The Open Agriculture Journal




ISSN: 1874-3315 ― Volume 14, 2020
RESEARCH ARTICLE

Inhibition of Mycelial Growth of Rhizoctonia Solani by Chitosan in vitro and in vivo



Sabah R. Mohammed1, 2, *, Elsayed M. Zeitar3, Ivan D. Eskov1
1 Plant Protection Department, Faculty of Agriculture, Saratov State Vavilov Agrarian University, Saratov, 410012, Russia
2 Plant Protection Department, Faculty of Agriculture, Damanhour University, Damanhour, 22511, Egypt
3 Plant Pathology Department, Faculty of Agriculture, Damanhour University, Damanhour, 22511, Egypt

Abstract

Objective:

Evaluate the antifungal effect of chitosan against Rhizoctonia solani in vitro and the possible mechanisms of its induced activity in potato tubers to control black scurf disease.

Methods:

The in vitro influence of chitosan at different concentrations on mycelial growth of R. solani was tested by using the poisoned food technique in PDA medium. The effect of these concentrations on the development of lesion diameters in tubers inoculated with R. solani mycelium was assayed for 30 days. The concentration that showed the greatest inhibitory effect on lesion diameters was tested to assess the induced activity of defense-related enzymes in the infected tubers.

Results:

In the poisoned food technique, chitosan at 1% completely inhibited the growth of R. solani mycelium. In vivo tests showed that chitosan treatment at 0.5% effectively controlled the black scurf in tubers inoculated with R. solani mycelium. Chitosan increased the activities of defense-related enzymes such as Peroxidase (POD), Polyphenol Oxidase (PPO) and Phenylalanine Ammonia-lyase (PAL) in treated tubers of tested cultivars.

Conclusion:

This work demonstrated that chitosan directly inhibited the growth of R. solani, and potentially elicited defense reaction in potato tubers.

Keywords: Chitosan, Induced resistance, Potato, Rhizoctonia solani, Defense enzymes, Black scurf, Peroxidase.


Article Information


Identifiers and Pagination:

Year: 2019
Volume: 13
First Page: 156
Last Page: 161
Publisher Id: TOASJ-13-156
DOI: 10.2174/1874331501913010156

Article History:

Received Date: 18/07/2019
Revision Received Date: 20/09/2019
Acceptance Date: 13/10/2019
Electronic publication date: 20/12/2019
Collection year: 2019

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© 2019 Mohammed et al.

open-access license: This is an open access article distributed under the terms of the Creative Commons Attribution 4.0 International Public License (CC-BY 4.0), a copy of which is available at: (https://creativecommons.org/licenses/by/4.0/legalcode). This license permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


* Address correspondence to this author at the Plant Protection Department, Faculty of Agriculture, Saratov State Vavilov Agrarian University, Saratov, 410012, Russia; Tel: +79026665260; E-mail: srabie106@gmail.com





