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PubMed2003

The complete genome sequence of the Arabidopsis and tomato pathogen Pseudomonas syringae pv. tomato DC3000.

We report the complete genome sequence of the model bacterial pathogen Pseudomonas syringae pathovar tomato DC3000 (DC3000), which is pathogenic on tomato and Arabidopsis thaliana. The DC3000 genome (6.5 megabases) contains a circular chromosome and two plasmids, which collectively encode 5,763 ORFs. We identified 298 established and putative virulence genes, including several clusters of genes encoding 31 confirmed and 19 predicted type III secretion system effector proteins. Many of the virulence genes were members of paralogous families and also were proximal to mobile elements, which collectively comprise 7% of the DC3000 genome. The bacterium possesses a large repertoire of transporters for the acquisition of nutrients, particularly sugars, as well as genes implicated in attachment to plant surfaces. Over 12% of the genes are dedicated to regulation, which may reflect the need for rapid adaptation to the diverse environments encountered during epiphytic growth and pathogenesis. Comparative analyses confirmed a high degree of similarity with two sequenced pseudomonads, Pseudomonas putida and Pseudomonas aeruginosa, yet revealed 1,159 genes unique to DC3000, of which 811 lack a known function.

Arabidopsis↗
PubMed1984

Cloned avirulence gene of Pseudomonas syringae pv. glycinea determines race-specific incompatibility on Glycine max (L.) Merr.

A genomic library of Pseudomonas syringae pv. glycinea race 6 DNA was constructed in the mobilizable cosmid vector pLAFR1 and maintained in Escherichia coli HB101. Completeness of the library was estimated by assaying clones for the expression of ice-nucleating activity in E. coli. Ice-nucleation activity was represented approximately once in every 600 clones. Six hundred eighty random race 6 cosmid clones were mobilized from E. coli by plasmid pRK2013 in individual conjugations to a race 5 strain of P. s. glycinea. A single clone (pPg6L3) was detected that changed the race specificity of race 5 from virulent (compatible) to avirulent (incompatible) on the appropriate soybean cultivars. The clone was also mobilized from E. coli into race 1 and race 4 strains of P. s. glycinea, and it conferred on these transconjugants the same host range incompatibility as the wild-type race 6 strain. The cosmid clone was mapped by restriction endonucleases, and two adjacent EcoRI fragments were identified by transposon Tn5 mutagenesis to be important in determining race specificity. Southern blot analysis showed that the two EcoRI fragments are unique to race 6 and are not present in the other races tested. The cosmid clone pPg6L3 was also mobilized to Pseudomonas fluorescens and Rhizobium japonicum. However, neither these isolates nor E. coli harboring pPg6L3 elicited a hypersensitive reaction in soybean leaves.

Journal Article↗
PubMed1991

Copper resistance in Pseudomonas syringae mediated by periplasmic and outer membrane proteins.

Copper-resistant strains of Pseudomonas syringae pathovar tomato accumulate copper and develop blue colonies on copper-containing media. Three of the protein products of the copper-resistance operon (cop) were characterized to provide an understanding of the copper-resistance mechanism and its relationship to copper accumulation. The Cop proteins, CopA (72 kDa), CopB (39 kDa), and CopC (12 kDa), were produced only under copper induction. CopA and CopC were periplasmic proteins and CopB was an outer membrane protein. Leader peptide sequences of CopA, CopB, and CopC were confirmed by amino-terminal peptide sequencing. CopA, CopB, and CopC were purified from strain PT23.2, and their copper contents were determined. One molecule of CopA bound 10.9 +/- 1.2 atoms of copper and one molecule of CopC bound 0.6 +/- 0.1 atom of copper. The Cop proteins apparently mediate sequestration of copper outside of the cytoplasm as a copper-resistance mechanism.

Amino Acid Sequence↗
PubMed2000

Cultivar-specific avirulence and virulence functions assigned to avrPphF in Pseudomonas syringae pv. phaseolicola, the cause of bean halo-blight disease.

