Induccion a la ressitencia sistemica contra el fuego bacteriano

8/19/2019 Induccion a la ressitencia sistemica contra el fuego bacteriano

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ABSTRACT

Induction of Systemic Resistance against bacterial blight of cotton

caused by Xanthomonas axonopodis pv. malvacearum (E.. Smith! "auterin et al .

Gayathri Subbiah* and R. Bhaskaran**

*Assistant Professor, Department of Biotechnology, St. Peters !ngineering

"ollege, A#adi, "hennai.

** Professor, Department of Plant Pathology, $amil %adu Agricultural &ni#ersity,"oimbatore.

Bacterial blight caused by Xanthomonas axonopodis p#. malvacearum is the

serious problem in cotton culti#ation in 'ndia and else(here. Resistance can be

systemically induced in plants lacking the gene for resistance by inoculation (ith non)

pathogens, by restricted inoculation (ith pathogens, or by treatment (ith chemicals.

A study (as undertaken to e#ol#e a management practice for bacterial blight of

cotton caused by Xanthomonas axonopodis p#. malvacearum by induced systemic

resistance using biotic Phylloplane micro organisms and bacterial endophytes+ and

abiotic elicitors. Among the phylloplane micro organisms, Trichoderma viride plf + and

Pseudomonas sp. plb (ere effecti#e in inhibiting the gro(th of the pathogen in vitro.!ndophytes (ere isolated from root, stem and leaf of t(o to si- (eek old cotton

seedlings. !ndo R, !ndo S/ and !ndo 0/ (as effecti#e under in vitro conditions.


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Among the abiotic elicitors salicylic acid, ben1oic acid, dipotassium phosphate,

napthalene acetic acid, glycine, hydrogen pero-ide, o-alic acid, copper sulphate and

indole acetic acid+, tested against the bacterial blight disease, salicylic acid 234 ppm

controlled the disease by producing the least number of lesions per leaf compared to other

chemicals.

Pot culture and field e-periments re#ealed that the per cent disease intensity (as

reduced in salicylic acid sprayed and Pseudomonas sp. plb sprayed plants. $his

treatment also produced increase in the biochemical constituents total sugars, reducing

sugars, non)reducing sugars, protein, phenol, acti#ities of phenylalanine ammonia lyase,

polyphenol o-idase, pero-idase and catalase+ in treated plants.


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I#TR$%&CTI$#

"otton is one of the most important commercial crop of the (orld. 't constitutes nearly 4 per cent

of the ra( material for the te-tile industry. 'n 'ndia, it is culti#ated in an area of .53 lakh ha (ith a

production of 6.44 lakh bales in the year 244)244/. $amil %adu has an area of 6.4 lakh ha (ith a production of .34 lakh bales in the year 244)244/. Diseases are one of the chief constraints in cotton

production. Among the se#eral fungal, bacterial and #iral diseases occurring in cotton, bacterial blight

caused by Xanthomonas axonopodis p#. malvacearum Smith+ 7auterin et al . is a ma8or disease causing

considerable yield losses both in rainfed and summer irrigated cotton. 'n 'ndia, the annual loss ranges from

3 to 23 per cent.

9anagement of the disease using chemicals like copper o-ychloride and antibiotics like

streptomycin sulphate and agrimycin is possible but apart from ha1ards of pollution, the chances of

selection of resistant mutants of the pathogen is high. Biological control of the plant diseases employing

phylloplane and rhi1osphere microflora and more recently by indigenous endophytic bacterial flora is

gaining momentum especially (ith increasing a(areness on the need for use of ecofriendly management

practices. $he plants ha#e e#ol#ed se#eral lines of defense besides pre)e-isting physical and chemical

barriers :sbourn, 6;;loepper 6;;3a and 6;;3b+. Se#eral studies ha#e sho(n that the

interaction bet(een plants and some endophytic bacteria (as associated (ith beneficial effects such as

plant gro(th promotion and biocontrol potential against plant pathogens. 9oreo#er, their uni=ue ability to

sur#i#e in plants (ith no or little microbial competition makes them potential candidates for biological

control 9isaghi and Donndelinger, 6;;4+.