1. INTRODUCTION

Rhizoctonia solani Kühn [teleomorph: Thanatephorus cucumeris (Frank) Donk] AG-3 PT, the pathogen of stem canker and black scurf in potato (Solanum tuberosum L.), is a worldwide plant disease [1Banville GJ, Carling DE, Otrysko BE. Rhizoctonia disease on potato Rhizoctonia species: Taxonomy, molecular biology, ecology, pathology and disease control 1996; 321-30.
[http://dx.doi.org/10.1007/978-94-017-2901-7_29]
]. Yield losses from R. solani can reach 50%, resulting in marked economic losses for farmers [2Keiser A. Rhizoctonia solani a fungal disease with multiple symptoms: means of preventive and curative control. Potato Research for a Production of Quality Information Day 2008.]. Control of R. solani relies mainly on fungicides application [3El Bakali AM, Martín MP. Black scurf of potato. Mycologist 2006; 20(4): 130-2.
[http://dx.doi.org/10.1016/j.mycol.2006.03.006]
] such as azoxystrobin, flutolanil, and pencycuron [4Campion C, Chatot C, Perraton B, Andrivon D. Anastomosis groups, pathogenicity and sensitivity to fungicides of Rhizoctonia solani isolates collected on potato crops in France. Eur J Plant Pathol 2003; 109(9): 983-92.
[http://dx.doi.org/10.1023/B:EJPP.0000003829.83671.8f]
, 5Djébali N, Belhassen T. Field study of the relative susceptibility of eleven potato (Solanum tuberosum L.) varieties and the efficacy of two fungicides against Rhizoctonia solani attack. Crop Prot 2010; 29(9): 998-1002.
[http://dx.doi.org/10.1016/j.cropro.2010.06.012]
] which are not always efficient due to the development of fungicide resistant communities. Furthermore, pesticides are environmental and human health concern. Consequently, more momentum has gained in search of sustainable solutions to chemical control. Chitosan (CH) (poly-β-(1 4) N-acetyl-d-glucosamine) is a modified, natural safe biopolymer derived by deacetylation of chitin, the second most abundant natural polymer in the world [6Rabea EI, Badawy ME, Steurbaut W, Stevens CV. In vitro assessment of N-(benzyl) chitosan derivatives against some plant pathogenic bacteria and fungi. Eur Polym J 2009; 45(1): 237-45.
[http://dx.doi.org/10.1016/j.eurpolymj.2008.10.021]
, 7El Hadrami A, Adam LR, El Hadrami I, Daayf F. Chitosan in plant protection. Mar Drugs 2010; 8(4): 968-87. Available at: http://www.ncbi.nlm. nih.gov/pubmed/20479963
[http://dx.doi.org/10.3390/md8040968] [PMID: 20479963]
]. Chitosan is very useful for several industries, such as cosmetology, biotechnology, food, pharma- cology and medicine [8Hamed I, Özogul F, Regenstein JM. Industrial applications of crustacean by-products (chitin, chitosan, and chitooligosaccharides): A review. Trends Food Sci Technol 2016; 48: 40-50.
[http://dx.doi.org/10.1016/j.tifs.2015.11.007]
, 9Choi C, Nam J-P, Nah J-W. Application of chitosan and chitosan derivatives as biomaterials. J Ind Eng Chem 2016; 33: 1-10.
[http://dx.doi.org/10.1016/j.jiec.2015.10.028]
]. Since the 1980s, farmers have used chitosan as biopesticide, biofertilizer and agricultural film in seed and fruit coating [10Malerba M, Cerana R. Chitosan Effects on Plant Systems. Int J Mol Sci 2016; 17(7): 996. Available at: http://www.ncbi.nlm.nih.gov/pubmed/27347928
[http://dx.doi.org/10.3390/ijms17070996] [PMID: 27347928]
]. Chitosan has been demonstrated to control postharvest diseases on several horticultural commo- dities such as apples, pears, kiwifruit, strawberries, tomato, raspberries, table grape and others [11Du J, Gemma H, Iwahori S. Effects of chitosan coating on the storage of peach, Japanese pear, and kiwifruit. J Jpn Soc Hortic Sci 1997; 66(1): 15-22.
[http://dx.doi.org/10.2503/jjshs.66.15]
-15Meng X, Li B, Liu J, Tian S. Physiological responses and quality attributes of table grape fruit to chitosan preharvest spray and postharvest coating during storage. Food Chem 2008; 106(2): 501-8.
[http://dx.doi.org/10.1016/j.foodchem.2007.06.012] [PMID: 2604 9245]
]. Chitosan protects rice, tomato, tobacco, and lettuce plants against R. solani infection [16Abd-El-Kareem F, El-Mougy NS, El-Gamal NG, Fotouh Y. Use of chitin and chitosan against tomato root rot disease under greenhouse conditions. Res J Agric Biol Sci 2006; 2: 147-52.-19Kurzawińska H. Potential use of chitosan in the control of lettuce pathogens. Polish Chitin Society. Monograph 2007; 12: 173-8.]. However, to our recent knowledge, there is no available information about the antifungal activity of chitosan against potato black scurf pathogen R. solani. This study aims to evaluate the antifungal activity and the induced effect of chitosan treatment on potato tuber resistance to R. solani under in vitro and in vivo conditions.

2. MATERIALS AND METHODS

2.1. Potato Tubers

Tubers of cultivars Sante and Kolobok were harvested from Moiseev farm, Bazarno-Karabulaksky District, Saratov region, Russia. The tubers based on size and showing no visible signs of disease infection or physical injuries were packed in net bags, transported to the laboratory, and stored at (16 ± 2) °C. Before treatment, the tubers were superficially disinfected with 2% sodium hypochlorite for 3 min, washed several times with sterilized water to wash out the remaining disinfectant solution and then air-dried.

2.2. Pathogen

Tubers with typical symptoms of black scurf were used to isolate the pathogen. Diseased tissue was cut into small pieces (1 cm2) and surface-disinfected in 70% ethanol for 30 s, followed by treatment with 2% (vol/vol) sodium hypochlorite for 3 min. Superficially sterilized pieces were rinsed several times in sterilized water to wash out the remaining disinfectant solution and cut aseptically into two pieces. These pieces were then dried between sterilized filter papers and then were placed on Petri dishes containing fresh Potato Dextrose Agar medium (PDA) (Difco Laboratories, St. Louis) supplemented with streptomycin sulphate 5 mg L-1. The plates were incubated at 24°C for 4 days, and single tips of fungal mycelium were transferred to sterile PDA for purification [20Papavizas G, Davey C. Isolation of Rhizoctonia solani Kuehn from naturally infested and artificially inoculated soils. Plant Dis Rep 1959; 43: 404-10.]. Several pure cultures of Rhizoctonia solani isolates were identified by light microscopy as described by Sherwood [21Sherwood R. Physiology of Rhizoctonia solani 1970.]. Rhizoctonia mycelia were stored at 4°C.