The avrPphF gene was cloned from Pseudomonas syringae pathovar phaseolicola (PPH:) races 5 and 7, based on its ability to confer avirulence towards bean cultivars carrying the R1 gene for halo-blight resistance, such as Red Mexican. avrPphF comprised two open reading frames, which were both required for function, and was located on a 154 kb plasmid (pAV511) in PPH: Strain RW60 of PPH:, lacking pAV511, displayed a loss in virulence to a range of previously susceptible cultivars such as Tendergreen and Canadian Wonder. In Tendergreen virulence was restored to RW60 by avrPphF alone, whereas subcloned avrPphF in the absence of pAV511 greatly accelerated the hypersensitive resistance reaction caused by RW60 in Canadian Wonder. A second gene from pAV511, avrPphC, which controls avirulence to soybean, was found to block the activity of avrPphF in Canadian Wonder, but not in Red Mexican. avrPphF also conferred virulence in soybean. The multiple functions of avrPphF illustrate how effector proteins from plant pathogens have evolved to be recognized by R gene products and, therefore, be classified as encoded by avirulence genes.

Base Sequence↗
PubMed2002

The Hrp pilus of Pseudomonas syringae elongates from its tip and acts as a conduit for translocation of the effector protein HrpZ.

The type III secretion system (TTSS) is an essential requirement for the virulence of many Gram-negative bacteria infecting plants, animals and man. Pathogens use the TTSS to deliver effector proteins from the bacterial cytoplasm to the eukaryotic host cell, where the effectors subvert host defences. Plant pathogens have to translocate their effector proteins through the plant cell wall barrier. The best candidates for directing effector protein traffic are bacterial appendages attached to the membrane-bound components of the TTSS. We have investigated the protein secretion route in relation to the TTSS appendage, termed the Hrp pilus, of the plant pathogen Pseudomonas syringae pv. tomato. By pulse expression of proteins combined with immunoelectron microscopy, we show that the Hrp pilus elongates by the addition of HrpA pilin subunits at the distal end, and that the effector protein HrpZ is secreted only from the pilus tip. Our results indicate that both HrpA and HrpZ travel through the Hrp pilus, which functions as a conduit for the long-distance translocation of effector proteins.

Bacterial Outer Membrane Proteins↗
PubMed1998

Isolation of new Arabidopsis mutants with enhanced disease susceptibility to Pseudomonas syringae by direct screening.

To identify plant defense components that are important in restricting the growth of virulent pathogens, we screened for Arabidopsis mutants in the accession Columbia (carrying the transgene BGL2-GUS) that display enhanced disease susceptibility to the virulent bacterial pathogen Pseudomonas syringae pv. maculicola (Psm) ES4326. Among six (out of a total of 11 isolated) enhanced disease susceptibility (eds) mutants that were studied in detail, we identified one allele of the previously described npr1/nim1/sai1 mutation, which is affected in mounting a systemic acquired resistance response, one allele of the previously identified EDS5 gene, and four EDS genes that have not been previously described. The six eds mutants studied in detail (npr1-4, eds5-2, eds10-1, eds11-1, eds12-1, and eds13-1) displayed different patterns of enhanced susceptibility to a variety of phytopathogenic bacteria and to the obligate biotrophic fungal pathogen Erysiphe orontii, suggesting that particular EDS genes have pathogen-specific roles in conferring resistance. All six eds mutants retained the ability to mount a hypersensitive response and to restrict the growth of the avirulent strain Psm ES4326/avrRpt2. With the exception of npr1-4, the mutants were able to initiate a systemic acquired resistance (SAR) response, although enhanced growth of Psm ES4326 was still detectable in leaves of SAR-induced plants. The data presented here indicate that eds genes define a variety of components involved in limiting pathogen growth, that many additional EDS genes remain to be discovered, and that direct screens for mutants with altered susceptibility to pathogens are helpful in the dissection of complex pathogen response pathways in plants.