A ne( technology for plant disease control is based on the acti#ation of the plants o(n defense

system (ith the aid of lo( molecular (eight synthetic molecules that induce systemic ac=uired resistance

in plants against a (ide range of microbial pathogens. $he systemic ac=uired resistance SAR+ is an

important component of plants defense against diseases (here initial infection pro#ides systemic

resistance to subse=uent infection by a #ariety of bacterial, fungal and #iral pathogens Gaffney et al .,

6;;+. 'nduced resistance depends on recognition of a pathogen or stress by the plant. $his generates a

cascade of e#ents, e#entually leading to the e-pression of defense mechanisms, (hich include physical

barriers, metabolites and proteins that interfere (ith spread of the in#ading microorganism. 'n recent years,


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a large number of chemicals, viz., salicylic acid, actigard, phosphates, calcium, hydrogen pero-ide,

8asmonic acid, methyl 8asmonate, ethylene, acetyl salicylic acid, ben1othiadia1ole, 2,


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'ATERIAS A#% 'ET)$%S

Collection of isolates of Xanthomonas axonopodis pv. malvacearum

"otton lea#es of the culti#ars 0RA 36


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(as measured after /5 h. $he filter paper disc treated (ith sterile distilled (ater ser#ed as control and

streptomycin sulphate 644 ppm and culture filtrate of Pseudomonas fluorescens Pf 6+ ser#ed as standard

checks.

&se of endophytes

Isolation of bacterial endophytes

'solation of endophytic bacteria (as made from root, stem and lea#es of healthy cotton seedlings of

S7PR 2 #ariety follo(ing those gi#en by 9isaghi and Donndelinger, 6;;4.

Efficacy of bacterial endophytes against Xanthomonas axonopodis pv. malvacearum

$he endophytic bacteria (ere tested against the pathogen by paper disc assay method as described

earlier.

Screening of abiotic elicitors under glass house conditions against bacterial blight of cotton

$he abiotic elicitors viz ., dipotassium phosphate, salicylic acid, indole acetic acid, glycine,

hydrogen pero-ide, napthalene acetic acid, o-alic acid, copper sulphate and ben1oic acid (ere selected for

this study. 'n order to find out the phytoto-ic effects, if any, the plants (ere sprayed (ith the abiotic

elicitors at different concentrations ranging from 34 ppm to 234 ppm and the symptoms of to-icity if any

(ere recorded after a (eek. $o assess their potential for the management of bacterial blight in cotton, the

selected chemicals (ere sprayed at )/)leaf stage and after 2/ h, the plants (ere inoculated (ith the

bacterial blight pathogen. $he symptom e-pression (as obser#ed from 63 days after inoculation and

number of lesions formed (ere assessed. $he chemical that produced less number of lesionsCleaf (as

taken for further study.

*reparation of formulation of antagonistic organisms for pot culture and field studies

$alc based formulation of Pseudomonas, 7idhyasekaran and 9uthamilan, 6;;3+, peat based

formulation of Bacillus Raguchander et al ., 6;;5+ and talc based formulation of Trichoderma (ere

prepared.

Effect of biocontrol agents and abiotic elicitors on bacterial blight of cotton in pot culture

$he effecti#e phylloplane microorganisms, endophytic bacteria and abiotic elicitor (ere taken for

pot culture e-periments. $he pots of 4)cm diameter (ere filled (ith three kg of black soil and fifteen

cotton seeds (ere so(n. Seed treatment (ere done (ith rifampicin mutant endophytic bacteria 2/ h before

so(ing. hen the plants (ere 4 days old, the follo(ing spray treatments (ere gi#en and 2/ h later

pathogen inoculation (as done.


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$6 ) Spraying of phylloplane Bacillus sp.

