International Journal of Agricultural

Science and Research (IJASR)

ISSN(P): 2250-0057; ISSN(E): 2321-0087
Vol. 4, Issue 6, Dec 2014, 95-104
TJPRC Pvt. Ltd

ROLE OF PHENOLS AND ANTIOXIDANT ENZYMES IN BIOCONTROL OF FUSARIUM

OXYSPORUM CAUSING FUSARIUM WILT OF ARACHIS HYPOGEAE. L (GROUNDNUT)
P. RAJESWARI
Department of Biochemistry and Molecular Biology, Pondicherry University Puducherry, India

ABSTRACT
The present study examined the effects of culture filtrates of Trichoderma viride, Trichoderma harzianum,
Pseudomonas fluorescens on Fusarium oxysporum infecting Arachis hypogaea. Fusarium wilt diseases caused by the
fungus Fusarium oxysporum lead to significant yield losses of crops. Biocontrol agents such as such as, Trichoderma
viride, Trichoderma harzianum and Pseudomonas fluorescens are applied to control Fusarium wilt of Arachis hypogaea.
Experiments were conducted on the effects of culture filtrates of Trichoderma viride (1%), Trichoderma harzianum
(1.5%), Pseudomonas fluorescens (2%) on the induction of phenol accumulation content and antioxidant enzymes in
Arachis hypogaea infected with Fusarium oxysporum. The Phenol and orthodihydroxy phenol content was recorded to be
higher than that of healthy plants in all the treatments, infected plants treated with Trichodermaviride(1%), Trichoderma
harzianum (1.5%), and Pseudomonas fluorescens (2%) and an increase (3- fold) in the infected plants than that of healthy
plants. Arachis hypogaea plants treated with Trichodermaviride (1%), Trichoderma harzianum(1.5%) and Pseudomonas
fluorescens (2%) showed significant increase in antioxidant enzymes, catalase, peroxidase, and polypohenoloxidase and an
increase (3- 4.5 fold) in the infected plants than that of healthy plants. This present study indicates that applying biocontrol
agents can be a ecologically viable method in plant disease management as it induces the plants own defense mechanism.

KEYWORDS: Fusarium Wilt, Phenols, Catalase, Peroxidase, Polyphenoloxidase

INTRODUCTION
Arachis hypogaea L. (Groundnut) is an important crop in many areas of the world and is found throughout the
tropical, subtropical, and warm temperate regions between 40 north and 40 south latitudes. Fusarium wilt caused by
Fusarium oxysporum has a gradual lethal implication for a groundnut plant. Fusarium oxysporum, the soil borne pathogen
causes vascular wilt diseases in a wide variety of economically important crops (Beckman, 1987). It has been known as
most distributed and important fungal disease on field crops for many years. Vascular wilt has been a major limiting factor
in the production of many agricultural and horticultural crops. Of the vascular wilt-causing Fusaria, Fusarium oxysporum
is the most important species (Agrios, 1997; Smith et al, 1988). Pradeep and Jambhale (2002) suggested that phenolic
compounds and related oxidative enzymes are mostly considered as one of the important biochemical parameters for
disease resistance. They differentiate resistant and susceptible genotypes and as such, would be helpful in discriminating
the genotypes biochemically, in addition to their cytological status. Trichoderma is a genus which include species of
free-living soil fungi, opportunistic, avirulent plant symbionts (Harman et al, 2004), asymptomatic endophytes
(Wilberforce et al, 2003) and parasites of other fungi (Harman, 2006). Trichoderma species possess high reproductive
capacity, ability to survive under very unfavourable conditions, efficiency in the utilization of nutrients, and capacity to
modify the rhizosphere (Woo et al, 2005). These fungi are well known for their ability to produce a wide range of
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antibiotic substances and for their ability to parasitize other fungi. These are highly interactive in roots, soil and foliar
environments and have been used to control many crop pathogens. Trichoderma spp. produce at least three classes of
compounds (viz. peptides, proteins and low molecular weight compounds) that elicit plant defense responses. There have
been studies on the application of antagonistc microbes, such as Pseudomonas spp, for control of Fusaruium wilt
(Tu and Chang, 1983; Duijff et al, 1999 and Leonardo et al, 2006). Thus, the present study was carried out to determine the
induction of phenolic compounds and defense enzymes in F. oxysporum infected Arachis hypogaea plants in response to
the application of Trichoderma spp and Pseudomonas fluorescens

