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MS Mincho-�3� fg3&�ArialI�Arial Unicode MS5�Tahoma5$�TahomaB��hF(!F(!��ۆ��T������T��'�0���0���(Journal of Plant Diseases and ProtectioncsMohamed Seleim
��)Journal of Plant Diseases and ProtectioncsNormal.dotmMohamed Seleim2@@��J�@la�Zn�@�h��	�@�h��	�����՜.��+,��D��՜.��+,��\���M 	��08�Caolan80	t$�U��Z��`2���$*4^����		*	�Wb���2	:F,r��Peroxidase and polyphenoloxidase activities as biochemical markers for biocontrol efficacy to control tomato bacterial wilt 
Mohamed A. Seleim1* Kamal A. Abo-Elyousr2, Abd-Alal A. Mohamed1, Hanan A. Al-Marzoky3
1Department of Plant Pathology, Faculty of Agriculture, Al-Azhar University (Assiut Branch), 71524 Assiut, Egypt.2Department of Plant Pathology, Faculty of Agriculture, Assiut University, 71526 Assiut, Egypt. 3Agricultural Botany Department, Faculty of Agriculture, Suez Canal University 41522 Ismailia, Egypt.
* Corresponding author: Mohamed A. Seleim, Department of Plant Pathology, Faculty of Agriculture, Al-Azhar University (Assiut Branch), 71524 Assiut, Egypt,�E-mail:� HYPERLINK "mailto:mohamedseleim@gmail.com"mohamedseleim@gmail.com.
Abstract
We study the effect of certain bioagents to control bacterial wilt of tomato under greenhouse and field conditions and study the effect of these bioagents in induction of some enzyme activity in planta e.g. Peroxidase and polyphenoloxidase. Under greenhouse condatition the effect of Pseudomonas putida and P. fluorescens, and their combination were study, we found that both of them recorded 60 and 66.67% of disease reduction, respectively and the combination treatment reduced the disease was 53.33%. Also under filed condatition P. putida was the best in reduction of the disease followed by the combination and then P. fluorescens. P. fluorescens treatment recorded the highest yield increasing percentagein the two trails and the rest of treatment came in the next rank. The results of the present study revealed that there was significant increase in the activity of PO and PPO and in tomato plants treated with PGPR strains P. fluorescens and P. Putida. Our results support that peroxidase and polyphenoloxidase activities can be used as biochemical marker for biocontrol efficacy to control tomato bacterial wilt.
Keywords: Tomato bacterial wilt, Pseudomonas putida, P. fluorescens, Biochemical changes, induced resistance. 

Introduction
Bacterial wilt caused by Ralstonia solanacearum is become a serious bacterial disease and a major constraint in the production of tomatoes [1]. Infected tomato plants maybe stunted or completely wilted, resulting in poor fruit quality such as small fruit size and significant yield loss about 10%. In Egypt, this disease was first identified in tomato in 2008[2]. Bacterial wilt is among the most difficult diseases to control [3] and the efficacy of current strategies for managementof this disease is limited. No conventional pesticides are known to provide effective control of this soilborne pathogen [4]. The use of plant growth promoting rhizobacteria (PGPR) has become a common practice in many regions of the world. Although significant control of plant pathogens has been demonstrated by PGPR in laboratory and greenhouse studies, results in the field have been inconsistent. Recent progress in our understanding of their diversity, colonizing ability, and mechanism of action, formulation and application should facilitate their development as reliable biocontrol agents against plant pathogens [5]. For many years, the role of oxidative enzymes and of their metabolic products in the defense mechanisms of infected plants has been studied [6]. Peroxidase activity in diseased plants and its effects on resistance or susceptibility in many host-pathogen interactions have been studied [7].The objective of this research was to study peroxidase and polyphenoloxidase activities as biochemical markers after treated by certain biocontrol agents to control tomato bacterial wilt. 
