A Review Study on the Postharvest Decay Control of Fruit by *Trichoderma*

*Ramsés González-Estrada, Francisco Blancas-Benítez, Beatriz Montaño-Leyva, Cristina Moreno-Hernández, Luz Del Carmen Romero-Islas, Jovita Romero-Islas, Rosa Avila-Peña, Anelsy Ramos-Guerrero, Angel Fonseca-Cantabrana and Porfirio Gutierrez-Martinez*

### **Abstract**

This chapter consists of an overview with the most relevant results about the efficacy of *Trichoderma* on postharvest disease control. The results of investigations demonstrate that this fungus can control several phytopathogens in different fruits. Postharvest losses represent a major problem in several countries. The constant application of fungicides not only at field but also at postharvest stage has led to microbial resistance cases, which make the control of these pathogens difficult. Biological control is a promising alternative to chemical fungicide applications. In this sense, an eco-friendly alternative and effective approach for controlling diseases is the use of microbial antagonists like *Trichoderma*, which have several mechanisms of action to stop disease development. A crucial treat in biological control is related to the maintenance of microbial viability and efficacy, that is why other technologies like their incorporation into edible films and coatings, nanotechnology, microbial mixtures, among others have been applied in combination with *Trichoderma* successfully. An enhancement in biocontrol activity is achieved when alternative systems are combined like GRAS substances, biopolymers, and other antagonists. Thus, *Trichoderma* is an eco-friendly alternative to threat postharvest diseases as an alternative to chemical treatments.

**Keywords:** *Trichoderma*, postharvest, pathogens, fruits, alternative systems

### **1. Introduction**

Postharvest diseases represent a major problem through the world causing significant losses at postharvest stage [1]. Postharvest treatments play an important role in the quality preservation of commodities; however, in developed countries, the inadequate storage and transportation systems favor the establishment of diverse pathogens [2]. Traditionally, postharvest disease management is carried out by the application of chemical fungicides; however, environmental and health issues as well as microbial resistance play an important role in the development

of new strategies for controlling diseases [3]. Biological control is an eco-friendly alternative for postharvest disease control. Antagonists can be isolated from diverse sources like soil, fruits, leaves, and from extreme conditions as marine environments [2, 4]. *Trichoderma* is recognized due to their effectiveness in controlling several pathogens in diverse fruits like strawberry (*Botrytis cinerea*), citrus (*Penicillium italicum*), kiwifruit (*Botrytis cinerea*), banana (*Colletotrichum musae*), guava (*Rhizopus* spp.), among others [2]. The efficacy of *Trichoderma* is related to several mechanism of action reported like competition, antibiosis, parasitism (involving lytic enzymes), and the induction of plant defenses [5–7]. Biocontrol activity of *Trichoderma* can be enhanced by the combination of this antagonist with other control systems like the use of GRAS substances, encapsulation in polymeric matrices (chitosan), physical methods, etc. The aim of this chapter was to summarize information about the efficacy and application of *Trichoderma* alone or in combination with other alternative methods in different fruits against the most important postharvest pathogens.

### **2. Fruits: importance at international level**

In recent years, the demand for food has increased dramatically, while the world population has increased by 70%, food consumption per capita has increased more than 20%. According to some reports, it is projected that the production of crops for the year 2030 is 70% higher than the productions that are currently available [8]. Fruits and vegetables will play an important role by providing essential nutrients for the population's diet, both in developed and developing countries, in addition to being associated with the reduction of the risk of suffering from different chronic-degenerative diseases [9]. The United States dominates the international fruit and vegetable trade market and is the first place in terms of imports and exports of food of plant origin, while the European Union, as a whole, is the second importer and exporter of this type of food. On the other hand, some Latin American countries, such as Chile, have become one of the main suppliers in the international fresh fruit market. The increase in the production and commercialization of fruits and vegetables has been increasing, reaching a global production of nearly 1 billion tons of fruits and vegetables [10]. Mexico ranks sixth in the fruit-producing countries, along with China, India, Brazil, the United States, and Italy. China, India, and Brazil concentrate about 30% of the total production of fruits worldwide; however, much of this production is destined for local consumption, so the impact on the world market is minimal [10, 11]. The low international trade attends to the lack of fruit conservation mechanisms, which can suffer damage during the transfer. This implies significant postharvest losses, thus reducing the possibilities of export and international marketing of a large number of vegetable crops.