$2 ) Spraying of phylloplane Pseudomonas sp. plb

$ ) Spraying of phylloplane Trichoderma viride$/ ) Seed treatment (ith root endophytic bacterium !ndo R+

$3 ) Seed treatment (ith stem endophytic bacterium !ndo S/+

$


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?or the analysis of the results, the procedures described by Gome1 and Gome1 6;5/+ (ere

follo(ed.

RES&TS

$he pathogenicity conducted stated that the "oimbatore isolate '2+ produced the ma-imum disease

grade of .< follo(ed by isolate from "heckanuarani '6+.

Se#en fungal and se#en bacterial isolates (ere obtained from the phylloplane of healthy cotton

lea#es.

Efficacy of phylloplane microflora against Xanthomonas axonopodis pv. malvacearum.

Among the fungal isolates, plf Trichoderma viride recorded the ma-imum inhibition 1one of

65.4 mm follo(ed by commercial formulation isolate of Trichoderma viride 6.44 mm+ this (as on par

(ith streptomycin sulphate. Among the bacterial isolates, plb Pseudomonas sp. 3+ recorded the

ma-imum inhibition 1one of 65.3 mm $able 6+.

Efficacy of bacterial endophytes against Xanthomonas axonopodis pv. malvacearum

!ight bacterial endophytes each from root and stem and se#en leaf endophytes (ere isolated.

$he endophytic bacteria (ere tested against the pathogen by paper disc assay method.

Pseudomonas fluorescens Pf 6+ (as used as standard check. Data presented in $able 2 re#ealed that !ndo

R 65.3 mm+, !ndo S/ 65.3 mm+ and !ndo 0/ 65./4 mm+ recorded the ma-imum inhibition 1one.

Screening of abiotic elicitors under glass house conditions against bacterial blight of cotton

'n order to find out an effecti#e abiotic elicitor, #arious chemicals (ere screened under glasshouse

at different concentrations. %one of the chemicals tested as elicitors sho(ed any phytoto-icity. $he data

presented in $able , sho( that salicylic acid at 234 ppm produced least number of lesions per leaf 3.44+

follo(ed by SA 244 ppm 62.44+. @ydrogen pero-ide at 234 ppm produced 6.


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sulphate spray 4.6 per cent (ere taken as standard checks. 'n order to record the biochemical changes

induced in plants, the leaf samples (ere collected at different inter#als of time and disease intensity (as

recorded 63 days after inoculation.

Effect of biotic and abiotic elicitors on bacterial leaf blight

Data presented in $able / sho( that 234 ppm salicylic acid spray significantly reduced the lesion

number per leaf ;.4+ and percent disease inde- 23.44+, follo(ed by spray application of phylloplane

Pseudomonas plb + 64.3 lesion numberCleaf (ith 4 per cent disease inde-+. P. fluorescens Pf !+ and

spray application of Trichoderma viride (ere on par. $he disease intensity in all the three endophyte

treatments (ere on par PD' /3.44+.

Biochemical changes in elicitor treated plants in response to inoculation +ith the bacterial blight

pathogen

$(enty four h after treatment of biotic and abiotic elicitors, the lea#es (ere inoculated (ith

X. axonopodis pv. malvacearum. Biochemical changes (ere recorded from 1ero day to 63 days after

pathogen inoculation.

Sugars

$he data presented in the ?ig 6 sho( that, all the treatments recorded decrease in total sugar

content upto ; days after inoculation. Among the treatments, salicylic acid spray sho(ed #ery high

decrease in total sugars on ;th and 62th day after pathogen inoculation (hen compared to the initial le#el.

$he initial le#el of total sugars in this treatment (as 5.