Materials and Methods
Tomato plants
Tomato Seeds (Lycopersicon esculentum Mill.) cv. G.S were surface-sterilized with 2% sodium hypochlorite for 2 min. [11], thoroughly washed with sterilized water, and planted into pots of sterilized soil. After 4 weeks, seedlings were cultivated in the greenhouse in terracotta pots (diameter 30 cm) filled with experimental soil and grown at 25 35oC. 
Source of R. solanacearum
A virulent strain of R. solanacearum biovar 2 isolated from tomato plants, showing typical symptoms of bacterial wilt and identified by the morphological and biochemical characteristics whose pathogenicity on tomato plants had been confirmed in previously work was used in this study. R. solanacearum was isolated on tetrazolium chloride (TTC) agar [8] and cultured routinely on nutrient agar at 28�C for 48 h and temporarily stored in sterile distilled water at 4oC.
Source of rhizobacterial strains
Pseudomonas fluorescens and Pseudomonas putida which isolated from tomto rhizosphere and identified by morphological and biochemical characterization [9-11] were used. Inhibitory activities of rhizhobacteria against R. solanacearum were confirmed by following the dual inoculation technique [13]. Rhizobacteria were cultured on nutrient agar (NA) supplemented with 5% sucrose and stored for further studies.
Preparation of rhizobacterial inoculum
Two days grown culture of two rhizobacteria (Pseudomonas fluorescens and Pseudomonas putida) were harvested from agar plates, sterile deionized water was used for suspending the bacterial cells, and the concentration of the inoculum was determined spectrophotometrically at 600 nm to adjuste the concentration to 108cfu/ml. 
Biocontrol of bacterial tomato wilt under greenhousecondition
In a greenhouse trails, tomato seedlings cv. GS roots were treated separately with each treatment for 30 minutes just before transplanting. 100 ml of pathogen inoculum was added around tomato plants at the time of transplanting. After 4 weeks from inoculation, disease index percentage was recorded using the following formula:
Disease index (%) = [" (ni � vi) � (V � N)] �100

Where ni = number of plants with respective disease rating; vi = disease rating; V = the highest disease rating (5); and N = the number of plants observed [4]. 

Disease rating was calculated as following scale: 
1=no symptoms.
2=one leaf wilted.
3=two to three leaves wilted.
4=four or more leaves wilted.
5=whole plant wilted.
 Treatments were maintained with seven replications and the pots were arranged in completely randomized design.The treatments details are given below:
T1 :Pseudomonas putida (Pp)
T2 :Pseudomonas fluorescens (Pf)
T3 :Pp + Pf
T4 :Control (only Ralstonia solanacearum )
Assay of defense related enzymes 
Sample collection
Samples were collected from individual treatments to study the induction of defense enzymes in response to biocontrol agents treatments under greenhouse conditions. Stems from different treatments were collected and used for analysis.
Enzyme extract
The stem sample, collected from treated and pathogen inoculated tomato plants were immediately homogenized with liquid nitrogen. One g of powdered sample was extracted with 2 ml of sodium phosphate buffer, 0.1M (pH 7.0) at 4�C. The homogenate was centrifuged for 20 min at 10,000 rpm. Protein extracts prepared from tomato tissues were used for estimation of defense enzymes (peroxidase (PO) and polyphenol oxidase (PPO)).The supernatants (crude enzyme extract) were immediately used for determination enzymes activities and total protein. Each treatment consisted in four replications (plants). An aliquot of the extract was used to determine the protein content using bovine serum albumin as standard [14].
Assay of peroxidase (PO)
Assay of PO activity was carried out as described; the reaction mixture consisted of 2.5 ml of a mixture containing 0.25 per cent (v/v) guaiacol in 0.01M. Sodium phosphate buffer, pH 6.0 and 0.1 M hydrogen peroxide. Enzyme extract (0.1 ml) was added to initiate the reaction which was followed colorimetrically at 470 nm. Crude enzyme preparations were diluted to give changes in absorbance at 470 nm of 0.1 to 0.2 absorbance units/min. Boiled enzyme was used as blank. Activity was expressed as the increase in absorbance at 470 nm min-1 mg-1 of protein [15].