### **3. Postharvest losses: causes and consequences**

Currently, the losses of food in the world are about 1300 million tons per year; of which, in Latin America, the loss is, on average, 127 million tons of food per year; and with respect to fruits and vegetables, their loss is up to 55% in Latin America [12]. During the postharvest handling of fruits and vegetables, the losses range from 25 to 60% of the total production. This is due to several factors, but one of the most important factors is the poor handling of the products, where mechanical damage and diseases caused by pathogens play an important role [13]. Nowadays, the application of chemical fungicides is widely used as a strategy to control postharvest pathogens; however, even when they are effective,

**65**

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma*

available products and affect the financial benefits [15].

**4. Alternative methods for controlling postharvest diseases**

(*P. expansum*), table grapes (*Botrytis cinerea*), among others [16].

Heat treatments are one of the physical methods that have been used most for the conservation of minimally processed postharvest fruits, either alone or combined with other eco-friendly alternatives [15]. Heat treatments have been applied in several fruits against different pathogens with good results, like peach (*Monilinia fructicola*), apple (*Penicillium expansum*), grapefruit (*P. digitatum*), sweet cherry

The use of essential oils to control postharvest diseases is gaining popularity due to its safety features, biodegradability, and that are eco-friendly compounds [17]. For a long time, it has been recognized that some essential oils have antimicrobial, antiviral, antifungal, antiparasitic, and insecticidal properties [17]. Besides, in order to protect them and favoring their efficacy for controlling postharvest pathogens, the essential oils can be combined with edible films and coating, as previously

The use of edible films and coatings is an alternative to the use of fungicides to preserve the postharvest quality of fruits and vegetables [20]. The application of coatings and edible films in foods is mainly in perishable products, such as horticultural products, due to their properties such as cost, availability, functional attributes, mechanical properties (flexibility, tension), optical properties (brightness and opacity), the barrier effect against the flow of gases, structural resistance to water and microorganisms, as well as sensory acceptability [21]. Besides, edible films and coatings have a high potential to incorporate active ingredients such as antibrowning agents, colorants, flavors, nutrients, spices, antimicrobial compounds, and antagonists; which may favor extending shelf life of the product and reducing the risk of pathogen growth [22]. In this sense, incorporation of different additives into polymeric matrices in several fruits coated has been evaluated successfully in controlling

There is an international tendency to reduce the use of fungicides in fruit and vegetable products and to develop safer alternatives that reduce postharvest deterioration in fruits [26–28]. One option is the development of alternative control

postharvest diseases and maintaining the fruit quality [18, 19, 23–25].

**5. Biological control at postharvest stage**

the environmental impact of these methods is negative since they are toxic. On the other hand, there are several reports about the adaptation of pathogens (resistance) that reduce the antifungal activity and difficult their control; thus, it is necessary to investigate new, secure, and effective alternatives for controlling postharvest diseases [13, 14]. Fungal contamination can occur during the handling of fruits and vegetables at field, postharvest handling, storage, and transport and cause the deterioration of products, and as a consequence, decrease the amount of

*DOI: http://dx.doi.org/10.5772/intechopen.82784*

**4.1 Physical treatments**

**4.2 Essential oils**

reported with good results [17–19].

**4.3 Edible films and coatings**

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

the environmental impact of these methods is negative since they are toxic. On the other hand, there are several reports about the adaptation of pathogens (resistance) that reduce the antifungal activity and difficult their control; thus, it is necessary to investigate new, secure, and effective alternatives for controlling postharvest diseases [13, 14]. Fungal contamination can occur during the handling of fruits and vegetables at field, postharvest handling, storage, and transport and cause the deterioration of products, and as a consequence, decrease the amount of available products and affect the financial benefits [15].

### **4. Alternative methods for controlling postharvest diseases**

### **4.1 Physical treatments**

*Trichoderma - The Most Widely Used Fungicide*

important postharvest pathogens.