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le#el of .34 to 6.44 mgCg on 62th day after inoculation of X. axonopodis pv. malvacearum and an

increase on 63th day 65.4 mg glucoseCg leaf tissue+. Phylloplane Pseudomonas sp. plb spray recorded

decrease in reducing sugar content from the initial le#el of 2.44 to 6.24 mg glucoseCg leaf tissue on ; th

day and later increased to 26.64 mg Cg on 63 th day after inoculation. Spray application of Pf 6 recorded

decrease in sugar content on ;th day 6.;4 mg Cg+ and increase on 63 th day 26.


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day+ to 36.23 gCg fresh leaf tissue on ;th day after inoculation and subse=uently decreased to /.34 gCg at

the end of 63th day. Spray application of Pf 6 recorded increased protein content of 34.34 gCg on ; th day

from the initial le#el of 6.24 gCg "ontrol recorded increase in protein content on ; th day 6.23 gCg

fresh leaf tissue+ and later decreased to 25.


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*olyphenol o,idase (**$!

Data presented in the ?ig sho( that PP: acti#ity increased in all the treatments from 1ero to ; th

day after inoculation. Among the treatments, spray application of salicylic acid, phylloplane Pseudomonas

plb spray and Pf 6 spray (ere on par (ith each other. PP: acti#ity (as significantly high on ; th day after

pathogen inoculation (hen compared to other sampling periods. Salicylic acid spray treatment recorded

the highest PP: acti#ity mean #alue 4.4


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62./ and 6.< PD' in field ' and '' follo(ed by streptomycin sulphate 4.6 E+ spray 63.;4 in field ' and

6apas yield

(as the highest in salicylic acid spray treatment 24.; =Cha+ follo(ed by phylloplane Pseudomonas sp. plb

spray 24.5 =Cha+ and both (ere on par. Pf 6 treatment recorded 65.3 =Cha and streptomycin sulphate

spray 6


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!nyedi et al. 6;;2+ reported that salicylic acid induced genes comparable (ith biologically

induced resistance acting as a signal molecule and co)ordinating plant defense gene e-pression. 9uller and

elt1ien 6;;4+ succeeded in inducing resistance in oats against "rechslera avenae using salicylic acid.

Shulae# et al . 6;;3+, 9olders et al . 6;;umar et al ., 244+.

?oliar application of synthetic acti#ators 2,< dichloroisonicotinic acid or ben1o 6,2,+ thiadia1ole

) carbothioic acid S) methyl ester in cotton plants significantly reduced the infection of X. axonopodis p#.

malvacearum "olson et al ., 2444+.

'n the present study, phylloplane bacteria Bacillus sp., Pseudomonas sp. plb and bacterial

endophytes !ndo R, !ndo S/ and !ndo 0/+ (ere selected as the antagonists based on in vitro

antagonism. Pre)inoculation of non)pathogen has been utili1ed for induction of resistance in plants

Gregersen and Petersen, 6;5;+. 9Piga et al . 6;;+ and 7an ees et al . 6;;+ stated that fluorescent

pseudomonads induce resistance against many plant pathogens. 9aurhofer et al . 6;;5+ demonstrated that

salicylic acid produced by P. fluorescens in rhi1osphere is in#ol#ed in induced systemic resistance. $hey

stated that e-pression of salicylic acid biosynthetic genes pch # and pch B+ in P. fluorescens strain P

significantly impro#ed its ability to induce systemic resistance in tobacco against tobacco necrosis #irus.

ei et al . 6;;


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Biochemical changes induced in cotton plants by biotic and abiotic elicitors

$he accumulation of phenol and acti#ities of en1ymes are kno(n to be associated (ith

biochemical disease resistance. $he biochemical mechanisms in#ol#ed in plant disease resistance is a

comple- phenonmenon.

