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Disease symptoms Symptoms include water‐soaked areas surrounded by chlorosis turning into necrotic spots on all aerial parts of plants. On tomato fruits, small, water‐soaked, or slightly raised pale‐green spots with greenish‐white halos are formed, ultimately becoming dark brown and slightly sunken with a scabby or wart‐like surface. Host range Main and economically important hosts include different types of tomatoes and peppers. Alternative solanaceous and nonsolanaceous hosts include Datura spp., Hyoscyamus spp., Lycium spp., Nicotiana rustica, Physalis spp., Solanum spp., Amaranthus lividus, Emilia fosbergii, Euphorbia heterophylla, Nicandra physaloides, Physalis pubescens, Sida glomerata, and Solanum americanum. Taxonomic status of the pathogen Domain, Bacteria; phylum, Proteobacteria; class, Gammaproteobacteria; order, Xanthomonadales; family, Xanthomonadaceae; genus, Xanthomonas; species, X. euvesicatoria, X. hortorum, X. vesicatoria. Synonyms (nonpreferred scientific names) Bacterium exitiosum, Bacterium vesicatorium, Phytomonas exitiosa, Phytomonas vesicatoria, Pseudomonas exitiosa, Pseudomonas gardneri, Pseudomonas vesicatoria, Xanthomonas axonopodis pv. vesicatoria, Xanthomonas campestris pv. vesicatoria, Xanthomonas cynarae pv. gardneri, Xanthomonas gardneri, Xanthomonas perforans. Microbiological properties Colonies are gram‐negative, oxidase‐negative, and catalase‐positive and have oxidative metabolism. Pale‐yellow domed circular colonies of 1–2 mm in diameter grow on general culture media. Distribution The bacteria are widespread in Africa, Brazil, Canada and the USA, Australia, eastern Europe, and south‐east Asia. Occurrence in western Europe is restricted. Phytosanitary categorization A2 no. 157, EU Annex designation II/A2. EPPO codes XANTEU, XANTGA, XANTPF, XANTVE. In this review we provide a historical perspective as well as an updated overview on the aetiology, epidemiology, and management strategies of bacterial spot of tomato and pepper.

Salmonella enterica is ubiquitous in the plant environment, persisting in the face of UV stress, plant defense responses, desiccation, and nutrient limitation. These fluctuating conditions of the leaf surface result in S. enterica population decline. Biomultipliers, such as the phytopathogenic bacterium Xanthomonas hortorum pv. gardneri (Xhg) , alter the phyllosphere to the benefit of S. enterica . Specific Xhg- dependent changes to this niche that promote S. enterica persistence remain unclear, and this work focuses on identifying factors that lead to increased S. enterica survival on leaves. Here, we show that the Xhg transcription activator-like effector AvrHah1 is both necessary and sufficient for increased survival of S. enterica on tomato leaves. An Xhg avrHah1 mutant fails to influence S. enterica survival while addition of avrHah1 to X. vesicatoria provides a gain of function. Our results indicate that although Xhg stimulates a robust immune response from the plant, AvrHah1 is not required for these effects. In addition, we demonstrate that cellular leakage that occurs during disease is independent of AvrHah1. Investigation of the interaction between S. enterica, Xhg, and the plant host provides information regarding how an inhospitable environment changes during infection and can be transformed into a habitable niche.

Four phenotypic xanthomonad groups have been identified that are pathogenic to pepper, tomato, or both hosts. These include groups A and C which are found in Xanthomonas axonopodis pv. vesicatoria, group B found in X. vesicatoria, and group D found in ‘X. gardneri’. We present DNA:DNA hybridization data in which X. axonopodis pv. vesicatoria group A and C strains have less than 70% DNA relatedness with each other, with the type strain of X. axonopodis, and with the currently classified species within Xanthomonas and, therefore, should be removed from this species and given species status. We present information that the A strains most closely resemble the strains originally isolated by Doidge in 1921. In an attempt to avoid confusion in nomenclature as stated in Principle 1 of the Bacteriological Code, we propose that the A strains of X. axonopodis pv. vesicatoria be renamed as X. euvesicatoria (ATCC11633 T = NCPPB2968 T = ICMP 109 T = ICMP 98 T). Use of the euvesicatoria epithet should be reserved for strains originally identified by Doidge, which she designated Bacterium vesicatorium (Ann. Appl. Biol. 7: 407–430, 1921) in the original description when she referred to those strains as being feebly amylolytic. The name X. perforans sp. nov. is proposed for the C group of strains previously designated as X. axonopodis pv. vesicatoria (ATCC BAA-983 T = NCPPB 4321 T). We also propose that ‘X. gardneri’, which has less than 70% DNA relatedness with any of the Xanthomonas species and which has never had taxonomic status, be named X. gardneri (ATCC 19865 T = NCPPB 881 T) to reflect the specific epithet proposed by Sutic [17] in 1957.