2.3. Chitosan

Chitosan, edible level with an average molecular weight of 150 kDa and degree of deacetylation 80%, was purchased from Chitosan Technologies Limited Company, Engels city, Russia.

2.4. Influence of Chitosan Treatment on Mycelial Growth of Rhizoctonia solani In Vitro

The effect of chitosan on mycelial growth was assessed by inoculating mycelial disks (5 mm in diameter) from the edge of a 4-day-old-culture of the fungus on the center of 90-mm Petri dishes containing 20 ml PDA medium amended with chitosan at (0, 0.125, 0.25, 0.5, or 1%). Plates were incubated in the dark at 25 °C and the mycelial growth was determined by measuring the colony diameter when the mycelium reached to the edges of the control plate. Each treatment was replicated using three plates, and the experiment was performed three times.

2.5. Influence of Chitosan Treatment On Lesion Diameters of Tubers Inoculated with Rhizoctonia Solani

Tubers were wounded (3 mm deep and 3 mm wide) at the equator with a sterile dissecting needle and then mycelial plugs of 4-day-old R. solani culture were inoculated into wounded sites with the hyphal side down. After 3 h of inoculation, tubers were treated with different concentrations of chitosan 0, 0.125, 0.25, 0.5, or 1% (w/v) dissolved in 0.5 mol L-1 glacial acetic acid, and the pH of CH solutions was adjusted to 5.5-6.0 with 1M aqueous NaOH. The treated tubers were incubated in plastic boxes (190 mm× 157 mm × 90 mm) with sterile water to maintain high relative humidity and stored at room temperature (20±2°C). The diameter of the lesions was measured after 30 d of inoculation. Each treatment was applied to three replicates of 15 tubers. The experiment was repeated twice.

2.6. Influence of Chitosan Treatment on Defense-Related Enzymes

Three grams of fresh weight were taken from 3-4 mm below the treated sides (Rhizoctonia with chitosan) and untreated ones (Rhizoctonia lacking chitosan) with a stainless-steel cork borer after 0, 1, 2, 3, 4, 5, 6, and 7 d of treatment with 0.5% chitosan and then were ground to a fine powder in liquid N2 and used for extraction of Peroxidase (POD), Polyphenol Oxidase (PPO) and Phenylalanine Ammonia-lyase (PAL), which are three defense-related enzymes in potato tubers. POD activity was assayed as described by Venisse et al. [22Venisse JS, Malnoy M, Faize M, Paulin JP, Brisset MN. Modulation of defense responses of Malus spp. during compatible and incompatible interactions with Erwinia amylovora. Mol Plant Microbe Interact 2002; 15(12): 1204-12. Available at: http://www.ncbi.nlm.nih.gov/ pubmed/12481992
[http://dx.doi.org/10.1094/MPMI.2002.15.12.1204] [PMID: 1248 1992]
] by measuring the absorbance at 470 nm. PPO activity was estimated according to Jiang et al. [23Aili J, Shiping T, Yong X. Effect of CA with high-O2 or high-CO2 concentrations on postharvest physiology and storability of sweet cherry. Acta Bot Sin 2002; 44(8): 925-30.] by measuring the absorbance at 420 nm. PAL activity was assayed according to the method of Assis et al. [24Assis JS, Maldonado R, Muñoz T. Escribano MaI, Merodio C. Effect of high carbon dioxide concentration on PAL activity and phenolic contents in ripening cherimoya fruit. Postharvest Biol Technol 2001; 23(1): 33-9.
[http://dx.doi.org/10.1016/S0925-5214(01)00100-4]
] by measuring the absorbance at 290 nm. The content of total protein was determined by the method of Bradford [25Bradford MM. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem 1976; 72(1-2): 248-54. [DOI].
[http://dx.doi.org/10.1016/0003-2697(76)90527-3] [PMID: 942051]
] using bovine serum albumin as standard.

2.7. Statistical Analysis

All statistical analyses were performed using the CoStat 6.45 software program. To test the effect of chitosan treatment, the data were Analyzed by One-way Analysis of Variance (ANOVA). Mean separations were performed by the least significant difference (LSD) test. Significance was defined as P< 0.05.