Sugars

Sugars are the building blocks of the defense chemicals phenols and their deri#ati#es

7idhyasekaran, 6;55+. 'nduction of high le#el of sugar by biotic and abiotic elicitors treatment, pro#ides

the substrate needed for the synthesis of defense chemicals. $he results obtained in the present study,

supports this hypothesis, since the phenol le#el increased throughout the sampling period upto 2/ h+ in the

elicitor treated cotton plants. >ehlenbuk et al . 6;;/+ reported that the culture filtrates of Bacillus subtilis

induced resistance against biotrophic leaf pathogens, by altering the carbohydrate metabolism in barley

after the attack by po(dery milde(. $he application of culture filtrates led to increased mo#ement of

assimilates from flag leaf into the ears, (here sugar content also increased, (hich (as used for the

synthesis of defense related compounds.

Protein

Salicylic acid spray and plb sho(ed increase in protein content upto ; days after pathogen

inoculation. 'nduced systemic resistance in tobacco after inoculation (ith Pseudomonas tabaci (as

follo(ed by an increase in the concentration of PR proteins $u1un et al., 6;5;+. Boller 6;53+ e-pressed

that proteins are associated in the defense of plant against fungi and bacteria by their action on the cell

(alls of in#ading pathogens. ard et al . 6;;6+ stated that salicyclic acid application induced gene

families of PR proteins PR)6, PR)2, PR), PR)/, PR)3, PR)H 6+ in tobacco against fungal and bacterial

pathogens.

Phenols

Phenylalanine ammonia lyase PA0+ is the key en1yme of phenylpropanoid metabolism in higher

plants leading to the synthesis of phenols 9assala et al., 6;54+. Salicylic acid spray and phylloplane

Pseudomonas plb spray recorded the ma-imum increase in phenol content after pathogen inoculation.

$he production of phenolic compounds depends upon phenylalanine ammonia lyase en1yme acti#ity

Graham and Graham, 6;;6+. $hanga#elu et al. 244+ reported increase in phenolic content in leaf tissues

to < days after P. fluorescens strain Pf 64 treatment (hen challenge inoculated (ith $usarium

oxysporum in banana.


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Enzyme assay

'ncrease in en1yme acti#ities due to elicitor treatment is one of the resistance response against the

in#ading pathogens. $he en1ymes phenylalanine ammonia lyase PA0+, pero-idase P:+, polyphenol

o-idase PP:+ and catalase play an important role in lignin biosynthesis 7ance et al ., 6;54+. $hese

en1ymes also bring about the production of molecules like phytoale-ins and phenolic compounds that

elicit the steps of induction of resistance >een and Ioshika(a, 6;5J 9auch et al ., 6;55J 7an 0oon et al .,

6;;/+. A large number of en1ymes ha#e been associated (ith 'SR including P:, PA0, lipo-ygenase, K 6,

glucanase and chitinase Ie et al ., 6;;4J >och et al ., 6;;2J Schneider and &llrich, 6;;/ and 7an 0oon,

6;;+.

Phenylalanine ammonia lyase

Salicylic acid or plb treated cotton plants (ere inoculated (ith the bacterial blight pathogen,

there (as increase in PA0 acti#ity. Podile and 0a-mi 6;;5+ stated that the product of PA0, cinnamic acidis directly linked to cell lignification processes and the highest le#el of PA0 acti#ity usually occur about 2/

h after initial infection. @arllen et al . 244/+ studied the effect of rhi1obacteria B646R, B262R and A:


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gene e-pression sho(ed #ariation in transcript accumulation during both compatible and incompatible

interactions for pod %, pod &, pod ' and pod ( genes. Pod ' and pod ( (ere more intensely up regulated

during resistance than during disease and in control, (hile in pod &, it is specifically do(n regulated

during hypersensiti#e reaction @R+ after o-idati#e burst. Pod % (as induced by the pathogen infection

and (eakly stimulated in the control. $he increased P: acti#ity may o-idise phenols to to-ic =uinones and

such o-idation products may inhibit the pathogen de#elopment. Pero-idase itself (as found to inhibit the

pathogen gro(th Loseph et al., 6;;5+. P: along (ith PA0 may also accelerate lignin synthesis.