Assay of polyphenoloxidase (PPO)
A sample of one g was homogenized in 2 ml of 0.1 M sodium phosphate buffer (pH 6.5) at 4�C. The homogenate was centrifuged at 20,000 rpm for 15 min. The supernatant served as enzyme source and polyphenoloxidase activity was determined as given; the reaction mixture consisted of 1.5 ml of 0.1 M sodium phosphate buffer (pH 6.5) and 200 �l of the enzyme extract. To start the reaction, 200 �l of 0.1M catechol was added and the activity was expressed as change in absorbance at 495 nm at 30-s intervals for 3 min.The enzymeactivity was expressed as changes in absorbance min-1 mg-1 of protein [16].
Biocontrol of bacterial tomato wilt under field conditions
This experiment was carried out in the experimental farm of Plant PathologyDepartment, Faculty of Agriculture, Assiut University, Egypt. Treatments were distributed in a complete randomized block designwith four replicates, the experimental plot area was 25 m2 containing fourteen rows, each row was 3.5 meter length and distance between rows was 50 cm. Tomato seedlings roots were treated by biocontrol agent at the time of transplantingas shown before.100 ml of pathogeninoculum was added around tomato plants. Disease index was recorded as shown in greenhouse trail. Tomato plants were grown up regular watering and after care. Also, the other agriculturalpractices were carried out as the recommended tomato production program of the Egyptian Ministry ofAgriculture.At the end time of experiment, the total yield of each treatment was assessmentper plant.
Statistical analysis
The data from the experiments were subjected to statistical analysis and interpreted by MSTAT-C using LSD test (P=0.05) [17].
Results
Greenhouse experiments
Treatments were screened under greenhouse conditions for their control efficacy against R. solanacearum causing bacterial wilt in tomato and the results showed that all treatments reduce the disease significantly. Pseudomonas putida and Pseudomonas fluorescens, both of them recorded 60 and 66.67% of disease reduction, respectively. Under combination treatment, the disease reduction percentage was 53.33% (Table 1). 
Field trails 
In field trials, the disease control percentage varied from 51.76 % to 88.24%. Results in Table (2) showed that Pseudomonas putida exhibited the disease control percentage 69.41 and 70.59%, respectively in two trails and Pseudomonas fluorescens exhibited 75.29 and 88.24%. Under combination treatment, the disease control percentage was 87.06% and 94.12%, respectively in two trails. 
Yield-increasing percentages have calculated at the end of experiments. As the data in Table (3), Pseudomonas fluorescens treatment recorded the highest yield increasing percentagein the two trails and the rest of treatment came in the next rank.
Assay of defense related enzymes
Assay of peroxidase (PO)
Assay of peroxidase activity in Tomato plants inoculated with R. solancearum showed differences among the various treatments. Increased activity of PO was observed with all treatments upon challenge inoculation withpathogen, when compared to control and it was significantly different (Table 4). PO activity varied from 40.86% to 52.44%. The plants treated with Pseudomonas fluorescens followed by Pseudomonas putida showed higher PO activity irrespective of the pathogen, 52.44 and 49.97 % respectively. Lowest PO activity was observed in combination treatment with 40.86%.
Assay of polyphenoloxidase (PPO)
Data in Table (5) showed that all treatments tested indicated significant increase in the activity of PPO when compared to control plants. PPO activity varied from 39.08% to 62.61%. Maximum PPO activity was noticed in Pseudomonas putida treatment which showed 62.61% increase over control followed by Pseudomonas fluorescens, 42.66% showed increase over control. Lowest PPO activity was observed in combination treatment which registered only 39.08%.
Discussion
Our biological efficacy studies conducted in greenhouse indicated that plants treated with P. fluorescens and P. putida and their combination significantly reduced disease compared to control, similar results were recorded in the conducted studies by [18-21]. Under field conditions, results illustrated that Pseudomonas fluorescens and P. putida and their combination exhibited significantly disease reduction percentage of tomato bacterial wilt. Also, treatments were found to be effective in increasing yield per plant. Our results were in agreement with similar previous studies [22, 26]. The mechanism involved in PGPR-mediated plant growth promotion is directly by production of plant growth regulators (auxins, cytokinins, gibberellins) and facilitation of the uptake of nutrients (nitrogen fixation, solubilization of phosphorus) [37, 38]. The indirect promotion of plant growth occurs when PGPR lessen or prevent the deleterious effects of plant pathogens on plants by production of inhibitory substances (antibiotics, antifungal metabolites, iron-chelating siderophores, cell wall-degrading enzymes and competition for sites on roots) or by increasing the natural resistance of the host (induced systemic resistance) [27, 28]. 