**2. Fruits: importance at international level**

of new strategies for controlling diseases [3]. Biological control is an eco-friendly alternative for postharvest disease control. Antagonists can be isolated from diverse sources like soil, fruits, leaves, and from extreme conditions as marine environments [2, 4]. *Trichoderma* is recognized due to their effectiveness in controlling several pathogens in diverse fruits like strawberry (*Botrytis cinerea*), citrus (*Penicillium italicum*), kiwifruit (*Botrytis cinerea*), banana (*Colletotrichum musae*), guava (*Rhizopus* spp.), among others [2]. The efficacy of *Trichoderma* is related to several mechanism of action reported like competition, antibiosis, parasitism (involving lytic enzymes), and the induction of plant defenses [5–7]. Biocontrol activity of *Trichoderma* can be enhanced by the combination of this antagonist with other control systems like the use of GRAS substances, encapsulation in polymeric matrices (chitosan), physical methods, etc. The aim of this chapter was to summarize information about the efficacy and application of *Trichoderma* alone or in combination with other alternative methods in different fruits against the most

In recent years, the demand for food has increased dramatically, while the world population has increased by 70%, food consumption per capita has increased more than 20%. According to some reports, it is projected that the production of crops for the year 2030 is 70% higher than the productions that are currently available [8]. Fruits and vegetables will play an important role by providing essential nutrients for the population's diet, both in developed and developing countries, in addition to being associated with the reduction of the risk of suffering from different chronic-degenerative diseases [9]. The United States dominates the international fruit and vegetable trade market and is the first place in terms of imports and exports of food of plant origin, while the European Union, as a whole, is the second importer and exporter of this type of food. On the other hand, some Latin American countries, such as Chile, have become one of the main suppliers in the international fresh fruit market. The increase in the production and commercialization of fruits and vegetables has been increasing, reaching a global production of nearly 1 billion tons of fruits and vegetables [10]. Mexico ranks sixth in the fruit-producing countries, along with China, India, Brazil, the United States, and Italy. China, India, and Brazil concentrate about 30% of the total production of fruits worldwide; however, much of this production is destined for local consumption, so the impact on the world market is minimal [10, 11]. The low international trade attends to the lack of fruit conservation mechanisms, which can suffer damage during the transfer. This implies significant postharvest losses, thus reducing the possibilities of

export and international marketing of a large number of vegetable crops.

Currently, the losses of food in the world are about 1300 million tons per year; of which, in Latin America, the loss is, on average, 127 million tons of food per year; and with respect to fruits and vegetables, their loss is up to 55% in Latin America [12]. During the postharvest handling of fruits and vegetables, the losses range from 25 to 60% of the total production. This is due to several factors, but one of the most important factors is the poor handling of the products, where mechanical damage and diseases caused by pathogens play an important role [13]. Nowadays, the application of chemical fungicides is widely used as a strategy to control postharvest pathogens; however, even when they are effective,

**3. Postharvest losses: causes and consequences**

**64**

Heat treatments are one of the physical methods that have been used most for the conservation of minimally processed postharvest fruits, either alone or combined with other eco-friendly alternatives [15]. Heat treatments have been applied in several fruits against different pathogens with good results, like peach (*Monilinia fructicola*), apple (*Penicillium expansum*), grapefruit (*P. digitatum*), sweet cherry (*P. expansum*), table grapes (*Botrytis cinerea*), among others [16].

### **4.2 Essential oils**

The use of essential oils to control postharvest diseases is gaining popularity due to its safety features, biodegradability, and that are eco-friendly compounds [17]. For a long time, it has been recognized that some essential oils have antimicrobial, antiviral, antifungal, antiparasitic, and insecticidal properties [17]. Besides, in order to protect them and favoring their efficacy for controlling postharvest pathogens, the essential oils can be combined with edible films and coating, as previously reported with good results [17–19].