0ignification has been suggested to inhibit the pathogens in a number of (aysM

i. by pro#iding resistance to en1ymes of the pathogen.

ii. by restricting diffusion of nutrients and (ater from the plant cell to the bacterium or

iii. by inhibiting the bacterial gro(th due to accumulation of phenolic precursors and the

subse=uent free radicals produced during polymeri1ation Ride, 6;5+.

)atalase

'n the elicitor treated and pathogen inoculated cotton lea#es, salicylic acid spray and phylloplane

Pseudomonas sp. plb spray (ere on par (ith regard to increasing the catalase acti#ity. !arlier reports

also suggest that, catalase acti#ity (as enhanced after days of *romyces phaseoli inoculation in bean

plants 9ontalbini, 6;;6+. Amaresh et al.2446+ found increase in catalase acti#ity four days after salicylic

acid treatment in +hizoctonia solani inoculated co(pea plants. @o(e#er, Scandalios 6;;+ has reported

that catalase may act as a free radical sca#enger and as a suppressor in a number of host)pathogen and

elicitor interactions.

'anagement of bacterial leaf blight of cotton in field

?ield e-periments conducted sho(ed that the spray application of salicylic acid and

phylloplane Pseudomonas sp. plb recorded least disease incidence compared to control follo(ed by

spray application of Pseudomonas fluorescens Pf 6+ and streptomycin sulphate. Preinoculation of

phylloplane bacteria protect the plants from pathogen infection reduction in disease se#erity both in terms

of time and intensity+. San8ay and Parashar 2442a+ reported that foliar spray of Pseudomonas fluorescens

pro#ided the ma-imum disease control of cotton bacterial blight /.


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ITERAT&RE CITE%

Ains(orth, G.". 6;ing, !.:.,

ard, 9.>. and D.!. Raney. 6;3/. $(o simple media for demonstration of pyocyanin and fluorescin. -ournal of

aboratory and Clinical 'edicine. / 46)4.

Amaresh, "., An8ali. A., Pranab, >.9. and S. Pradeep. 2446. 'nfluence of salicylic acid on protein content and catalase

acti#ity in relation to systemic ac=uired resistance in co(pea against root rot. Indian *hytopath.0 1/ 25/)25.

Bho(mik, B., Singh, R.P., Layaraman, L. and L.P. 7erma. 2442. Population dynamics of cotton endophytic Pseudomonas,

their antagonism and protecti#e action against the ma8or pathogens of cotton. Indian *hytopath.0 11/ 62/)62.

Boller, $. 6;53. 'nduction of hydrolases as defense reaction against pathogens. In/ Cellular and molecular biology of plant

stress. !dited by >ey, L.0. and $. >osuge. pp 2/)2


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nman, R.!. 6;. and S.9. ong. 6;;5. Antifungal effects of hydrogen pero-ide and pero-idase on spore germination

and mycelial gro(th of Pseudocercospora species. Can. -. Bot.0 79/ 266;)262/.

>een, %.$. and 9. Ioshika(a. 6;5. N 6, endo glucanases from soybean releases elicitor acti#e carbohydrates from fungus

cell (alls. *lant *hysiol.0 76/

/


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9c'nroy, L.A. and L.. >loepper. 6;;3b. Sur#ey of indigenous bacterial endophytes from cotton and s(eet corn. *lant Soil0

675/ )/2. Ryals, L.A., %euensch(ander, &.@., illits, 9.G., 9olina, A., Steiner, @enry Iork and 9.D. @unt.

6;;en. 'icrobiol.0 687/ 6;)25.

9illar, R.0. and 7.L. @iggins. 6;4. Association of cyanide (ith infection of birdsfoot trefoil by /temphylium loti.

*hytopathology0 98/ 64/)664.

9isaghi, '.L. and ".R. Donndelinger. 6;;4. !ndophytic bacteria in symptom free cotton plants. *hytopathology0 ;8/ 545)

566.

