Classical and augmentative biological control against ... - IOBC-WPRS

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Ruocco et al. Ismaiel 2009). The mechanisms that Trichoderma uses to antagonize phytopathogenic fungi include competition, colonization, antibiosis and direct mycoparasitism (Harman 2006, 2011; Howell 2003). This antagonistic potential serves as the basis for effective biological control applications of different Trichoderma strains as an alternative method to chemicals for the control of a wide spectrum of plant pathogens (Harman et al. 1991; Lorito et al. 2010). The colonization of the root system by rhizosphere competent strains of Trichoderma results in increased development of root and/or aerial systems and crop yields (Bae et al. 2011; Chacon et al. 2007; Kubicek et al. 1998; Yedidia et al. 2003). Trichoderma has also been described as being involved in other biological activities such as the induction of plant systemic resistance (Shoresh et al. 2010; Tucci et al. 2011) and antagonistic effects on plant pathogenic nematodes (Jegathambigai et al. 2008; Sharon et al. 2001).Some strains of Trichoderma have also been noted to be aggressive biodegraders in their saprophytic phases, in addition to acting as competitors to fungal pathogens, particularly when nutrients are a limiting factor in the environment (Worasatit et al. 1994). These facts strongly suggest that in the plant root environment Trichoderma actively interacts with the components in the soil community, the plant, bacteria, fungi, other organisms, such as nematodes or insects, that share the same ecological niche (Lorito et al. 2010). Trichoderma spp. are important participants in the nutrient cycle. They aid in the decomposition of organic matter and make available to the plant many elements normally inaccessible. Yedidia et al. (2001) noted that the presence of the fungus increased the uptake and concentration of a variety of nutrients (copper, phosphorus, iron, manganese and sodium) in the roots of plants grown in hydroponic culture, even under axenic conditions. These increased concentrations indicated an improvement in plant active-uptake mechanisms. Corn that developed from seeds treated with T. harzianum strain T-22 produced higher yields, even when a fertilizer containing 40% less nitrogen was applied, than the plants developed from seed that was not treated with T-22 (Harman 2000). This ability to enhance production with less nitrate fertilizer, provides the opportunity to potentially reduce nitrate pollution of ground and surface water, a serious adverse consequence of large-scale maize culture. In addition to effects on the increase of nutrient uptake and the efficiency of nitrogen use, the beneficial fungi can also solubilize various nutrients in the soil, that would be otherwise unavailable for uptake by the plant (Altomare et al. 1999b). The cross-talk that occurs between the fungal BCA and the plant is important both for identification of each component to one another and for obtaining beneficial effects. Somehow, the plant is able to sense, possibly by detection of the released fungal compounds, that Trichoderma is not a hostile presence, therefore the plant defence system is not activated as it is when there is pest attack and the BCA is recognized as a plant symbiont rather than a plant pathogen (Woo and Lorito, 2006). Molecules produced by Trichoderma and/or its metabolic activity also have potential for promoting plant growth (Chacón et al., 2007; Vinale et al. 2008a; 2008b; Yedidia et al. 1999). Applications of T. harzianum to seed or the plant resulted in improved germination, increased plant size, augmented leaf area and weight, greater yields (Altomare et al. 1999a; Harman et al. 2004c, b; Inbar and Chet 1995; Tucci et al. 2011; Vinale et al. 2008a). Numerous studies indicated that metabolic changes occur in the root during colonization by Trichoderma spp., such as the activation of pathogenesis-related proteins (PR-proteins), which induce in the plant an increased resistance to subsequent attack by numerous microbial pathogens (Table 12) 46
Chapter 6 Table 12: Evidence for, and effectiveness of, induced resistance in plants by Trichoderma species (Harman et al., 2004a). The induction of systemic resistance (ISR) observed in planta determines an improved control of different classes of pathogens (mainly fungi and bacteria), which are spatially and temporally distant from the Trichoderma inoculation site. This phenomenon has been observed in many plant species, both dicotyledons (tomato, pepper, tobacco, cotton, bean, cucumber) and monocotyledions (corn, rice). For example, Trichoderma induces resistance towards Botrytis cinerea in tomato, tobacco, lettuce, pepper and bean plants, with a symptom reduction ranging from 25 to 100% (Tucci et al. 2011). Moreover, Trichoderma determined an overall increased production of defence-related plant enzymes, including various peroxidases, chitinases, β-1,3-glucanases, and the lipoxygenasepathway hydroperoxide lyase (Harman et al. 2004c; Howell et al. 2000; Yedidia et al. 1999) of T. harzianum strain T-39, the active ingredient of the commercial product TricodexTM. Thus far, Trichoderma is able not only to produce toxic compounds with a direct antimicrobial activity against pathogens, but also to generate fungal substances that are able to stimulate the plant to produce its own defence metabolites. In fact, the ability of T. virens to induce phytoalexin accumulation and localized resistance in cotton has already been discussed (Hanson and Howell 2004). In cucumber, root colonization by strain T-203 of T. asperellum caused an increase in phenolic glucoside levels in the leaves; the aglycones, which are phenolic glucosides with the carbohydrate moieties removed, are strongly inhibitory to a range of bacteria and fungi (Yedidia et al. 2003). 47
Page 1 and 2:
IOBC OILB WPRS SROP In te rn atio n
Page 3 and 4:
Preface One of the Research Activit
Page 5 and 6: LORITO Matteo UNINA, Dip. Arboricol
Page 7 and 8: List of Tables (continued) Table 17
Page 9 and 10: Contents Chapter 1 Potential of bio
Page 11 and 12: Chapter 1 Potential of biological c