3. RESULTS

3.1. Influence of Chitosan Treatment on Mycelial Growth of Rhizoctonia solani In Vitro

Chitosan significantly inhibited the growth of R. solani mycelium in a concentration-dependent manner. The mycelial growth was completely inhibited by chitosan at 1% (Fig. 1).

3.2. Influence of Chitosan Treatment On Lesion Diameters of Tubers Inoculated with Rhizoctonia Solani

Treatment with chitosan significantly reduced the lesion diameter of R. solani AG-3 PT inoculated tubers after 30 d of incubation and the reduction enhanced with increasing the concentration. However, no significance was found between 0.5 and 1% concentrations in both cultivars (Fig. 2A, B). The CH treatment at 0.5% reduced the lesion diameter by 67.1 and 62.8%, respectively, in Kolobok and Cante cultivars.

Fig. (1)
Influence of chitosan concentrations on mycelial growth of Rhizoctonia solani. Bars represent standard deviations of the means. Values followed by different letters are significantly different according to LSD test at P < 0.05.


Fig. (2)
Influence of chitosan at different concentrations on lesion diameters of inoculated tubers of tow cultivars. A, Kolobok. B, Cante. Data are expressed as a mean of triplicate samples (± SD). Significant differences (P> 0.05) as indicated by LSD test are shown by different letters.


Fig. (3)
Influence of chitosan at 0.5% on the activity of defense-related enzymes POD (A, D), PPO (B, E) and PAL (C, F) in two cultivars (A, B, and C, Kolobok. D, E, and F, Cante). Data is presented as mean (± SD) based on three replicates.


3.3. Influence of Chitosan Treatment on Defense-Related Enzymes

POD was activated following treatment with CH 0.5% in both cultivars Kolobok and Cante. In Kolobok cultivar, a significant increase in POD activity was early observed after 1 day. This response was followed by a strong increase in the activity up to the 4th day (Fig. 3A). In contrast, this response was weaker and delayed in Cante cultivar, the first peak of POD activity appeared only after 3 days of treatment (Fig. 3D). The induction in POD activity reached its higher increase 4 and 5 days after treatment in Kolobok and Cante cultivars, respectively (Fig. 3A, D).

PPO activity pattern was higher in the treated tubers than the control in both cultivars. In Kolobok cultivar, PPO activity showed a different pattern between CH treated and non-treated tubers. Peaks were observed at 3 and 6 days in treated tubers (Fig. 3B). In Cante cultivar, the PPO activity followed the same trend and fluctuation in both CH-treated and untreated control tubers. However, the PPO activity showed a higher induction in CH-treated samples (range of 1.3-1.9 fold increase) than that in the untreated control samples (Fig. 3E).

PAL activity increased with incubation time in CH-treated tubers of both cultivars, reaching maximum values on 3 and 6 days in Kolobok and Cante cultivars, respectively (Fig. 3C, F).