MATERIALS AND METHODS

Trichoderma viride, Trichoderma harzianum and Pseudomonas fluorescens were obtained from Institute of
Microbial Technology (IMTECH), Chandigarh, India and were used for the present study. The pathogen Fusarium
oxysporum was obtained from the infected leaves of Arachis hypogaea and was obtained from the infected leaves of
Arachis hypogaea and was purified by single conidium isolation method. The purified culture was stored in the slants of
PSA. Fusarium oxysporum was grown on PSA for 30 days and further grown in Czapeks medium for 7 days and filtrate
was taken. Trichoderma viride and Trichoderma harzianum were grown on Malt Extract agar and Pseudomonas
fluorescens on ABM Medium and further grown on Czapeks medium in conical flask. It was further centrifuged and
culture filtrate was taken.
Four plants of A. hypogaea raised from the seeds (JLR variety) were grown in each of six earthen pots
(25cm diameter) upto 75 DAS (Day after Sowing) and grouped into three sets. Control-first set of two pots was sprayed
with distilled water on 30 DAS and left without any treatment. Infected- second set of two pots was sprayed with culture of
pathogen, Fusarium oxysporum on 30 DAS and left without any treatment. Infected-treated - third set was sprayed with
pathogen on 30 DAS. These infected plants were sprayed with OIC (Optimum Inhibitoty Concentration) of culture filtrates
of antagonistic microorganisms, Trichoderma viride (1%), Trichoderma harzianum (1.5%) and Pseudomonas fluorescens
(2%) on 40 DAS. On 50 DAS, the leaves of control, infected and infected treated plants were collected for the estimation
of biochemical parameters such as as total phenols, orthodihydroxy phenols, antioxidant enzymes, i.e, catalase, peroxidase,
and polyphenol oxidase.
The total phenol content was estimated according to the method of Bray and Thorpe (1954). To 1 ml of alcoholic
extract, 1 ml of Folin-Ciocalteau reagent and 2 ml of 20% sodium carbonate were added and shaken well. The mixture was
heated in a boiling water bath for 1 min and cooled under running tap water. The blue solution was diluted to 25 ml with
distilled water and read at 650 nm in Systronics Spectrophotometer. Phenols were quantified using catechol as standard.
The Ortho Di-hydroxy phenol content was estimated according to the method proposed by Johnson and Shoal
(1952). To 1 ml of alcoholic extract, 1 ml of 0.5 N HCl and 1 ml of Arnow's reagent were added. To this, 2 ml of 1 N
NaOH and 10 ml of distilled water were added. A pink colour appeared immediately on adding NaOH. The colour
intensity was reduced by diluting it to 25 ml with distilled water and the absorbance read at 515 nm. The O.D. phenols
were calculated using a standard curve with catechol.
Modified method of Luck (1974) was employed for the assay of CAT. To 50 l of the enzyme extract, 3 ml of
hydrogen peroxide-phosphate buffer (pH 7.0) was added. The time required for decrease in absorbance from 0.45 to 0.40
was noted. The enzyme solution, which contained hydrogen peroxide free phosphate buffer was used as control.

Impact Factor (JCC): 4.3594

Index Copernicus Value (ICV): 3.0

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Role of Phenols and Antioxidant Enzymes in Biocontrol of Fusarium

oxysporum Causing Fusarium Wilt of Arachis hypogeae. L (Groundnut)

The activity was expressed as units. The change in the absorbance of 0.001/min/ml of enzyme was assigned as one unit.
Peroxidase activity was measured by the method of Hampton (1962). To 1 ml of 0.001 M pyrogallol, 1.8 ml of
distilled water was added in a cuvette and the absorbance was adjusted to zero at 470 nm. Immediately 0.1 ml of 2%
(0.588 M) H202 and 0.1 ml of enzyme were added. The contents were mixed well and placed in Systronics
Spectrophotometer. The change in the absorbance at every 30-second interval for 3 minutes was measured. Suitable control
with heat killed enzyme was maintained.
Polyphenol oxidase activity was measured by the method of Matta and Dimond (1963). To 0.5 ml of enzyme, 0.5
ml of the phosphate buffer (pH 7.0) and 1.5 ml of distilled water were added and the absorbance was adjusted to zero at
495 nm in Systronics Spectrophotometer and immediately 0.5 ml of 0.1 M catechol was added into the cuvette and the
changes in the absorbance at every 30 seconds intervals upto 3 min was recorded. Control was maintained with heat-killed
enzyme.