Induced resistance is a state of enhanced defensive capacity against broad spectrum of pests and pathogens developed by a plant when appropriately stimulated [29]. The resulting elevated resistance due to biotic agents is referred to as induce systemic resistance (ISR) whereas that by other than biological control agents is called systemic acquired resistance [30].
In our study, we concentrated on biotic inducers for inducing the defense molecules challenged with Ralstonia solanacearum in tomato. The ISR in this study was primarily focused for the defense related proteins, viz. PO and PPO. The results of the present study revealed that there was significant increase in the activity of PO and PPO and in tomato plants treated with PGPR strains P. fluorescens and P. putida and rest treatments. Similar studies, which showed an increase in PO and PPO activity, were reported by others in increased activity of PO and PPO enzymes in tomato plants treated by PGPR against bacterial wilt caused by Ralstonia solanacearum [21]. The present study also indicated enhanced activity of PO and PPO enzymes due to different treatments, which might have prevented the establishment of R. solanacearum within the tomato roots. Similar increase in plant growth and reduction in bacterial wilt in tomato plants treated with native isolates of P. fluorescens was observed and also increased activity of PO and PPO enzymes [31]. Our results support that peroxidase and polyphenoloxidase activities can be used as biochemical marker for biocontrol efficacy to control tomato bacterial wilt.
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Table 1: Disease incidence and disease reduction percentage of treated tomato plant under greenhouse conditions
TreatmentsDisease incidence (%)Disease reduction (%)Pseudomonas putida (Pp)30 b60Pseudomonas fluorescens (Pf)25 b66.67Pp + Pf35 b53.33Control 75 a0LSD test (P=0.05)28.34* Values in the same column followed by a similar letter are not significantly different as determined by the LSD test (P=0.05).
Table 2: Disease incidence and disease reduction percentage of treated tomato plant under field conditions
Treatments Season 2011Season 2012Disease incidence (%)Disease reduction (%)Disease incidence (%)Disease reduction (%)Pseudomonas putida (Pp)25 c69.41 25 b70.59 Pseudomonas fluorescens (Pf)20 c75.29 10 b88.24 Pp + Pf40 b51.76 25 b70.59 Control 85 a0 70 a0 LSD test (P=0.05)11.9219.22* Values in the same column followed by a similar letter are not significantly different as determined by the LSD test (P=0.05).
Table 3: Influence of different treatments on yield under field conditions
Treatments Season 2011Season 2012Yield (kg per plant)Yield increase (%)Yield (kg per plant)Yield increase (%)Pseudomonas putida (Pp)0.852 c85.163 2.835 b128.629 Pseudomonas fluorescens (Pf)1.102 a139.511 3.188 a157.056 Pp + Pf0.996 b116.576 3.205 a158.468 Control 0.460 d0 1.238 c0 LSD test (P=0.05)0.0470.23* Values in the same column followed by a similar letter are not significantly different as determined by the LSD test (P=0.05).Table 4: Effect of different treatments on PO and PPO enzymes in tomato plants, inoculated with pathogen under greenhouse conditions
TreatmentPOPPO�%in absorbanceIOC%�%in absorbanceIOC%Pseudomonas putida (Pp)0.1321 b
49.97
0.0177 a
42.66Pseudomonas fluorescens(Pf)  0.1343 a52.440.0156 b62.61
Pp + Pf0.1241 c40.860.0152 b39.08Control 0.0881 d00.0109 e0LSD test (P=0.05)5.583.79Values in the same column followed by a similar letter are not significantly different as determined by the LSD test (P=0.05).



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