4. DISCUSSION

As a natural elicitor and antifungal agent, chitosan is a promising alternative for the management of postharvest plant diseases [26Bautista-Baños S, Hernandez-Lauzardo AN, Velazquez-Del Valle MG, et al. Chitosan as a potential natural compound to control pre and postharvest diseases of horticultural commodities. Crop Prot 2006; 25(2): 108-18.
[http://dx.doi.org/10.1016/j.cropro.2005.03.010]
]. In our present study, chitosan was effective in inhibition R. solani mycelial growth. These observations confirmed similar data on the antifungal effect of chitosan on mycelial growth of R. solani [27Freddo ÁR, Mazaro SM, Brun EJ, Wagner Júnior A. Chitosan as fungistatic mycelial growth of Rhizoctonia solani Kuhn. Cienc Rural 2014; 44(1): 1-4.
[http://dx.doi.org/10.1590/S0103-84782014000100001]
, 28Hirano S, Nagao N. Effects of chitosan, pectic acid, lysozyme, and chitinase on the growth of several phytopathogens. Agric Biol Chem 1989; 53(11): 3065-6.], and other several phytopathogenic fungi, such as Fusarium solani [29Eweis M, Elkholy SS, Elsabee MZ. Antifungal efficacy of chitosan and its thiourea derivatives upon the growth of some sugar-beet pathogens. Int J Biol Macromol 2006; 38(1): 1-8.
[http://dx.doi.org/10.1016/j.ijbiomac.2005.12.009] [PMID: 16413607]
], Colletotrichum sp [30de Oliveira KAR, Berger LRR, de Araújo SA, Câmara MPS, de Souza EL. Synergistic mixtures of chitosan and Mentha piperita L. essential oil to inhibit Colletotrichum species and anthracnose development in mango cultivar Tommy Atkins. Food Microbiol 2017; 66: 96-103.http://www.ncbi.nlm.nih.gov/pubmed/28576378
[http://dx.doi.org/10.1016/j.fm.2017.04.012] [PMID: 28576378]
]. This recent study also showed that chitosan can effectively manage black scurf in potato tubers, by induction of defense-related enzymes such as POD, PPO. These results support the findings of CH-enhanced activities of POD and PPO against Sclerotinia sclerotiorum in carrots [31Qing W. ZUO J-h, Qian W, Yang N, GAO Lp. Inhibitory effect of chitosan on growth of the fungal phytopathogen, Sclerotinia sclerotiorum, and sclerotinia rot of carrot. J Integr Agric 2015; 14(4): 691-7.
[http://dx.doi.org/10.1016/S2095-3119(14)60800-5]
], Fusarium sulphureum in potato [32Yang B, LI YC, HAN RF, GE YH. Postharvest chitosan treatment induces resistance in potato against Fusarium sulphureum. Agric Sci China 2008; 7(5): 615-21.
[http://dx.doi.org/10.1016/S1671-2927(08)60060-7]
], and Alternaria kikuchiana and Physalospora piricola in pear [33Meng X, Yang L, Kennedy JF, Tian S. Effects of chitosan and oligochitosan on growth of two fungal pathogens and physiological properties in pear fruit. Carbohydr Polym 2010; 81(1): 70-5.
[http://dx.doi.org/10.1016/j.carbpol.2010.01.057]
]. POD participates in the cell wall building processes, for instance, suberization, phenols oxidation, and lignification of host plant cells during the defense reaction against pathogenic agents [34Sommer A, Ne’eman E, Steffens JC, Mayer AM, Harel E. Import, targeting, and processing of a plant polyphenol oxidase. Plant Physiol 1994; 105(4): 1301-11.
[http://dx.doi.org/10.1104/pp.105.4.1301] [PMID: 7972497]
]. PPO is involved in the oxidation of polyphenols into quinones (antimicrobial compounds) and cells lignification in the infected plants [35Kolattukudy P, Mohan R, Bajar MA, Sherf BA. Plant peroxidase gene expression and function 1992.
[http://dx.doi.org/10.1042/bst0200333]
]. In this experiment, the activity of PAL has been increased in CH-treated tubers. PAL is the first enzyme in the phenylpropanoid pathway and is involved in the syntheses of phytoalexins, phenols, and lignin which have defense functions in the host plants [36Pellegrini L, Rohfritsch O, Fritig B, Legrand M. Phenylalanine ammonia-lyase in tobacco. Molecular cloning and gene expression during the hypersensitive reaction to tobacco mosaic virus and the response to a fungal elicitor. Plant Physiol 1994; 106(3): 877-86.
[http://dx.doi.org/10.1104/pp.106.3.877] [PMID: 7824656]
].

CONCLUSION

This study indicated that chitosan directly inhibited the R. solani growth, and potentially elicited defense reaction in potato tubers.

ETHICS APPROVAL AND CONSENT TO PARTICIPATE

Not applicable.

HUMAN AND ANIMAL RIGHTS

No animals/humans were used for studies that are the basis of this research.

CONSENT FOR PUBLICATION

Not applicable.

AVAILABILITY OF DATA AND MATERIALS

Not applicable.

FUNDING

None.

CONFLICT OF INTEREST

The authors declare no conflict of interest, financial or otherwise.

ACKNOWLEDGEMENTS

The authors are grateful to the Biotechnology Lab., Agronomy Faculty, Saratov State Vavilov Agrarian University for the support and technical assistance of this research.

