Abstract
Analysis of a Pathogen and Circadian Clock controlled Arabidopsis gene, PCC1, revealed five paralogs in the genome of this model plant on chromosomes 1, 2 and 3. The expression pattern of these five members of the PCC1 super-family under various conditions and after particular stimuli was followed by Northern blot analysis with gene-specific oligo probes. One particular paralogue (At1g05340) was found to be associated with susceptibility to pathogens and altered root length and was chosen for further in-depth study. The locus was renamed “BPR1” (Botrytis and Pseudomonas syringae pv. phaseolicola susceptibility and Root length). BPR1 expression was very low in unstressed plants, but was induced by exogenous application of salicylic acid (SA). BPR1 transcripts were detectable by 2 h after infiltration of Arabidopsis with an avirulent strain of Pseudomonas syringae and one day after infiltration with a higher concentration of virulent P. syringae. Transcript levels remained elevated until the second day after treatment. Inoculation with Botrytis cinerea caused accumulation of BPR1 transcripts for 4 consecutive days whereas PCC1 expression decreased. BPR1 and PCC1 expression did not increase after inoculation with Alternaria brassicicola. Pseudomonas-induced BPR1 expression was observed in several defence signalling mutants (eds1, eds5, sid2 and npr1) and the jasmonic acid signalling pathway mutant (coi1). In contrast, Pseudomonas-induced PCC1 expression did not occur in these defence signalling mutants. However, Pseudomonas induced neither BPR1 nor PCC1 expression in NahG plants. Exogenous application of methyljasmonate and ethylene did not induce BPR1 expression by 4 h after treatment. BPR1 induction was dependent on the resistance gene RPS2 and on NDR1. In contrast, PCC1 induction was not dependent on the RPS2 resistance gene. Organ specific expression studies showed BPR1 was expressed in roots, flowers and infected leaves but not in uninfected leaves and stems. By contrast, PCC1 expression was seen only in infected and uninfected leaves. Histochemical localization of GUS activity in transgenic Col-0 plants demonstrated BPR1 expression in vascular tissue of roots, sepals, and petals and in filaments, developing ovules, stigmas, and infected leaves. Over-expression of BPR1 was achieved by use of the constitutive 35S promoter and a knock out BPR1 line was characterized which had a T-DNA insertion in the promoter region of the BPR1 gene. Four-week-old 35S::BPR1 plants exhibited chlorotic symptoms 8 days after P. syringae pv. phaseolicola infiltration. Bacterial re-isolation experiments revealed at least 7 fold more bacteria in over-expression lines than in the wild-type. 35S::BPR1 plants showed increased susceptibility to Botrytis cinerea in comparison to wild-type and knockout BPR1 plants. In a symptom category from 1 (low) to 5 (intensive), most over-expression plants were placed to categories 4 and 5, in contrast the majority of wild-type and knockout plants which were placed into categories 2 and 3.Roots of 35S::BPR1 plants were almost 30% longer than knockout BPR1. The data from the root length measurements were analyzed by completely randomized design which came from 2 over-expression lines, knockout and wild-type plants. The means of the root lengths were compared by Duncan’s test and differences were found to be significant at the P<1% level, BPR1 knock-out, wild-type and over expression plants comprised 3 separate classes: a, b and c from the shortest to longest roots, respectively. To study genetic polymorphisms in the PCC1 superfamily we chose 10 accessions (Ct-1, Oy-0, Bur-0, Cvi-0, St-0, Shakdara, Mt-0, Mh-1, Ms-0, Ta-0) of Arabidopsis for which McKhann et al. (2004) have shown in genetic differences in four genes. Member-specific primers were chosen from the 5`UTR and 3`UTR regions. Polymorphisms were seen for all PCC1 superfamily members. A PCC1 natural mutant was found in some accessions which showed a 145 bp smaller PCR band. Sequence analysis revealed a deletion of the end of exon 2, of the entire intron 2 and the beginning and the end of exon 3. Additionally there were 4 single nucleotide substitutions in exon 2 and exon 3. Rhythmic transcript fluctuations of the PCC1 natural mutant suggest normal expression at least at the RNA level. BPR1 protein was expressed as a GST (Gluthation S transferase) fusion in E. coli to produce polyclonal antibody. To study BPR1 protein interactions, the lexA yeast-two-hybrid system was used. The BPR1-bait vector was tested in yeast for protein expression and self-activation. A lexA yeast two hybrid cDNA library from pathogen-challenged Arabidopsis was amplified. A preliminary experiment indicated that BPR1 protein might interact with At2g39010, an aquaporin that is known as an intrinsic plasma membrane protein with water channel activity, and water transport protein. In summary, BPR1 and PCC1 appear to have a rather complementary instead of a redundant expression pattern. From my results, BPR1 appears to be a susceptibility factor for certain pathogens in Arabidopsis.