Arabidopsis↗
PubMed2002

In vitro freezing in microtitre plates applied to tobacco plants transformed with the inaZ gene of Pseudomonas syringae.

High throughput assays have been developed to measure the ice nucleation activity of transgenic tobacco, Nicotiana tabacum L. cv. Petit Havana SR1 plants expressing the ice nucleation gene, inaZ, from Pseudomonas syringae at a young seedling stage, as well as in leaf tissue. Both assays are carried out in 96-well microtitre plates. The first assay involves direct seeding in vitro, one seed per microtitre plate well containing Murashige-Skoog agar. When seedlings reach the two-leaf stage, they are exposed to freezing temperatures by floating the plates on a circulating alcohol bath set at temperatures colder than -9 degrees C. The second assay involves placing small leaf discs individually in microtitre plate wells containing sterile distilled water. The assays complement each other, give highly reproducible results, are technically simple and enable the detection of freezing events in large numbers of plants. The utility and limitations of these assays are discussed.

Bacterial Outer Membrane Proteins↗
PubMed1993

An inducible NADP(+)-dependent D-phenylserine dehydrogenase from Pseudomonas syringae NK-15: purification and biochemical characterization.

An inducible NADP(+)-dependent D-phenylserine dehydrogenase [EC 1.1.1.-], which catalyzes the oxidation of the hydroxyl group of D-threo-beta-phenylserine, was purified to homogeneity from a crude extract of Pseudomonas syringae NK-15 isolated from soil. The enzyme consisted of two subunits identical in molecular weight (about 31,000). In addition to D-threo-beta-phenylserine, it utilized D-threo-beta-thienylserine, D-threo-beta-hydroxynorvaline, and D-threonine as substrates but was inert towards other isomers of beta-phenylserine and threonine. It showed maximal activity at pH 10.4 for the oxidation of D-threo-beta-phenylserine, and it required NADP+ as a natural coenzyme. NAD+ showed a slight coenzyme activity. The enzyme was inhibited by p-chloromercuribenzoate, HgCl2, and monoiodoacetate but not by the organic acids such as tartronate. The Michaelis constants for D-threo-beta-phenylserine and NADP+ were 0.44 mM and 29 microM, respectively. The N-terminal 27 amino acids sequence was determined. It suggested that the NADP(+)-binding site was located in the N-terminal region of the enzyme.

Alcohol Oxidoreductases↗
PubMed2003

The rapid induction of glutathione S-transferases AtGSTF2 and AtGSTF6 by avirulent Pseudomonas syringae is the result of combined salicylic acid and ethylene signaling.

The expression of two members of the glutathione S-transferase (GST) multigene family was studied in Arabidopsis plants inoculated with an avirulent strain of Pseudomonas syringae pv. tomato (Pst). Accumulation of AtGSTF2 and AtGSTF6 transcripts started 4 and 2 h after inoculation, respectively, and clearly preceded the induction of the pathogenesis-related PR-1 gene. The aim of this work was to find the reason for the faster induction of the two GSTs compared with classical salicylic acid (SA)-regulated PR-proteins. Expression studies in Pst-inoculated SA-signaling mutants NahG and npr1 revealed that induction of both GSTs was SA-dependent and partially NPR1-independent. The induction of AtGSTF2 by Pst was also strongly repressed in the ethylene insensitive etr1 mutant. Both GSTs were induced by low amounts of SA (0.1 mM) and ethylene (0.1 ppm) while PR-1 gene expression was unaffected by ethylene. Interestingly, ethylene was about 50-fold less effective in NahG compared with wild-type plants thus suggesting a potentiation effect of SA on ethylene-induced accumulation of AtGST transcripts. Increased AtGST expression in plants inoculated with Pst correlated with increased production of SA and ethylene. However, the initial phase of AtGSTF6 induction was independent of SA- and ethylene-signaling. The jasmonate (JA)-insensitive mutant jar1 showed normal induction kinetics for both GSTs. Our data support the hypothesis that full expression of the pathogen-induced AtGSTF2 and, to a lesser extent AtGSTF6, is the result of combined SA- and ethylene-signaling and that early AtGSTF6 expression depends on additional unknown signaling mechanisms.