RESULTS
The infected plants recorded maximum total phenol (35.62 mg/g), which was 3-fold compared with that of control
plants (11.89 mg/g). An increase of about 1-2% was recorded in all the treatments. The total phenol content in the T. viride,
T. harzianum and P. fluorescens sprayed plants was recorded as 17.82 mg/g, 23.76 mg/g, 14.27 mg/g. (Figure 1)

Figure 1: Total Phenols

The values within a column followed by different letters are significantly different according to Tukeys HSD
multiple range test (TMRT) at 5% level of significance (n=3)
a

p< 0.001 as compared to control

The infected plants recorded maximum O.D phenol (17.91 mg/g), which was 3-fold compared with that of control
plants (6.31 mg/g). An increase of about 1-2% was recorded in all the treatments. The O.D phenol content in the T. viride,
T. harzianum and P. fluorescens sprayed plants was recorded as 11.91 mg/g, 14.07 mg/g, 10.74 mg/g, respectively
(Figure 2).

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Figure 2: Ortho Dihydroxy Phenols

The values within a column followed by different letters are significantly different according to Tukeys HSD
multiple range test (TMRT) at 5% level of significance (n=3)
a

p< 0.001 as compared to control

Highest CAT activity was observed in the infected plants (38.76 SAU) which was 2-times higher than that of

control plants (17.63 SAU). The CAT activity in the T. viride, T. harzianum and P. fluorescens sprayed plants was
recorded as 24.89, 26.70, 23.07 SAU respectively. Slight increase in CAT activity over control plants was observed in all
the treatments (Figure 3).

Figure 3: Catalase
SAU= Specific activity units
a

p< 0.001 as compared to control

1unit= change in the absorbance of 0.001/min/ml of enzyme

The values within a column followed by different letters are significantly different according to Tukeys HSD
multiple range test (TMRT) at 5% level of significance (n=3)
The infected leaves showed maximum activity (102.73 SAU) compared to that of control plants (23.17 SAU).
The PO activity in the T. viride, T. harzianum and P. fluorescens- sprayed plants was recorded as 32.40, 36.15, 31.29 SAU
respectively. An increase of about 1.5 times in PO activity over control plants was observed in all the treatments.
The infected leaves showed maximum activity (150.83 SAU) compared to that of control plants (40.40 SAU). (Figure 4)

Impact Factor (JCC): 4.3594

Index Copernicus Value (ICV): 3.0

Role of Phenols and Antioxidant Enzymes in Biocontrol of Fusarium

oxysporum Causing Fusarium Wilt of Arachis hypogeae. L (Groundnut)

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Figure 4: Peroxidase
SAU= Specific activity units
a

p< 0.001 as compared to control

The values within a column followed by different letters are significantly different according to Tukeys HSD
multiple range test (TMRT) at 5% level of significance (n=3)
The PPO activity in the T. viride, T. harzianum and P. fluorescens- sprayed plants was recorded as 52.85, 61.53,
47.35 SAU respectively. An increase of about 1.5 -2 times in PPO activity over control plants was observed in all the
treatments (Figure 5).

Figure 5: Polyphenoloxidase
SAU= Specific activity units
a

p< 0.001 as compared to control

1unit= change in the absorbance of 0.001/min

The values within a column followed by different letters are significantly different according to Tukeys HSD
multiple range test (TMRT) at 5% level of significance (n=3)