### **4.3 Edible films and coatings**

The use of edible films and coatings is an alternative to the use of fungicides to preserve the postharvest quality of fruits and vegetables [20]. The application of coatings and edible films in foods is mainly in perishable products, such as horticultural products, due to their properties such as cost, availability, functional attributes, mechanical properties (flexibility, tension), optical properties (brightness and opacity), the barrier effect against the flow of gases, structural resistance to water and microorganisms, as well as sensory acceptability [21]. Besides, edible films and coatings have a high potential to incorporate active ingredients such as antibrowning agents, colorants, flavors, nutrients, spices, antimicrobial compounds, and antagonists; which may favor extending shelf life of the product and reducing the risk of pathogen growth [22]. In this sense, incorporation of different additives into polymeric matrices in several fruits coated has been evaluated successfully in controlling postharvest diseases and maintaining the fruit quality [18, 19, 23–25].

### **5. Biological control at postharvest stage**

There is an international tendency to reduce the use of fungicides in fruit and vegetable products and to develop safer alternatives that reduce postharvest deterioration in fruits [26–28]. One option is the development of alternative control

systems for controlling postharvest pathogens on fruits: as physical treatments (hot water), application of substance of vegetable and animal origin, or the use of antagonistic microorganisms [2, 29, 30]. Biological control is a promising alternative with high application potential for controlling postharvest pathogens. Some of the advantages of biological control compared to the use of fungicides are the absence of toxic waste in the fruits, friendly relationship with the environment, as well as in safety and health for the people who handle the products [14]. For several decades, a large number of microorganisms with potential characteristics of antagonistic organisms have been isolated from different sources (leaves, fruits, marine environment) [4, 31]. Bacteria, yeasts, and fungi are the main antagonistic organisms that have been used at postharvest stage. Countries such as the United States, South Africa, Spain, Italy, and Israel, among others, have developed and, in some cases, commercialized antagonistic microorganisms for their commercial use in postharvest, for the control of different diseases caused by the main pathogenic fungi of fruits of temperate, tropical, and subtropical origin [27, 32, 33]. The most widely used antagonists have an outstanding capacity for rapid growth and colonization of the fruit, nutrient acquisition, ability to withstand extreme temperature conditions, solar radiation, control capacity at low concentrations, as well as genetic stability and development capacity in economic cultivation means, ease of application in the management of postharvest. The main mechanisms of action present in the antagonists are antibiosis, production of lytic enzymes, competence by nutrients and space, or the induction of resistance mechanisms [1, 34].

There is a large number of reports of *in vitro* and *in vivo* tests, using bacteria such as *Pseudomonas cepacia* Van Hall, *Pseudomonas syringae*, *Bacillus subtilis*; as well as yeasts such as *Candida sake*, *Rhodotorula glutinis, Debaryomyces hansenii* for the control of the main pathogenic fruit fungi in postharvest, such as *Penicillium digitatum*, *Penicillium italicum, Alternaria, Aspergillus, Botrytis, Fusarium, Geotrichum, Gloeosporium, Monilinia, Penicillium, Mucor, Colletotrichum*, and *Rhizopus* [4, 27, 32, 33, 35]. In addition to bacteria and yeasts used in the biological control postharvest pathogens, the application of *Trichoderma*, alone or in combination with other alternative control systems in different fruits for the control of pathogenic fungi of fruits in the fruit, has been evaluated with favorable results [36, 37].

### **6.** *Trichoderma***: application on fruits**

*Trichoderma* has been applied as postharvest biocontrol agent in different crops such as papayas, strawberries, tomatoes, apples, pears, and bananas (**Table 1**). Existing different species of *Trichoderma* with high antagonistic capacity are *T. asperellum*, *T. viride*, and *T. harzianum.* Several authors have investigated different species of *Trichoderma* with the objective to find the most effective biocontrol agent for each crop and pathogen. The efficacy of six species of *Trichoderma* was evaluated on papaya fruit reducing the diameter lesions and disease incidence caused by *Colletotrichum gloeosporioides* by the application of *T. asperellum* and *T. viride* [6]. In a similar study on mango fruit, good efficacy was reported for controlling anthracnose applying *T. harzianum* obtaining a lower disease incidence (41.7% disease incidence) in fruits compared to control [38]. At present, *Trichoderma* is produced at industrial level as active component of biological products "biopesticides"; other ingredients that conform the biopesticides are the edible polymers, which can form coating for easy adhesion to the fruit and give to the product protection and stability during its shelf life. The application of biopesticides is widely used in agriculture and can be applied by immersion or spraying during the industrialization of