REFERENCES

[1] Banville GJ, Carling DE, Otrysko BE. Rhizoctonia disease on potato Rhizoctonia species: Taxonomy, molecular biology, ecology, pathology and disease control 1996; 321-30.
[http://dx.doi.org/10.1007/978-94-017-2901-7_29]
[2] Keiser A. Rhizoctonia solani a fungal disease with multiple symptoms: means of preventive and curative control. Potato Research for a Production of Quality Information Day 2008.
[3] El Bakali AM, Martín MP. Black scurf of potato. Mycologist 2006; 20(4): 130-2.
[http://dx.doi.org/10.1016/j.mycol.2006.03.006]
[4] Campion C, Chatot C, Perraton B, Andrivon D. Anastomosis groups, pathogenicity and sensitivity to fungicides of Rhizoctonia solani isolates collected on potato crops in France. Eur J Plant Pathol 2003; 109(9): 983-92.
[http://dx.doi.org/10.1023/B:EJPP.0000003829.83671.8f]
[5] Djébali N, Belhassen T. Field study of the relative susceptibility of eleven potato (Solanum tuberosum L.) varieties and the efficacy of two fungicides against Rhizoctonia solani attack. Crop Prot 2010; 29(9): 998-1002.
[http://dx.doi.org/10.1016/j.cropro.2010.06.012]
[6] Rabea EI, Badawy ME, Steurbaut W, Stevens CV. In vitro assessment of N-(benzyl) chitosan derivatives against some plant pathogenic bacteria and fungi. Eur Polym J 2009; 45(1): 237-45.
[http://dx.doi.org/10.1016/j.eurpolymj.2008.10.021]
[7] El Hadrami A, Adam LR, El Hadrami I, Daayf F. Chitosan in plant protection. Mar Drugs 2010; 8(4): 968-87. Available at: http://www.ncbi.nlm. nih.gov/pubmed/20479963
[http://dx.doi.org/10.3390/md8040968] [PMID: 20479963]
[8] Hamed I, Özogul F, Regenstein JM. Industrial applications of crustacean by-products (chitin, chitosan, and chitooligosaccharides): A review. Trends Food Sci Technol 2016; 48: 40-50.
[http://dx.doi.org/10.1016/j.tifs.2015.11.007]
[9] Choi C, Nam J-P, Nah J-W. Application of chitosan and chitosan derivatives as biomaterials. J Ind Eng Chem 2016; 33: 1-10.
[http://dx.doi.org/10.1016/j.jiec.2015.10.028]
[10] Malerba M, Cerana R. Chitosan Effects on Plant Systems. Int J Mol Sci 2016; 17(7): 996. Available at: http://www.ncbi.nlm.nih.gov/pubmed/27347928
[http://dx.doi.org/10.3390/ijms17070996] [PMID: 27347928]
[11] Du J, Gemma H, Iwahori S. Effects of chitosan coating on the storage of peach, Japanese pear, and kiwifruit. J Jpn Soc Hortic Sci 1997; 66(1): 15-22.
[http://dx.doi.org/10.2503/jjshs.66.15]
[12] Zhang D, Quantick PC. Antifungal effects of chitosan coating on fresh strawberries and raspberries during storage. J Hortic Sci Biotechnol 1998; 73(6): 763-7.
[http://dx.doi.org/10.1080/14620316.1998.11511045]
[13] de Capdeville G, Wilson CL, Beer SV, Aist JR. Alternative disease control agents induce resistance to blue mold in harvested ‘red delicious’ apple fruit. Phytopathology 2002; 92(8): 900-8.
[http://dx.doi.org/10.1094/PHYTO.2002.92.8.900] [PMID: 18942970]
[14] Liu J, Tian S, Meng X, Xu Y. Effects of chitosan on control of postharvest diseases and physiological responses of tomato fruit. Postharvest Biol Technol 2007; 44(3): 300-6.
[http://dx.doi.org/10.1016/j.postharvbio.2006.12.019]
[15] Meng X, Li B, Liu J, Tian S. Physiological responses and quality attributes of table grape fruit to chitosan preharvest spray and postharvest coating during storage. Food Chem 2008; 106(2): 501-8.
[http://dx.doi.org/10.1016/j.foodchem.2007.06.012] [PMID: 2604 9245]
[16] Abd-El-Kareem F, El-Mougy NS, El-Gamal NG, Fotouh Y. Use of chitin and chitosan against tomato root rot disease under greenhouse conditions. Res J Agric Biol Sci 2006; 2: 147-52.
[17] Rodríguez AF, Menéndez DC, Delgado EO, Díaz OL, Pino JC. Evaluation of chitosan as an inhibitor of soil-borne pathogens and as an elicitor of defence markers and resistance in tobacco plants. Span J Agric Res 2007; 5(4): 533-41.
[http://dx.doi.org/10.5424/sjar/2007054-274]
[18] Liu H, Tian W, Li B, et al. Antifungal effect and mechanism of chitosan against the rice sheath blight pathogen, Rhizoctonia solani. Biotechnol Lett 2012; 34(12): 2291-8. http://www.ncbi.nlm. nih.gov/pubmed/22932934