9olders, ., Buchala, A. and L.P. 9itrau-. 6;;


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Saha, S., Singh, R.P., 7erma, L. P. and L. Layaraman. 2446. Population dynamics of cotton phylloplane bacteria antagonistic

to(ards Xanthomonas campestris p#. malvacearum. Indian *hytopath.0 1/ /4;)/6.

Samuel, S.>. and $. Patro. 244. !#aluation of biocontrol potential of phylloplane bacteria against bacterial leaf spot of

greengram. Abst. of research papers presented during 33 th annual meeting at :smania &ni#ersity, @yderabad 344

44, Lan 6


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7an ees, S.".9., Pieterse, ".9.L., $dri8ssenaar, I.A., 7an t esterde, @artog, ?. and 0.". 7an 0oon. 6;;. Differential

induction of systemic resistance in #rabidiopsis by biocontrol bacteria. 'ol. *lant:'icrobe Interac.0 68/

6uc. 6;;4. Association of pathogenesis related proteins and acti#ities of pero-idase, K 6, ,

glucanase and =uitinase (ith systemic induced resistance to blue mould tobacco but not to systemic tobacco mosaic

#irus. *hysiol. 'ol. *lant *athol.0 6/ 32)36. >och, !., 9eier, B.9., !ihen, @.G. and A. Slusarenko. 6;;2. A

lipo-ygenase from lea#es of tomato 1ycopersicon esculentum 9ill+ is induced in response to plant pathogenic

pseudomonads. *lant *hysiol.0 44/ 36)3


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plf 2 6


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Bacterial strains

(Root endophytes!

Inhibition 2one (mm!? Bacterial strains (Stem

endophytes!

Inhibition 2one (mm!? Bacterial strains (eaf

endophytes!

Inhibition 2one (mm!?

!ndo R6


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Table 5. Effect of chemicals on bacterial leaf blight of cotton

Chemicals

esion number per leaf?

Concentration (ppm!

34 644 634 244 234 9ean

Ben1oic acid 2. 2.


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Table . Effect of biotic and abiotic elicitors on lesion number per leaf and per cent disease inde, (*%I!

Treatments esion number@leaf?*er cent disease inde,

(*%I! ??

Spraying of phylloplane Bacillus sp. 63.45 b /4.44 ;.2+Spraying of phylloplane Pseudomonas sp. plb 64.3a 4.44 .26+

Spraying of phylloplane $richoderma #iride 6./2 b 3.44


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6.2/+e 4.24+g

/Seed treatment of root endophytic bacteria !ndo R+ 2.54

6.52+ef

2


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6. Spraying of phylloplane Bacillus sp.

2. Spraying of phylloplane Pseudomonas sp. plb . Spraying of phylloplane Trichoderma viride/. Seed treatment of root endophytic bacteria !ndo R+

3. Seed treatment of stem endophytic bacteria !ndo S/+


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6. Spraying of phylloplane Bacillus sp.2. Spraying of phylloplane Pseudomonas sp. plb

. Spraying of phylloplane Trichoderma viride

/. Seed treatment of root endophytic bacteria !ndo R+



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6. Spraying of phylloplane Bacillus sp.2. Spraying of phylloplane Pseudomonas sp. plb . Spraying of phylloplane Trichoderma viride/. Seed treatment of root endophytic bacteria !ndo R+



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2. Spraying of phylloplane Pseudomonas sp. plb . Spraying of phylloplane Trichoderma viride




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2. Spraying of phylloplane Pseudomonas sp. plb


/. Seed treatment of root endophytic bacteria !ndo R+3. Seed treatment of stem endophytic bacteria !ndo S/+


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6. Spraying of phylloplane Bacillus sp.2. Spraying of phylloplane Pseudomonas sp. plb





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/. Seed treatment of root endophytic bacteria !ndo R+3. Seed treatment of stem endophytic bacteria !ndo S/+


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. Spraying of phylloplane Trichoderma viride/. Seed treatment of root endophytic bacteria !ndo R+


Induccion a la ressitencia sistemica contra el fuego bacteriano

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