We investigated whether the co-occurrence of phytopathogens (Clavibacter michiganensis subsp. michiganensis [Cmm] and Xanthomonas gardneri [Xg]) frequently encountered in tomato production and Salmonella enterica subsp. enterica serotype Typhimurium (strain JSG626) affects the persistence of these pathogens in tomato plant tissues during the early stages of plant growth. Cmm increased the recovery of Salmonella Typhimurium (up to 1.8 log CFU per plant at 21 days postinoculation [DPI]) from coinoculated tomato plants compared with plants inoculated with Salmonella Typhimurium alone (P < 0.05). Xg had no effect on Salmonella Typhimurium persistence in the plants. Increased persistence of Salmonella Typhimurium was also observed when it was inoculated 7 days after Cmm inoculation of the same plant (P < 0.05). In contrast, Salmonella Typhimurium reduced the population of both Cmm and Xg (up to 1.5 log CFU per plant at 21 DPI; P < 0.05) in coinoculated plants compared with plants inoculated with Cmm or Xg alone. The Xg population increased (1.16 log CFU per plant at 21 DPI; P < 0.05) when Salmonella Typhimurium was inoculated 7 days after Xg inoculation compared with plants inoculated with Xg alone. Our findings indicate that the type of phytopathogen present in the phyllosphere and inoculation time influence the persistence of Salmonella Typhimurium JSG626 and its interactions with phytopathogens cocolonized in tomato plants. Salmonella reduced the phytopathogen load in plant tissues, and Cmm enhanced the recovery of Salmonella from the coinoculated plant tissues. However, further investigations are needed to understand the mechanisms behind these interactions.

This study aimed to evaluate the effectiveness of exopolysaccharides (EPS) produced by Lactobacillus plantarum in controlling bacterial spot and eliciting defense mechanisms, and verify alterations in the tomato plant physiological responses. Tomato plants with five definitive leaves were sprayed with EPS, acibenzolar-S-methyl (ASM, positive control), or distilled water (negative control) and inoculated with Xanthomonas gardneri after 3 days. Local and systemic protection was evaluated 7, 15, and 21 days after inoculation. Biochemical defense mechanisms (peroxidases [POX], polyphenoloxidases [PPO], catalase [CAT], superoxide dismutase [SOD], accumulation of hydrogen peroxide [H2O2], and physiological changes) were quantified. In addition, histochemical analyses were examined through light and fluorescence microscopy in treated tomato plants, inoculated or not with X. gardneri. On average, EPS and ASM provided 55 and 96% reduction, respectively, of the bacteriosis severity in treated leaves, compared to the control plants. Increased activities of PPO, CAT, and SOD were found in the EPS-treated plants after being challenged with the pathogen. ASM caused increases in all enzymes analyzed, especially in inoculated plants. The rate of photosynthesis was three times higher, while stomatal conductance and transpiration were 36% lower in the plants sprayed with EPS and challenged with the pathogen, compared to the inoculated control plants. The photochemical efficiency and Soil Plant Analysis Development (SPAD) index did not change. Under light and fluorescence microscopy, it could be seen accumulation of cellulosic compounds in the epidermis of the leaves treated with EPS. Therefore, application of EPS can be considered an alternative for the control of bacterial spot in tomato plants. The study discusses how biochemical and physiological alterations caused by the polysaccharide in the plants contributed to reduce the severity of the disease.

Xanthomonas gardneri is one of the causal agents of bacterial spot (BS), an economically important bacterial disease of tomato and pepper. Field-deployable and portable loop-mediated isothermal amplification (LAMP)-based instruments provide rapid and sensitive detection of plant pathogens. In order to rapidly and accurately identify and differentiate X. gardneri from other BS-causing Xanthomonas spp., we optimized a new real-time monitoring LAMP-based method targeting the X. gardneri-specific hrpB gene. Specificity and sensitivity of real-time and colorimetric LAMP assays were tested on the complex of bacterial strains pathogenic to tomato and pepper and on plants infected by the pathogen. The assay detection limit was 1 pg/μL of genomic DNA with an assay duration of only 30 min. The use of portable and handheld instruments allows for fast analysis, reducing the diagnosis time, and can contribute to proper disease management and control of X. gardneri. Due to the high efficiency of this method, we suggest its use as a standard diagnostic tool during phytosanitary controls.