[http://dx.doi.org/10.1007/s10529-012-1035-z] [PMID: 22932934]
[19] Kurzawińska H. Potential use of chitosan in the control of lettuce pathogens. Polish Chitin Society. Monograph 2007; 12: 173-8.
[20] Papavizas G, Davey C. Isolation of Rhizoctonia solani Kuehn from naturally infested and artificially inoculated soils. Plant Dis Rep 1959; 43: 404-10.
[21] Sherwood R. Physiology of Rhizoctonia solani 1970.
[22] Venisse JS, Malnoy M, Faize M, Paulin JP, Brisset MN. Modulation of defense responses of Malus spp. during compatible and incompatible interactions with Erwinia amylovora. Mol Plant Microbe Interact 2002; 15(12): 1204-12. Available at: http://www.ncbi.nlm.nih.gov/ pubmed/12481992
[http://dx.doi.org/10.1094/MPMI.2002.15.12.1204] [PMID: 1248 1992]
[23] Aili J, Shiping T, Yong X. Effect of CA with high-O2 or high-CO2 concentrations on postharvest physiology and storability of sweet cherry. Acta Bot Sin 2002; 44(8): 925-30.
[24] Assis JS, Maldonado R, Muñoz T. Escribano MaI, Merodio C. Effect of high carbon dioxide concentration on PAL activity and phenolic contents in ripening cherimoya fruit. Postharvest Biol Technol 2001; 23(1): 33-9.
[http://dx.doi.org/10.1016/S0925-5214(01)00100-4]
[25] Bradford MM. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem 1976; 72(1-2): 248-54. [DOI].
[http://dx.doi.org/10.1016/0003-2697(76)90527-3] [PMID: 942051]
[26] Bautista-Baños S, Hernandez-Lauzardo AN, Velazquez-Del Valle MG, et al. Chitosan as a potential natural compound to control pre and postharvest diseases of horticultural commodities. Crop Prot 2006; 25(2): 108-18.
[http://dx.doi.org/10.1016/j.cropro.2005.03.010]
[27] Freddo ÁR, Mazaro SM, Brun EJ, Wagner Júnior A. Chitosan as fungistatic mycelial growth of Rhizoctonia solani Kuhn. Cienc Rural 2014; 44(1): 1-4.
[http://dx.doi.org/10.1590/S0103-84782014000100001]
[28] Hirano S, Nagao N. Effects of chitosan, pectic acid, lysozyme, and chitinase on the growth of several phytopathogens. Agric Biol Chem 1989; 53(11): 3065-6.
[29] Eweis M, Elkholy SS, Elsabee MZ. Antifungal efficacy of chitosan and its thiourea derivatives upon the growth of some sugar-beet pathogens. Int J Biol Macromol 2006; 38(1): 1-8.
[http://dx.doi.org/10.1016/j.ijbiomac.2005.12.009] [PMID: 16413607]
[30] de Oliveira KAR, Berger LRR, de Araújo SA, Câmara MPS, de Souza EL. Synergistic mixtures of chitosan and Mentha piperita L. essential oil to inhibit Colletotrichum species and anthracnose development in mango cultivar Tommy Atkins. Food Microbiol 2017; 66: 96-103.http://www.ncbi.nlm.nih.gov/pubmed/28576378
[http://dx.doi.org/10.1016/j.fm.2017.04.012] [PMID: 28576378]
[31] Qing W. ZUO J-h, Qian W, Yang N, GAO Lp. Inhibitory effect of chitosan on growth of the fungal phytopathogen, Sclerotinia sclerotiorum, and sclerotinia rot of carrot. J Integr Agric 2015; 14(4): 691-7.
[http://dx.doi.org/10.1016/S2095-3119(14)60800-5]
[32] Yang B, LI YC, HAN RF, GE YH. Postharvest chitosan treatment induces resistance in potato against Fusarium sulphureum. Agric Sci China 2008; 7(5): 615-21.
[http://dx.doi.org/10.1016/S1671-2927(08)60060-7]
[33] Meng X, Yang L, Kennedy JF, Tian S. Effects of chitosan and oligochitosan on growth of two fungal pathogens and physiological properties in pear fruit. Carbohydr Polym 2010; 81(1): 70-5.
[http://dx.doi.org/10.1016/j.carbpol.2010.01.057]
[34] Sommer A, Ne’eman E, Steffens JC, Mayer AM, Harel E. Import, targeting, and processing of a plant polyphenol oxidase. Plant Physiol 1994; 105(4): 1301-11.
[http://dx.doi.org/10.1104/pp.105.4.1301] [PMID: 7972497]
[35] Kolattukudy P, Mohan R, Bajar MA, Sherf BA. Plant peroxidase gene expression and function 1992.
[http://dx.doi.org/10.1042/bst0200333]
[36] Pellegrini L, Rohfritsch O, Fritig B, Legrand M. Phenylalanine ammonia-lyase in tobacco. Molecular cloning and gene expression during the hypersensitive reaction to tobacco mosaic virus and the response to a fungal elicitor. Plant Physiol 1994; 106(3): 877-86.
[http://dx.doi.org/10.1104/pp.106.3.877] [PMID: 7824656]