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Claims 20
Claim · model-written summary
Five PCC1 paralogs were found on chromosomes 1, 2 and 3.
- Taxon
- Arabidopsis
Arabidopsist0 - Gene
- PCC1e0
- Outcome
- five paralogse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“PCC1, revealed five paralogs in the genome of this model plant on chromosomes 1, 2 and 3”
t0“Arabidopsis thaliana”
Claim · model-written summary
At1g05340 was associated with pathogen susceptibility and altered root length.
- Taxon
- ArabidopsisArabidopsist0
- Gene
- At1g05340e0
- Outcome
- susceptibility to pathogens and altered root lengthe0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“One particular paralogue (At1g05340) was found to be associated with susceptibility to pathogens and altered root length”
t0“Arabidopsis thaliana”
Claim · model-written summary
Salicylic acid induced BPR1 expression.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- exogenous application of salicylic acid (SA)e0
- Gene
- BPR1e0
- Outcome
- BPR1 expressione0
- Direction
- inducede0
- Comparator
- unstressed plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“BPR1 expression was very low in unstressed plants, but was induced by exogenous application of salicylic acid (SA)”
t0“Arabidopsis thaliana”
Claim · model-written summary
Avirulent Pseudomonas induced BPR1 transcripts by 2 h.
- Taxon
- ArabidopsisArabidopsise0
- Treatment
- infiltration of Arabidopsis with an avirulent strain of Pseudomonas syringaee0
- Interactor
- Pseudomonas syringae
Pseudomonas syringaee0pathogen - Gene
- BPR1e0
- Outcome
- BPR1 transcriptse0
- Value
- 2 he0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“BPR1 transcripts were detectable by 2 h after infiltration of Arabidopsis with an avirulent strain of Pseudomonas syringae”
Claim · model-written summary
Virulent P. syringae induced BPR1 transcripts after one day.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- infiltration with a higher concentration of virulent P. syringaee0
- Interactor
- P. syringaePseudomonas syringaee0pathogen
- Outcome
- Transcript levelse0
- Direction
- remained elevatede0
- Value
- one daye0
Not reportedGeneComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“one day after infiltration with a higher concentration of virulent P. syringae. Transcript levels remained elevated until the second day after treatment”
t0“Arabidopsis thaliana”
Claim · model-written summary
Botrytis inoculation accumulated BPR1 transcripts for four days.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- Inoculation with Botrytis cinereae0
- Interactor
- Botrytis cinerea
Botrytis cinereae0pathogen - Gene
- BPR1e0
- Outcome
- BPR1 transcriptse0
- Direction
- accumulatione0
- Value
- 4 consecutive dayse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Inoculation with Botrytis cinerea caused accumulation of BPR1 transcripts for 4 consecutive days whereas PCC1 expression decreased”
t0“Arabidopsis thaliana”
Claim · model-written summary
Botrytis inoculation decreased PCC1 expression.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- Inoculation with Botrytis cinereae0
- Interactor
- Botrytis cinereaBotrytis cinereae0pathogen
- Gene
- PCC1e0
- Outcome
- PCC1 expressione0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Inoculation with Botrytis cinerea caused accumulation of BPR1 transcripts for 4 consecutive days whereas PCC1 expression decreased”
t0“Arabidopsis thaliana”
Claim · model-written summary
Alternaria inoculation did not increase BPR1 and PCC1 expression.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- inoculation with Alternaria brassicicolae0
- Interactor
- Alternaria brassicicola
Alternaria brassicicolae0pathogen - Gene
- BPR1 and PCC1e0
- Outcome
- BPR1 and PCC1 expressione0
- Direction
- did not increasee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“BPR1 and PCC1 expression did not increase after inoculation with Alternaria brassicicola”
t0“Arabidopsis thaliana”
Claim · model-written summary
Pseudomonas induced BPR1 in the listed defense signaling mutants.
- Taxon
- ArabidopsisArabidopsist0
- Material
- eds1, eds5, sid2 and npr1e0
- Interactor
- Pseudomonas
Pseudomonase0pathogen - Gene
- BPR1e0
- Outcome
- Pseudomonas-induced BPR1 expressione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Pseudomonas-induced BPR1 expression was observed in several defence signalling mutants (eds1, eds5, sid2 and npr1) and the jasmonic acid signalling pathway mutant (coi1)”
t0“Arabidopsis thaliana”
Claim · model-written summary
Pseudomonas induced BPR1 in coi1.