Arabidopsis↗
PubMed2004

Up-regulation and localization of asparagine synthetase in tomato leaves infected by the bacterial pathogen Pseudomonas syringae.

Nitrogen metabolism is one aspect of basic metabolism, which is still quite unknown in the field of plant-pathogen interactions. Evidence derived from previous studies conducted in our laboratory strongly suggests that during microbial pathogenesis an important nitrogen mobilization process takes place in diseased tissues. Here we describe the expression pattern of asparagine synthetase (AS; EC 6.3.5.4) in tomato leaves infected by the bacterial pathogen Pseudomonas syringae pv. tomato. Using an homologous AS cDNA probe isolated by RT-PCR from infected leaves, we have observed a high level induction of AS expression during the course of infection. Concomitantly, a single AS polypeptide also accumulated in response to bacterial infection. Furthermore, immunohistochemical analysis of AS in infected leaves revealed a strong immunostaining in phloem cells of the main vascular bundles and in secondary veins of the leaf blade. These data correlate with those previously reported for expression of a cytosolic isoform of glutamine synthetase (GS1) also induced during development of the infectious process. Taken together, our results suggest the existence of a GS1/AS pathway representing a metabolic route for transferring ammonium released from protein catabolism into asparagine, an amino acid that may have a major role in nitrogen mobilization from diseased tissues.

Amino Acid Sequence↗
PubMedDate not supplied

A gene from Pseudomonas syringae pv. glycinea with homology to avirulence gene D from P. s. pv. tomato but devoid of the avirulence phenotype.

A gene was cloned from Pseudomonas syringae pv. glycinea that hybridized to avirulence gene D (avrD), previously cloned from P. s. pv. tomato. Unlike avrD, the hypersensitive response (HR) was not elicited when the P. s. pv. glycinea gene was reintroduced into P. s. pv. glycinea race 4 on a broad host range plasmid and the bacteria were inoculated into soybean leaves. DNA sequence data disclosed that the P. s. pv. glycinea homologue of avrD encoded a protein containing 86% identical amino acids to avrD, with substitutions distributed throughout the protein. Two ORFs immediately downstream from the avrD homologue were more similar in P. s. pv. tomato and P. s. pv. glycinea, with 98 and 99% identical amino acids. Expression of the wildtype P. s. pv. glycinea gene and recombinant genes constructed between the P. s. pv. tomato avrD gene and its P. s. pv. glycinea homologue in both Escherichia coli and P. s. pv. glycinea indicated that the P. s. pv. glycinea gene product was formed less efficiently or was less stable than was the P. s. pv. tomato protein encoded by avrD. The data indicated that the P. s. pv. glycinea homologue represents a recessive allele of the P. s. pv. tomato avrD gene which has been modified by mutation such that it does not lead to an avirulence phenotype on the normal host plant, soybean.

Amino Acid Sequence↗
PubMedDate not supplied

Identification of a lysA-like gene required for tabtoxin biosynthesis and pathogenicity in Pseudomonas syringae pv. tabaci strain PTBR2.024.

Pseudomonas syringae pv. tabaci strain PTBR2.024 produces tabtoxin and causes wildfire disease on tobacco and green bean. PTBR7.000, a Tn5 mutant of PTBR2.024, does not produce tabtoxin, is nonpathogenic on tobacco, and is prototrophic. A 3-kb fragment from a genomic library of the parent strain PTBR2.024 complemented both mutant phenotypes. This 3-kb fragment contains two open reading frames (ORFs), ORF1 and ORF2, and two truncated ORFs, ORF3 and ORF4. The Tn5 insert in PTBR7.000 was mapped to ORF2, and complementation studies showed that an intact ORF2 was sufficient to restore tabtoxin production and pathogenicity. The deduced amino acid sequences of ORF2 and truncated ORF3 contain significant homology to bacterial lysine biosynthetic enzymes, diaminopimelate decarboxylase, and delta 1-piperidine-2,6-dicarboxylate succinyl transferase, respectively. ORF2, however, is not required for lysine biosynthesis. We designated the sequence corresponding to ORF2 as gene tabA and propose that the product of tabA is an enzyme in the tabtoxin biosynthetic pathway that recognizes a substrate analogue of a compound in the lysine biosynthetic pathway.