Endorsements



"Open access will revolutionize 21st century knowledge work and accelerate the diffusion of ideas and evidence that support just in time learning and the evolution of thinking in a number of disciplines."


Daniel Pesut
(Indiana University School of Nursing, USA)

"It is important that students and researchers from all over the world can have easy access to relevant, high-standard and timely scientific information. This is exactly what Open Access Journals provide and this is the reason why I support this endeavor."


Jacques Descotes
(Centre Antipoison-Centre de Pharmacovigilance, France)

"Publishing research articles is the key for future scientific progress. Open Access publishing is therefore of utmost importance for wider dissemination of information, and will help serving the best interest of the scientific community."


Patrice Talaga
(UCB S.A., Belgium)

"Open access journals are a novel concept in the medical literature. They offer accessible information to a wide variety of individuals, including physicians, medical students, clinical investigators, and the general public. They are an outstanding source of medical and scientific information."


Jeffrey M. Weinberg
(St. Luke's-Roosevelt Hospital Center, USA)

"Open access journals are extremely useful for graduate students, investigators and all other interested persons to read important scientific articles and subscribe scientific journals. Indeed, the research articles span a wide range of area and of high quality. This is specially a must for researchers belonging to institutions with limited library facility and funding to subscribe scientific journals."


Debomoy K. Lahiri
(Indiana University School of Medicine, USA)

"Open access journals represent a major break-through in publishing. They provide easy access to the latest research on a wide variety of issues. Relevant and timely articles are made available in a fraction of the time taken by more conventional publishers. Articles are of uniformly high quality and written by the world's leading authorities."


Robert Looney
(Naval Postgraduate School, USA)

"Open access journals have transformed the way scientific data is published and disseminated: particularly, whilst ensuring a high quality standard and transparency in the editorial process, they have increased the access to the scientific literature by those researchers that have limited library support or that are working on small budgets."


Richard Reithinger
(Westat, USA)

"Not only do open access journals greatly improve the access to high quality information for scientists in the developing world, it also provides extra exposure for our papers."


J. Ferwerda
(University of Oxford, UK)

"Open Access 'Chemistry' Journals allow the dissemination of knowledge at your finger tips without paying for the scientific content."


Sean L. Kitson
(Almac Sciences, Northern Ireland)

"In principle, all scientific journals should have open access, as should be science itself. Open access journals are very helpful for students, researchers and the general public including people from institutions which do not have library or cannot afford to subscribe scientific journals. The articles are high standard and cover a wide area."


Hubert Wolterbeek
(Delft University of Technology, The Netherlands)

"The widest possible diffusion of information is critical for the advancement of science. In this perspective, open access journals are instrumental in fostering researches and achievements."


Alessandro Laviano
(Sapienza - University of Rome, Italy)

"Open access journals are very useful for all scientists as they can have quick information in the different fields of science."


Philippe Hernigou
(Paris University, France)

"There are many scientists who can not afford the rather expensive subscriptions to scientific journals. Open access journals offer a good alternative for free access to good quality scientific information."


Fidel Toldrá
(Instituto de Agroquimica y Tecnologia de Alimentos, Spain)

"Open access journals have become a fundamental tool for students, researchers, patients and the general public. Many people from institutions which do not have library or cannot afford to subscribe scientific journals benefit of them on a daily basis. The articles are among the best and cover most scientific areas."


M. Bendandi
(University Clinic of Navarre, Spain)

"These journals provide researchers with a platform for rapid, open access scientific communication. The articles are of high quality and broad scope."


Peter Chiba
(University of Vienna, Austria)

"Open access journals are probably one of the most important contributions to promote and diffuse science worldwide."


Jaime Sampaio
(University of Trás-os-Montes e Alto Douro, Portugal)

"Open access journals make up a new and rather revolutionary way to scientific publication. This option opens several quite interesting possibilities to disseminate openly and freely new knowledge and even to facilitate interpersonal communication among scientists."


Eduardo A. Castro
(INIFTA, Argentina)

"Open access journals are freely available online throughout the world, for you to read, download, copy, distribute, and use. The articles published in the open access journals are high quality and cover a wide range of fields."


Kenji Hashimoto
(Chiba University, Japan)

"Open Access journals offer an innovative and efficient way of publication for academics and professionals in a wide range of disciplines. The papers published are of high quality after rigorous peer review and they are Indexed in: major international databases. I read Open Access journals to keep abreast of the recent development in my field of study."


Daniel Shek
(Chinese University of Hong Kong, Hong Kong)

"It is a modern trend for publishers to establish open access journals. Researchers, faculty members, and students will be greatly benefited by the new journals of Bentham Science Publishers Ltd. in this category."


Jih Ru Hwu
(National Central University, Taiwan)


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