was pathogenic to both Solanaceae species. Two different inoculation protocols were evaluated (substrate irrigation and stem cutting). All isolates which expressed pathogenicity when stem inoculated also did it when root inoculated, but not vice-versa. Therefore, the recommended test protocol for tomato and pepper breeding programmes is that based on root inoculation by irrigation.     

新疆加工型辣椒细菌性斑点病的发生和病原鉴定

 新疆加工型辣椒主要产区巴音郭楞蒙古自治州发生了一种严重危害辣椒的细菌性病害。从发病辣椒叶片中分离细菌,通过烟草过敏性反应、马铃薯软腐试验和接种辣椒等致病性测定,确定了13个致病菌株,各菌株之间致病力无明显差异。通过菌体形态、培养性状观察、生理生化反应、寄主范围测定,结合16S rDNA和rpoD基因扩增、序列测定和系统发育分析,将病原菌鉴定为丁香假单胞菌丁香致病变种(Pseudomonas syringae pv. syringae)。病原菌人工接种还能侵染番茄、茄子、马铃薯及黄瓜、四季豆、白菜、萝卜、芹菜等植物。P. syringae pv. syringae引起加工型辣椒细菌性斑点病在国内属首次报道。     

福建省辣椒疫霉菌群体结构表型特征分析

为了明确福建省辣椒疫霉菌群体遗传结构的表型特征,对分离自福建省15个市(县)的300株辣椒疫霉菌的交配型、甲霜灵敏感性及生理小种进行了测定。结果显示,288株为A2交配型,12株为A1交配型,出现频率分别为96%和4%,以A2占优势,在各采集点均有分布。263株对甲霜灵表现敏感,28株为中间型,9株为抗性,分别占总体的87.67%、9.33%和3.00%,甲霜灵敏感菌株为主要菌系。根据菌株对辣椒疫霉菌生理小种鉴别寄主的致病性测定结果,可将辣椒疫霉菌划分为3个生理小种,其中168株为小种3,126株为小种2,6株为小种1,分别占56%、42%和2%,小种3为福建辣椒疫霉菌优势小种。     

Studies of infection withxanthomonas campestris pv.vesicatoria,causal agent of bacterial scab of pepper in Israel

Xanthomonas campestris pv. vesicatoria, the causal agent of bacterial scab of pepper, was isolated in several regions in Israel. When artificial inoculation was practiced, pathogen growth was enhanced by high temperatures (30-36°C), and an inoculum concentration of 10^p6 colony-forming units (CFU) per ml was optimal for symptoms to develop on plants. Pre-inoculation treatments such as wounding the leaves by rubbing them with carborundum powder or spraying them with diluted wax solvents, markedly increased disease severity, but were not essential. Pre-inoculation conditioning at two different relative humidity levels (R.H. = 100% or R.H. < 40%) did not affect disease severity. Young leaves were more severely affected following infection than older leaves. Disease severity was similar with several isolates ofX. campestris pv. vesicatoria.     

Evaluation of proposed amended names of several pseudomonads and xanthomonads and recommendations

ABSTRACT In 1980, over 90% of all plant-pathogenic pseudomonads and xanthomonads were lumped into Pseudomonas syringae and Xanthomonas campestris, respectively, as pathovars. The term "pathovar" was created to preserve the name of plant pathogens, but has no official standing in nomenclature. Proposals to elevate and rename several pathovars of the genera Pseudomonas and Xanthomonas to the rank of species has caused great confusion in the literature. We believe the following changes have merit and expect to adopt them for publication in a future American Phytopathological Society Laboratory Guide for Identification of Plant Pathogenic Bacteria. Upon review of published data and the Rules of The International Code of Nomenclature of Bacteria, we make the following recommendations. We reject the proposal to change the name of P. syringae pvs. phaseolicola and glycinea to P. savastanoi pvs. phaseolicola and glycinea, respectively, because both pathogens are easily differentiated phenotypically from pv. savastanoi and convincing genetic data to support such a change are lacking. We accept the elevation of P. syringae pv. savastanoi to the rank of species. We accept the reinstatement of X. oryzae to the rank of species with the inclusion of X. oryzicola as a pathovar of X. oryzae and we accept the species X. populi. We agree with the elevation of the pvs. cassavae, cucurbitae, hyacinthi, pisi, and translucens to the rank of species but not pvs. melonis, theicola, and vesicatoria type B. We recommend that all type A X. vesicatoria be retained as X. campestris pv. vesicatoria and all type B X. vesicatoria be named X. exitiosa. We reject the newly proposed epithets arboricola, bromi, codiaei (poinsettiicola type B), hortorum, sacchari, and vasicola and the transfer of many pathovars of X. campestris to X. axonopodis. The proposed pathovars of X. axonopodis should be retained as pathovars of X. campestris.