Acyltransferases↗
PubMedDate not supplied

Pathological and molecular characterizations of alfalfa interactions with compatible and incompatible bacteria, Xanthomonas campestris pv. alfalfae and Pseudomonas syringae pv. pisi.

We report on the interactions of alfalfa with Xanthomonas campestris pv. alfalfae and Pseudomonas syringae pv. pisi. A hypersensitive response was observed when leaves were infiltrated with P. s. pv. pisi, which remained strictly limited to the injected zone. The compatible interaction with X. c. pv. alfalfae was characterized by water-soaking symptoms and the spreading of the bacterium into the leaf blade. Analyses of transcript accumulation were conducted with cDNAs encoding enzymes involved in phytoalexin synthesis: chalcone synthase (CHS), chalcone isomerase (CHI), and isoflavone reductase (IFR). In incompatible interactions the maximum accumulation of the CHS, CHI, and IFR transcripts was observed 6 hr postinfection. In the compatible interaction, the induction of these transcripts was delayed until 25-30 hr postinfection, and the level of their accumulation was considerably lower. Extending this molecular analysis to the root system showed that the reaction of roots during an incompatible interaction was quite comparable to that of leaves. To complete these analyses, expression of genes encoding pathogenesis-related (PR) proteins in leaves was also analyzed by polymerase chain reaction. High-level accumulation of a 0.8-kb transcript encoding a PR protein was observed 6 to 30 hr postinfection in the incompatible interaction.

Acyltransferases↗
PubMedDate not supplied

Role of the phytotoxin coronatine in the infection of Arabidopsis thaliana by Pseudomonas syringae pv. tomato.

The role of the phytotoxin coronatine in the virulence of Pseudomonas syringae pv. tomato in Arabidopsis thaliana was evaluated by comparing symptom development, in planta bacterial multiplication, and the induction of defense-related genes in Arabidopsis plants inoculated with the coronatine-producing (Cor+) P. s. pv. tomato strain DC3000 and the coronatine-defective (Cor-) strain DC3661 by either infiltration or dipping methods. The Cor+ strain, P. s. pv. tomato DC3000, caused severe disease symptoms and multiplied by 4-6 logs after inoculation by either infiltration or dipping. P. s. pv. tomato DC3661 failed to produce any disease symptoms and multiplied by only 1-1.5 logs in dipped plants, whereas it caused mild symptoms and multiplied 6 logs over the 4-day experimental period in plants inoculated by infiltration. Parallel experiments using a natural host, tomato, yielded similar results. Analysis of the accumulation of mRNAs encoded by several distinct defense-related genes in Arabidopsis leaves infiltrated with either DC3000 or DC3661 demonstrated that the Cor- strain consistently induced higher levels of these transcripts. These results demonstrate that coronatine production is required under more natural inoculation conditions for the successful infection of Arabidopsis by DC3000, and that coronatine may play a critical role during the early stages of infection by suppressing the activation of defense-related genes.

Amino Acids↗
PubMed1998

A race-specific insertion of transposable element IS801 in Pseudomonas syringae pv. phaseolicola.

The isolation and cloning of a random amplified polymorphic DNA-polymerase chain reaction (RAPD-PCR) band specific for Pseudomonas syringae pv. phaseolicola race 1 allowed us to design a pair of primers that amplify 1.2-kb race 1-specific and 2.7-kb race 2-specific fragments, providing a rapid method for the identification of races by standard PCR methods. Restriction analysis revealed identical endonuclease sites in both fragments but the race 2 fragment contains a 1.5-kb insertion, identified as transposable element IS801 by sequence comparison. One complete and one partial open reading frame (ORF), each with a high probability of encoding a protein, have been identified in the 1.2-kb fragment common to both race 1 and race 2 sequences. As IS801 disrupts the partial ORF in the race 2 fragment, the complete sequence of this ORF has been obtained as well as its promoter region. The possibility that it may encode an avirulence gene is discussed as well as the role of transposable elements in pathogen evolution.