**67**

**Table 1.**

Trichoderma*.*

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma*

*theobromae*

*stolonifer*

*cinerea*

*expansum*

*expansum*

*cinerea*

*expansum*

*cinerea*

*expansum*

*cinerea*

*gloeosporioides*

*Percentage of inhibition of different postharvest diseases by the application of different species of* 

Blue mold *Penicillium* 

Blue mold *Penicillium* 

*gloeosporioides*

Gray mold *Botrytis* 

Blue mold *Penicillium* 

**Disease Pathogen** *Trichoderma* **species Disease** 

cepa T22

*T. harzianum* Rifai cepa T67

*Trichoderma* spp*.* cepa 8009

*T. reesei* cepa T34 71.8

**inhibition (%)**

5.3

33.3

72.5

70.6

59.3

40.6

28.7

47.6

*T. longibrachiatum* 65.0 Valenzuela et al. [6] *T. viride* 77.3

*T. harzianum* 96.9 Dal Bello

*T. harzianum* 50.5 Batta [40]

*T. viride* 60.7 Mortuza

*T. harzianum* 62.9 Batta [36]

*T. harzianum* 60 Prabakar

*T. atroviride* cepa P1 70.6 Quaglia et al. [44] *T. harzianum* Rifai

[43] *T. harzianum* 56.2

**Reference**

and Ilag

et al. [38]

et al. [45]

agricultural products [39]. The incorporation of *T. harzianum* into edible coatings as biopesticide produced higher inhibition of the pathogens *Botrytis cinerea* and *Penicillium expansum* compared to the application as simple conidial suspension of the antagonist on the fruits [40]. The same author previously reported the same effect on other fruits such as pears, grapes, apples, strawberries, kiwis, and peaches [36, 41]. Microbial antagonists not only have antifungal properties, but also can act as inductor agents; some studies report that their application can induce biochemical defense responses by the interaction antagonist-fruit. In this sense, the application of *T. virens* decreases the blue mold incidence of apple fruits caused by *Penicillium expansum* by an increase in the enzymatic activity of peroxidase, catalase, β-1,3-glucanase, and the concentration of phenolic compounds, related as defense mechanisms against pathogens [42]. Studies realized in different plants such as tobacco, broccoli, tomato, lemon, apple, potato, and rice, reported that different *Trichoderma* species promote the expression of

*DOI: http://dx.doi.org/10.5772/intechopen.82784*

Banana Crown rot *Lasiodiplodia* 

Pear Rotten spots *Rhizopus* 

Mango Antracnosis *Colletotrichum* 

Apple Blue mold *Penicillium* 

Apple Gray mold *Botrytis* 

Pear Gray mold *Botrytis* 

Tomato Gray mold *Botrytis* 

Papaya Anthracnose *Colletotrichum* 

**Postharvest fruit**


### *A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

### **Table 1.**

*Trichoderma - The Most Widely Used Fungicide*

systems for controlling postharvest pathogens on fruits: as physical treatments (hot water), application of substance of vegetable and animal origin, or the use of antagonistic microorganisms [2, 29, 30]. Biological control is a promising alternative with high application potential for controlling postharvest pathogens. Some of the advantages of biological control compared to the use of fungicides are the absence of toxic waste in the fruits, friendly relationship with the environment, as well as in safety and health for the people who handle the products [14]. For several decades, a large number of microorganisms with potential characteristics of antagonistic organisms have been isolated from different sources (leaves, fruits, marine environment) [4, 31]. Bacteria, yeasts, and fungi are the main antagonistic organisms that have been used at postharvest stage. Countries such as the United States, South Africa, Spain, Italy, and Israel, among others, have developed and, in some cases, commercialized antagonistic microorganisms for their commercial use in postharvest, for the control of different diseases caused by the main pathogenic fungi of fruits of temperate, tropical, and subtropical origin [27, 32, 33]. The most widely used antagonists have an outstanding capacity for rapid growth and colonization of the fruit, nutrient acquisition, ability to withstand extreme temperature conditions, solar radiation, control capacity at low concentrations, as well as genetic stability and development capacity in economic cultivation means, ease of application in the management of postharvest. The main mechanisms of action present in the antagonists are antibiosis, production of lytic enzymes, competence by nutri-

ents and space, or the induction of resistance mechanisms [1, 34].

been evaluated with favorable results [36, 37].