There are reports of acibenzolar-S-methyl (ASM) having host fitness costs and variable levels of control of bacterial speck (Pseudomonas syringae pv. tomato) and bacterial spot (Xanthomonas gardneri) in tomato (Solanum lycopersicum). The plant growth regulator uniconazole (UNI) is associated with alleviating abiotic stress symptoms, and was tested as an additive to ASM to see if it would reduce ASM-associated fitness costs and improve the consistency of disease control. Field applied ASM (fASM) plus greenhouse applied UNI (gUNI) was less consistent than fASM alone, as the combination reduced disease incidence in only two of three years versus fASM alone that reduced disease incidence in three of three years. However, fASM alone never increased total yield compared to the non-treated control, whereas fASM+gUNI increased it in one of three years, which was not associated with changes in disease intensity or relative chlorophyll levels. Greenhouse applied ASM (gASM) plus gUNI reduced disease incidence in one of three years, whereas gASM alone was never effective. This is the first report that gASM can result in long term disease control reducing disease severity up to 13 weeks post-application, indicating long-term effects of gASM are possible. The lack of improved consistency for disease control or improved yield with ASM combined with UNI compared to ASM alone indicates that other additives need to be tested. Also, further research is needed to discover why the ASM + UNI combination did provide improvements under certain field conditions.

Many phytopathogenic bacteria inject virulence effector proteins into plant cells. To identify novel virulence effectors of the bacterial plant pathogen Xanthomonas, a worldwide collection of pepper (Capsicum annuum) pathogenic Xanthomonas strains was studied. Xanthomonas gardneri strains produced in pepper enhanced watersoaking, a phenotype that is typical of a compatible interaction. Transfer of X. gardneri library clones into a Xanthomonas euvesicatoria recipient strain revealed that enhanced watersoaking was attributable to avrHah1 (avirulence (avr) gene homologous to avrBs3 and hax2, No. 1), a novel avrBs3-like gene. avrHah1 is a novel member of the avrBs3 family that encodes tandemly arranged repeat units of both 34 and 35 amino acid lengths. Although AvrHah1 is only distantly related to AvrBs3, it was shown to trigger a Bs3-dependent hypersensitive response (HR). When fused to a nuclear export signal, AvrHah1 is no longer capable of triggering a Bs3 HR, indicating that nuclear targeting of AvrHah1 is crucial to its recognition. Phylogenetic analysis revealed that, although AvrBs3 and AvrHah1 are only distantly related, they share blocks of high homology within potentially solvent-exposed repeat units. Thus, these data suggest that the recognition specificity of AvrBs3-like proteins is predominantly determined by solvent-exposed residues, rather than by overall homology or repeat unit length.

Bacterial leaf spot of tomato and pepper (BLS), an economically important bacterial disease caused by four species of Xanthomonas (X. euvesicatoria (Xe), X. vesicatoria (Xv), X. gardneri (Xg), and X. perforans (Xp)), is a global problem and can cause over 50% crop loss under unfavorable conditions. Among the four species, Xe and Xv are prevalent worldwide. Characterization of the pathogens is crucial for disease management and regulatory purposes. In this study, we performed a multilocus sequence analysis (MLSA) with six genes (hrcN, dnaA gyrB, gapA, pdg, and hmbs) on BLS strains. Other Xanthomonas species were included to determine phylogenetic relationships within and among the tested strains. Four BLS species comprising 76 strains from different serological groups and diverse geographical locations were resolved into three major clades. BLS xanthomonads formed distinct clusters in the phylogenetic analyses. Three other xanthomonads, including X. albilineans, X. sacchari, and X. translucens pv. undolusa revealed less than 85%, 88%, and 89% average nucleotide identity (ANI), respectively, with the other species of Xanthomonas included in this study. Both antibody and MLSA data showed that Xv was clearly separated from Xe and that the latter strains were remarkably clonal, even though they originated from distant geographical locations. The Xe strains formed two separate phylogenetic groups; Xe group A1 consisted only of tomato strains, whereas Xe group A2 included strains from pepper and tomato. In contrast, the Xv group showed greater heterogeneity. Some Xv strains from South America were closely related to strains from California, while others grouped closer to a strain from Indiana and more distantly to a strain from Hawaii. Using this information molecular tests can now be devised to track distribution of clonal populations that may be introduced into new geographic areas through seeds and other infected plant materials.

Bacterial spot (BS), caused by four species of Xanthomonas: X. euvesicatoria, X. vesicatoria, X. perforans and X. gardneri in tomato (Solanum lycopersicum L.) results in severe loss in yield and quality by defoliation and the appearance of lesions on fruits, respectively. The combined industry standard for BS control (foliar applications Actigard® rotated with copper plus mancozeb) does not offer sufficient protection, especially when weather conditions favor disease spread. Development of tomato cultivars with BS resistance is thus an important measure to minimize losses. Hypersensitive and non-hypersensitive resistance has been identified in different wild accessions and cultivated tomato relatives and has been transferred to cultivated tomato. However, complete resistance is yet to be obtained. With the advent of next generation sequencing and precise genome editing tools, the genetic regions that confer resistance to bacterial spot can be targeted and enriched through gene pyramiding in a new commercial cultivar which may confer higher degree of horizontal resistance to multiple strains of Xanthomonas causing bacterial spot in tomato.