DISCUSSIONS
The total and ortho di-hydroxy phenol contents showed an increase (3- fold) in the leaves of infected plants
compared to those of control plants. Accumulation of phenolic compounds at the infection site has been correlated with the
restriction of pathogen development, since such compounds are toxic to pathogens. Also, phenolic compounds may impede
pathogen infection by increasing the mechanical strength of the host cell wall (Benhamou et al, 2000). Farkas and Kiraly
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(1962) and Jaypal and Mahadevan (1968) found that sharp increased phenol content in the infected plants might be due to
the fact that the accumulation of phenols in the infected tissue might come from the surrounding healthy leaves in order to
resist the advancement of the pathogen towards the other healthy cells.
Of all the treatments T. viride - sprayed plants (1%) showed a minimum increase in the total and ortho di- hydroxy
content followed by T. harzianum-sprayed plants and P. fluorescens- sprayed plants. Ojha and Chatterjee (2012) observed
that the phenol content was significantly higher in F. oxysporum - infected tomato plants treated with either salicylic acid
or Trichoderma harzianum. Induction of total phenol accumulation in the host plant treated with salicylic acid and
Trichoderma harzianum might play an important role in resistance and defense against F. oxysporum. Increase in phenolic
content was observed in plants raised from P. fluorescens- treated seeds when inoculated with R. solani. This might be due
to the induction of systemic resistance in the host plant due to treatments. The over-production of phenolic compounds
resists the advancement of the pathogen towards the other healthy cells. The higher phenol content due to Pseudomonas
treatment was made by Alstrom (1995) in tomato plant infected with P. syringe. Similar findings have been reported by
Ramamoorthy and Samiyappan (2001) that the higher content of phenol in the Pseudomonas-treated chilli plants infected
with C. capsici was mainly attributed to the fact that the phenols are fungitoxic in nature and their accumulation increased
the physical and mechanical strength to the host cell wall resulting in the inhibition of pathogen invasion.
Development of an antioxidant defense system in plants protects them against oxidative stress damage by either
partial suppression of active oxygen species production or the scavenging of active oxygen species which are generated
during plant pathogen interactions (Ye et al, 2006; Cavalcanti et al, 2007). Thus various antioxidant enzymes like
peroxidases and polyphenol oxidases can participate in reactive oxygen metabolism of the species during infection.
Morkunas and Gmerek (2007) stated that peroxidases may be some of the elements of the defense system in response to
pathogen like F. oxysporum. The enhanced antioxidant enzyme activities in the infected host tissue might be due to
induction of systemic resistance in response to the pathogen infection. These enzymes participate in rapid detoxification of
active oxygen species (superoxide anion (02- ); hydrogen peroxide (H2O2); singlet oxygen (O-) into water. The rapid
conversion inhibited the toxic effect caused by AOS to the host plant. In addition, the host cell under pathogenesis might
accelerate the terminal respiratory pathway, which may lead to increase in the CAT activity.
Antioxidant enzymes viz. peroxidase, polyphenol oxidase and catalase showed an increase (3- 4.5 fold) in the
infected plants compared to that of control plants. These results are supported by the Ojha and Chatterjee (2012) in case of
infection of fusarium wilt of tomato. As a consequence of infection, in tomato plants infected with Fusarium oxysporum
f. sp. lycopersici both the polyphenol oxidase and peroxidase enzyme activity was found to be much higher, and gradually
increased as the infection period progressed. Earlier researchers too observed enhanced activity of these enzymes in host
tissues in response to pathogenic infection (Das et al, 2003; Ojha et al, 2005; Chakraborty and Chatterjee, 2007;
Abo-Elyousr et al, 2008). Mohan et al. (1993a, b) reported that the higher activity of phenol oxidase (PO) and polyphenol
oxidase (PPO) in the infected tomato leaves might be due to their participation in the oxidation of phenolic residues into
cell wall polymers in the pathogen-infected cell. Kwon and Anderson (2001) reported enhanced Superoxide dismutase
( SOD) and Catalase ( CAT) isozymes activity in the wheat leaves infected with Fusarium.
The study shows that slight increase was found in catalase, peroxidase and polyphenol oxidase enzyme activities
in T. viride treatment, followed by T. harzianum-sprayed plants and P. fluorescens- sprayed plants compared to control
plants.
Impact Factor (JCC): 4.3594

Index Copernicus Value (ICV): 3.0

Role of Phenols and Antioxidant Enzymes in Biocontrol of Fusarium

oxysporum Causing Fusarium Wilt of Arachis hypogeae. L (Groundnut)