Base Sequence↗
PubMed1999

Identification of a locus in arabidopsis controlling both the expression of rhizobacteria-mediated induced systemic resistance (ISR) and basal resistance against Pseudomonas syringae pv. tomato.

Selected nonpathogenic rhizobacteria with biological disease control activity are able to elicit an induced systemic resistance (ISR) response that is phenotypically similar to pathogen-induced systemic acquired resistance (SAR). Ten ecotypes of Arabidopsis thaliana were screened for their potential to express rhizobacteria-mediated ISR and pathogen-induced SAR against the leaf pathogen Pseudomonas syringae pv. tomato DC3000 (Pst). All ecotypes expressed SAR. However, of the 10 ecotypes tested, ecotypes RLD and Wassilewskija (Ws) did not develop ISR after treatment of the roots with nonpathogenic Pseudomonas fluorescens WCS417r bacteria. This nonresponsive phenotype was associated with relatively high susceptibility to Pst infection. The F1 progeny of crosses between the non-responsive ecotypes RLD and Ws on the one hand, and the responsive ecotypes Columbia (Col) and Landsberg erecta (Ler) on the other hand, were fully capable of expressing ISR and exhibited a relatively high level of basal resistance, similar to that of their WCS417r-responsive parent. This indicates that the potential to express ISR and the relatively high level of basal resistance against Pst are both inherited as dominant traits. Analysis of the F2 and F3 progeny of a Col x RLD cross revealed that inducibility of ISR and relatively high basal resistance against Pst cosegregate in a 3:1 fashion, suggesting that both resistance mechanisms are monogenically determined and genetically linked. Neither the responsiveness to WCS417r nor the relatively high level of basal resistance against Pst were complemented in the F1 progeny of crosses between RLD and Ws, indicating that RLD and Ws are both affected in the same locus, necessary for the expression of ISR and basal resistance against Pst. The corresponding locus, designated ISR1, was mapped between markers B4 and GL1 on chromosome 3. The observed association between ISR and basal resistance against Pst suggests that rhizobacteria-mediated ISR against Pst in Arabidopsis requires the presence of a single dominant gene that functions in the basal resistance response against Pst infection.

Arabidopsis↗
PubMed1999

A new peroxidase cDNA from white clover: its characterization and expression in root tissue challenged with homologous rhizobia, heterologous rhizobia, or Pseudomonas syringae.

Temporal reverse transcription-polymerase chain reaction (RT-PCR) expression analyses were performed on Trprx2, a new white clover peroxidase, with roots challenged with homologous rhizobia, heterologous rhizobia, and a pathogen, Pseudomonas syringae. Low levels of Trprx2 expression were evident in all rhizobial treatments but in P.syringae-treated clover background expression was dramatically reduced within 1 h and was undetectable in treatments inoculated for more than 3 h. Spraying 4 mM salicylic acid onto seedlings increased Trprx2 expression. These data suggest a defensive role for Trprx2 in white clover and indicate active defense suppression by the pathogen.

Amino Acid Sequence↗
PubMed2000

avrPto enhances growth and necrosis caused by Pseudomonas syringae pv.tomato in tomato lines lacking either Pto or Prf.

AvrPto was introduced into three tomato genotypes with two biotic agents to study its role in compatible interactions. avrPto enhanced the capacity of the Pseudomonas syringae pv. tomato strain T1 to induce necrotic symptoms on tomato plants that lacked either Pto or Prf genes. The enhanced necrosis correlated with a small increase in bacterial growth. In planta expression of avrPto in isolation did not elicit necrosis in the absence of a functional Prf gene.

Genes, Plant↗