- Taxon
- ArabidopsisArabidopsist0
- Material
- coi1e0
- Interactor
- PseudomonasPseudomonase0pathogen
- Gene
- BPR1e0
- Outcome
- Pseudomonas-induced BPR1 expressione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Pseudomonas-induced BPR1 expression was observed in several defence signalling mutants (eds1, eds5, sid2 and npr1) and the jasmonic acid signalling pathway mutant (coi1)”
t0“Arabidopsis thaliana”
Claim · model-written summary
Pseudomonas induced no PCC1 expression in defense signaling mutants.
- Taxon
- ArabidopsisArabidopsist0
- Interactor
- PseudomonasPseudomonase0pathogen
- Gene
- PCC1e0
- Outcome
- Pseudomonas-induced PCC1 expression did not occure0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Pseudomonas-induced PCC1 expression did not occur in these defence signalling mutants”
t0“Arabidopsis thaliana”
Claim · model-written summary
Pseudomonas induced neither gene in NahG plants.
- Taxon
- ArabidopsisArabidopsist0
- Material
- NahG plantse0
- Interactor
- PseudomonasPseudomonase0pathogen
- Gene
- BPR1 nor PCC1e0
- Outcome
- neither BPR1 nor PCC1 expressione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Pseudomonas induced neither BPR1 nor PCC1 expression in NahG plants”
t0“Arabidopsis thaliana”
Claim · model-written summary
Methyljasmonate and ethylene did not induce BPR1 by 4 h.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- Exogenous application of methyljasmonate and ethylenee0
- Gene
- BPR1e0
- Outcome
- BPR1 expressione0
- Direction
- did not inducee0
- Value
- 4 he0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Exogenous application of methyljasmonate and ethylene did not induce BPR1 expression by 4 h after treatment”
t0“Arabidopsis thaliana”
Claim · model-written summary
BPR1 induction depended on RPS2 and NDR1.
- Taxon
- ArabidopsisArabidopsist0
- Gene
- RPS2 and on NDR1e0
- Outcome
- BPR1 inductione0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“BPR1 induction was dependent on the resistance gene RPS2 and on NDR1”
t0“Arabidopsis thaliana”
Claim · model-written summary
PCC1 induction did not depend on RPS2.
- Taxon
- ArabidopsisArabidopsist0
- Gene
- RPS2e0
- Outcome
- PCC1 inductione0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“PCC1 induction was not dependent on the RPS2 resistance gene”
t0“Arabidopsis thaliana”
Claim · model-written summary
BPR1 was expressed in roots, flowers and infected leaves but not uninfected leaves or stems.
- Taxon
- ArabidopsisArabidopsist0
- Gene
- BPR1e0
- Outcome
- expressed in roots, flowers and infected leaves but not in uninfected leaves and stemse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“BPR1 was expressed in roots, flowers and infected leaves but not in uninfected leaves and stems”
t0“Arabidopsis thaliana”
Claim · model-written summary
PCC1 expression was limited to leaves.
- Taxon
- ArabidopsisArabidopsist0
- Gene
- PCC1e0
- Outcome
- expression was seen only in infected and uninfected leavese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“PCC1 expression was seen only in infected and uninfected leaves”
t0“Arabidopsis thaliana”
Claim · model-written summary
BPR1 expression localized to vascular tissue and reproductive and infected leaf tissues.
- Taxon
- ArabidopsisArabidopsist0
- Gene
- BPR1e0
- Outcome
- expression in vascular tissue of roots, sepals, and petals and in filaments, developing ovules, stigmas, and infected leavese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“BPR1 expression in vascular tissue of roots, sepals, and petals and in filaments, developing ovules, stigmas, and infected leaves”
t0“Arabidopsis thaliana”
Claim · model-written summary
BPR1 overexpression lines had at least sevenfold more bacteria than wild type.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- over-expression linese0
- Outcome
- bacteriae0
- Direction
- moree0
- Value
- at least 7 folde0
- Comparator
- wild-typee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Bacterial re-isolation experiments revealed at least 7 fold more bacteria in over-expression lines than in the wild-type”
t0“Arabidopsis thaliana”
Claim · model-written summary
BPR1 overexpression plants had longer roots than knockout plants.
- Taxon
- ArabidopsisArabidopsist0
- Treatment
- 35S::BPR1 plantse0
- Outcome
- Rootse0
- Direction
- longere0
- Value
- almost 30%e0
- Comparator
- knockout BPR1e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Roots of 35S::BPR1 plants were almost 30% longer than knockout BPR1”
t0“Arabidopsis thaliana”
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