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The treatments generally induced systemic resistance in the host cell, which in turn enhanced activation of these
enzymes in the conversion of reactive oxygen species or radicals to water in order to reduce the infection. F. oxysporum infected plants treated with antagonistic organisms showed higher activity of peroxidase and polyphenol oxidase than the
control plants. Caruso et al. (2001) also experimentally supported the idea that peroxidase plays a defense role against
invading pathogens. Keen (1999) explained that the scavengers (SOD, PO, CAT) of active oxygen species act like
antibiotic against the invading pathogen. The enhanced activity of SOD, PO, CAT recorded in the treated cells generally
restricted the pathogen activity.
Jayalakshmi et al. (2009) reported that PPO activity was increased by the treatment with T. harzianum implicating
it in induced defense responses against root rot in chickpea. Abd-El-Khair et al. (2010) concluded that the levels of
chitinase, peroxidase and polyphenol oxidase activities increased in Trichoderma - treated bean plant compared to
untreated plants. Meena et al. (2001) reported that P. fluorescens induced the activities of PPO in response to infection by
Cercospora personatum in groundnut. The systemic induced resistance by Pseudomonas treatment (Van Loon et al, 1998)
includes increased cell wall structure modification in response to pathogen attack (Chen et al, 2000; Benhamou et al,
2000); accumulation of phenolic compounds (Ramamoorthy and Samiyappan, 2001); induced biochemical and
physiological changes in the treated plants such as enhanced chitinase activity against red rot of sugarcane (Viswanathan
and Samiyappan,1999) and synthesis of phytoalexin and other secondary metabolites (Maurhofer et al, 1994).
The treatments generally induced systemic resistance in the host cell, which in turn enhanced activation of these enzymes
in the conversion of reactive oxygen species or radicals to water in order to reduce the infection. Thus the rapid conversion
reduced the severity of the infection caused by the pathogen.

CONCLUSIONS
Trichoderma spp. as bio-control agents induced the accumulation of some enzymes chitinase, peroxidase and
polyphenol oxidase which played an important role in plant defense mechanisms against pathogen infection and
significantly reduced the disease. Pseudomonas fluorescens capable of inducing higher levels of defence enzymes, PO,
PPO and CAT in the conversion of reactive oxygen species or radicals to water in order to reduce the infection of
F. oxysporum and to induce systemic resistance to disease. This study conclude that inducing the plants own defense
mechanism by applying biocontrol agents can be an effective strategy in plant disease management.

REFERENCES
1.

Abd-El-Khair H, Khalifa, R. Kh. M. and Karima, Haggag H. E ( 2010) Effect of Trichoderma species on damping
off diseases incidence, some plant enzymes activity and nutritional status of bean plants. J. Amer. Sci. 6:122-134.

2.

Abo-Elyousr, A. M. K, Hussein, M. A. M, Allam, A. D. A. and Hassan, A. H. M (2008) Enhanced onion

resistance against stemphylium leaf blight disease, caused by Stemphylium vesicarium, by di-potassium phosphate
and benzothiadiazole treatments. Plant Pathol. J. 24: 171177.

3.

Agrios, G.N (1997) Plant Pathology (4th ed.) Academic Press, New York.

4.

Alstrom, S (1995) Evidence of disease resistance induced by rhizosphere Pseudomonad against Pseudomonas
syringae pv. phaseolicola. J. Gen. Appl. Microbiol. 41: 315-325.

5.

Beckman, C.H (1987) the nature of wilt diseases in plants. Amer. Phytopathol. Soc. St, Paul MN, USA. 175pp

www.tjprc.org

editor@tjprc.org

102

P. Rajeswari

6.

Benhamou N, Gagne, S, Quere, D. L. and Dehbi, L ( 2000) Bacterial mediated induced resistance in cucumber
beneficial effect of the endophytic bacterium Serratia plymuthica on the protection against infection by Pythium
ultimum. Phytopathol. 90: 4546.

7.

Bray, H.G. and Thorpe, W.V (1954) Analysis of phenolic compounds of interest in metabolism. Meth. Biochem.
Anal. 1: 27-52.

8.

Caruso, C, Chilosi, G, Leonard, L, Bertin, L, Magro, P, Buonocore, V. and Caporale (2001) A basic peroxidase
from wheat kernel with antifungal activity. Phytochem. 72: 248 254

9.