Page 13 and 14: Chapter 1 To allow for the analysis
Page 15 and 16: Chapter 1 Table 4: Microbial specie
Page 17 and 18: Chapter 1 Table 4 (continued) T. lo
Page 19 and 20: Chapter 1 Table 4 (continued) Cupri
Page 21 and 22: Chapter 1 other was aiming at ident
Page 23 and 24: Chapter 2 Table 5: References extra
Page 25 and 26: Chapter 2 Table 6: Biocontrol agent
Page 27 and 28: Table 7 (continued) Chapter 2 Hemip
Page 29 and 30: Chapter 2 Bacillus thuringiensis: 2
Page 31 and 32: Chapter 3 - financial; since the de
Page 33 and 34: Chapter 3 Pathogens and Nematodes a
Page 35 and 36: Chapter 3 [Remark: Estimating the s
Page 37 and 38: Chapter 3 100% 80% 60% P = Phytopha
Page 39 and 40: Chapter 3 0,25 4 0,20 % dedicated t
Page 41 and 42: Chapter 3 Table 9: Recent introduct
Page 43 and 44: Chapter 3 Table 9: Recent introduct
Page 45 and 46: Chapter 4 The survey was limited to
Page 47 and 48: Chapter 4 Table 11 (continued) Othe
Page 49 and 50: Chapter 4 SCLPs are included in ann
Page 51 and 52: Chapter 4 laminarin is available on
Page 53 and 54: Chapter 5 (EC) No 1109/2009. Tests
Page 55: Chapter 6 Identified difficulties a
Page 59 and 60: Table 13: Trichoderma-based prepara
Page 61 and 62: Chapter 6 Many preparations have be
Page 63 and 64: Chapter 6 Persistence, physiologica
Page 65 and 66: Chapter 6 Harman, G. E. (2006) Over
Page 67 and 68: Chapter 6 Sharon, E., Bar-Eyal, M.,
Page 69 and 70: Chapter 7 purification, formulation
Page 71 and 72: Chapter 7 Business profitability Co
Page 73 and 74: Chapter 8 A survey was carried out
Page 75 and 76: Chapter 8 80 70 60 %of supply %of a
Page 77 and 78: Chapter 8 efficacy and the price of
Page 79 and 80: Conclusions and perspectives reduct
Page 81 and 82: Appendices For Chapter 1 Appendix 1
Page 83 and 84: Appendix 1 B O Milsana + Brevibacil
Page 85 and 86: Appendix 1 Strawberry (target patho
Page 87 and 88: Appendix 1 Fruits - postharvest (ap
Page 89 and 90: Appendix 1 Legumes (Fabaceae) (targ
Page 91 and 92: Appendix 1 Miscellaneous crops (tar
Page 93 and 94: Appendix 1 Lecanicillium muscarium
Page 95 and 96: Appendix 1 Bedini, S., Bagnoli, G.,
Page 97 and 98: Appendix 1 Dik, A., and Wubben, J.
Page 99 and 100: Appendix 1 Gerlagh, M., Amsing, J.
Page 101 and 102: Appendix 1 Jing, W., Tu, K., Shao,
Page 103 and 104: Appendix 1 Li, G. Q., Huang, H. C.,
Page 105 and 106: Appendix 1 Morandi, M. A. B., Maffi
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Appendix 1 Rabea, E. I., Badawy, M.
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Appendix 1 Sutton, J. C., Liu, W.,
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Appendix 1 Zahavi, T., Cohen, L., W
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Appendix 2 Powdery mildew on grapes
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Appendix 2 Powdery mildew on cucurb
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Appendix 2 Askary, H. (1998). Patho
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Appendix 2 Malathrakis, N. E. (2002
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Appendix 3. Inventory of biocontrol
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Appendix 3 References on biocontrol
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Appendix 4. Inventory of biocontrol
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Appendix 4 Grapes (target pathogen
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Appendix 4 References on biocontrol
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Appendix 4 Robak, J., and Ostrowska
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Appendix 5 Cherry (target pathogens
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Appendix 5 Successful inhibition in
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Appendix 5 Smilanick, J. L., Denis-
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Appendix 6 showed protein content o
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Appendix 6 El-Khallal, S. M. (2007)
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Appendix 6 Liu, Q., J. C. Yu, et al
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Appendix 6 of tomato wilt pathogen
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Appendix 6 incognita, respectively.
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Appendix 7. Number of references re
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Appendix 8. Collection of data on a
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Appendix 8 Camporese & Duso, 1996 T
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Appendix 8 8.5 Fungi [5 species] Re
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Appendix 8 Berner & Schnetter, 2002
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Appendix 8 Morandi-Filho et al., 20
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Appendix 8 Marshall D.B. & Lester P
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Appendix 9 9.2. Details on the use
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Appendix 9 References 1. Abd-Rabou
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Appendix 9 50. Gariepy TD, Kuhlmann
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Appendix 9 97. Langewald J, Neuensc
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Appendix 9 146. Roltsch WJ, Meyerdi
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Appendix 10. Substances included in
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Appendix 10 Chemical Dimoxystrobin
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Appendix 10 Chemical Prosulfocarb '
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Appendix 10 Natural other Abamectin
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Appendix 11. Invertebrate beneficia
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Appendix 11 11.2. Invertebrate bioc
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Appendix 11 Coccinella septempuncta
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Appendix 11 Trichogramma brassicae
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Page 191 and 192:
Page 193 and 194:
Appendix 11 Diglyphus isaea Digline
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