Cavalcanti, F. R, Resende, M. L, Lima, S. P, Silveira, J. A. and Oliveira, J.T (2007) Activities of antioxidant
enzymes and photosynthetic responses in tomato pre-treated by plant activators and inoculated by Xanthomonas
vesicatoria. Physiol .Mol. Plant Pathol. 68: 198208.

10. Chakraborty, M. R. and Chatterjee, N.C (2007) Interaction of Trichoderma harzianum with Fusarium solani
during its pathogenesis and the associated resistance of the host. Asian J. Exp. Sci. 21: 353357.
11. Chen, C, Belanger, R. R, Benhamou, N. and Paulitz, T (2000) Defense enzymes induced in cucumber roots by
treatment with plant growth promoting rhizobacteria and Pythium aphanidermatum. Physiol.Mol.Plant Pathol.
56: 13-23
12. Das, S, Aggarwal, R. and Singh, D.V (2003) Differential induction of defense related enzymes involved in lignin
biosynthesis in wheat in response to spot bloth infection. Indian Phytopathol. 56: 129133.
13. Duijff, B.J, Recorbet, G, Peter, A, Bakker, H. M, Loper J. E and Lemanceau, P (1999) Microbial antagonism at
the root level is involved in the suppression of Fusarium wilt by the combination of non pathogenic Fusarium
oxysporum Fo47 and Pseudomonas putida WCS358. Phytopathol. 89: 1073-1079
14. Farkas, G. L. and Kiraly, Z (1962) Role of phenolic compounds in the physiology of plant diseases and disease
resistance. Phytopath. Z. 44: 105-150.
15. Hampton, R. E (1962) Activity of some soluble oxidases in carrot slices infected with Thielaviopsis basicola.
Phvtopath. 52: 497-499.
16. Harman, G.E, Howell, C.R, Viterbo, A, Chet, I. and Lorito, M (2004) Trichoderma species-opportunistic,
avirulent plant symbionts. Nature reviews. Microbiology 2: 43-56.
17. Harman, G. E (2006) Overview of mechanism and use of Trichoderma spp. Phytopathol. 96: 190-194.
18. Jayalakshmi S. K, Raju, S, Usha Rani, S, Benagi, V. I. and Sreeramulu, K ( 2009) Trichoderma harzianum L1 as
a potential source for lytic enzymes and elicitor of defense responses in chickpea (Cicer arietinum L.) against wilt
disease caused by Fusarium oxysporum f. sp. ciceri. Austr. J. Crop Sci. 3:44-52.
19. Jayapal, R. and Mahadevan, A (1968) Biochemical changes in banana leaves in response to leaf spot pathogens.
Indian Phytopath. 21: 43-48
20. Johnson, G. and Shoal, L. A (1952) Relation of Chlorogenic acid to scab resistance in potatoes. Science, 115:
627-629.

Impact Factor (JCC): 4.3594

Index Copernicus Value (ICV): 3.0

103

Role of Phenols and Antioxidant Enzymes in Biocontrol of Fusarium

oxysporum Causing Fusarium Wilt of Arachis hypogeae. L (Groundnut)

21. Keen, N. T (1999) Plant disease resistance: progress in basic understanding and practical application.
In: Advances in Botanical Research. Incorporating advances in plant pathology. Vol.30. (Ed.) T. A. Cellow,
Acad. Press, New York. pp. 291-322.
22. Kwon, S. I. and Anderson, A. J (2001) Differential production of superoxide dismutase and catalase isozymes
during infection of wheat by a Fusarium proliferatum like fungal isolate.
23. Physiol. Mol. Plant Pathol. 58: 73-81.
24. Leonardo, D, Blanca L.F, Landa, B. and Weller D. M (2006) Host crop affects rhizosphere colonization and
competitiveness of 2, 4- diacetylphloroglucinol-producing Pseudomonas fluoresens. Phytopathol. 96: 751-762.
25. Luck, H (1974) Methods in Enzymatic Analysis. (Bergmeyer 2nd ed.). Acad. Press, New York. pp. 885.
26. Matta, A. and Diamond, A. E (1963) Symptoms of Fusarium wilt in relation to quantity of Fusarium and enzyme
activity in tomato stems. Plant pathol.53: 574-578.
27. Maurhofer, M, Hase, C, Meuwly, P, Metraux, JP. and Defago, G (1994) Induction of systemic resistance of
tobacco to tobacco necrosis virus by the root- colonizing Pseudomonas fluorescens strains CHAO: influence of
the gacA gene and pyoverdine production. Phytopathol. 88: 678-684.
28. Meena, B, Marimuthu, T, Vidyasekaran, P. and Velazhahan, R (2001) Biological control of root rot of groundnut
with antagonistic Pseudomonas fluorescens strains. J. Plant Dis. Protect. 108:368-381.
29. Mohan, R, Bajar, A. M. and Kolattukudy, P. E (1993a) Induction of a tomato anionic peroxidase gene (tap 1) by
wounding in transgenic tobacco and activation of tap1: GUS and tap2: GUS chimeric gene fusions in transgenic
tobacco by wounding and pathogen attack. Plant Mol. Biol. 21: 341354.
30. Mohan, R, Vijayan, P. and Kolattukudy, P.E (1993b) Developmental and tissue-specific expression of a tomato
anionic peroxidase (tap1) gene by a minimal promoter, with wound and pathogen induction by an additional
5-flanking region. Plant Mol. Biol. 22:475490.
31. Morkunas, I. and Gmerek J (2007) The possible involvement of peroxidase in defense of yellow lupin embryos
axes against Fusarium oxysporum. J. Plant Physiol. 164: 497506.
32. Ojha, S, Chakraborty, M. R. and Chatterjee, N.C. (2005) Activities of phenol oxidizing enzymes in anthracnose
disease of Saracaasoca under pathogenesis. Indian Biol. 37: 911.
33. Ojha, Suprakash. and Chatterjee, Narayan Chandra (2012) Induction of resistance in tomato plants against
Fusarium oxysporum f.sp. lycopersici mediated through salicylic acid and Trichoderma harzianum. J. Plant Prot.
Res.52:220-225
34. Pradeep T, Jambhale N.D. (2002) Relationship between phenolics, polyphenol oxidase and peroxidase, and
resistance to powdery mildew in Zizyphus. Indian Phytopathol. 55 (2): 195196.
35. Ramamoorthy, V. and Samiyappan, R (2001) Induction of defence related genes in Pseudomonas fluorescens
treated chilli plants in response to infection by Colletotrichum capsici. J. Mol. Pl. Pathol. 31: 146-155.

www.tjprc.org

editor@tjprc.org

104

P. Rajeswari

36. Smith, I. M, Dunez, J, Phillips, D. H, Lelliott, R. A. and Archer, S.A. (Eds.) (1988) European Handbook of plant
diseases. Blackwell Scientific Publications: Oxford. pp.583.
37. Tu, C. C. and Chang, Y. H (1983) Soil microbial activity in relation to Fusarium wilt suppression soils and
conductive soil. In: Proc Republic of China Federal Republic of Germany Seminar Plant Nutrition Soil Sciences.
Natural Science Council, Taipei, pp.189-196.
38. Van Loon, L. C, Bakker, P. A. H. M. and Pieterse, C. M. J (1998) Systemic resistance induced by rhizosphere
bacteria. Annu. Rev. Phytopathol. 36: 453-483.
39. Viswanathan, R. and Samiyappan, R (1999) Identification of antifungal chitinases from sugarcane. ICAR. News
5:1-2.
40. Wilberforce, E. M, Boddy, L, Griffiths, R. and Griffith, G. W (2003) Agricultural management affects
communities of culturable root endophytic fungi in temperate grasslands. Soil Biol. Biochem.35:1143-1154.
41. Woo, S. L, Scala, F, Ruocco, M, Lorito, M (2005) The molecular biology of the interactions between
Trichoderma species, phytopathogenic fungi, and plants. Phytopathol. 96:181-185.
42. Ye, S. F, Zhou, H. Y, Sun, Y, Zou, L. Y, Yu, J. Q ( 2006) Cinnamic acid causes oxidative stress in cucumber
roots and promotes incidence of Fusarium wilt. Environ. Exp. Bot. 56: 255262.

Impact Factor (JCC): 4.3594

Index Copernicus Value (ICV): 3.0