**6.** *Trichoderma***: application on fruits**

There is a large number of reports of *in vitro* and *in vivo* tests, using bacteria such as *Pseudomonas cepacia* Van Hall, *Pseudomonas syringae*, *Bacillus subtilis*; as well as yeasts such as *Candida sake*, *Rhodotorula glutinis, Debaryomyces hansenii* for the control of the main pathogenic fruit fungi in postharvest, such as *Penicillium digitatum*, *Penicillium italicum, Alternaria, Aspergillus, Botrytis, Fusarium, Geotrichum, Gloeosporium, Monilinia, Penicillium, Mucor, Colletotrichum*, and *Rhizopus* [4, 27, 32, 33, 35]. In addition to bacteria and yeasts used in the biological control postharvest pathogens, the application of *Trichoderma*, alone or in combination with other alternative control systems in different fruits for the control of pathogenic fungi of fruits in the fruit, has

*Trichoderma* has been applied as postharvest biocontrol agent in different crops such as papayas, strawberries, tomatoes, apples, pears, and bananas (**Table 1**). Existing different species of *Trichoderma* with high antagonistic capacity are *T. asperellum*, *T. viride*, and *T. harzianum.* Several authors have investigated different species of *Trichoderma* with the objective to find the most effective biocontrol agent for each crop and pathogen. The efficacy of six species of *Trichoderma* was evaluated on papaya fruit reducing the diameter lesions and disease incidence caused by *Colletotrichum gloeosporioides* by the application of *T. asperellum* and *T. viride* [6]. In a similar study on mango fruit, good efficacy was reported for controlling anthracnose applying *T. harzianum* obtaining a lower disease incidence (41.7% disease incidence) in fruits compared to control [38]. At present, *Trichoderma* is produced at industrial level as active component of biological products "biopesticides"; other ingredients that conform the biopesticides are the edible polymers, which can form coating for easy adhesion to the fruit and give to the product protection and stability during its shelf life. The application of biopesticides is widely used in agriculture and can be applied by immersion or spraying during the industrialization of

**66**

*Percentage of inhibition of different postharvest diseases by the application of different species of*  Trichoderma*.*

agricultural products [39]. The incorporation of *T. harzianum* into edible coatings as biopesticide produced higher inhibition of the pathogens *Botrytis cinerea* and *Penicillium expansum* compared to the application as simple conidial suspension of the antagonist on the fruits [40]. The same author previously reported the same effect on other fruits such as pears, grapes, apples, strawberries, kiwis, and peaches [36, 41]. Microbial antagonists not only have antifungal properties, but also can act as inductor agents; some studies report that their application can induce biochemical defense responses by the interaction antagonist-fruit. In this sense, the application of *T. virens* decreases the blue mold incidence of apple fruits caused by *Penicillium expansum* by an increase in the enzymatic activity of peroxidase, catalase, β-1,3-glucanase, and the concentration of phenolic compounds, related as defense mechanisms against pathogens [42]. Studies realized in different plants such as tobacco, broccoli, tomato, lemon, apple, potato, and rice, reported that different *Trichoderma* species promote the expression of

genes dependent on the defense mechanism of plants. Some of the expressed genes were chit36, chit42, agn13.1, and gluc78, which correspond to defense enzymes against cellular attack [7].

### **7.** *Trichoderma***: mechanisms for controlling pathogens**

The biocontrol mechanisms attributed to *Trichoderma* spp. are: competition for nutrients, parasitism, antibiosis, secretion of enzymes, and the production of inhibitor compounds [46, 47]. This biocontrol agent attacks and penetrates fungal cells, causing an alteration with the consequent degradation of the cell wall, causing retraction of the plasma membrane and disorganization of the cytoplasm [48]. These mechanisms are favored by the ability of *Trichoderma* to colonize the rhizosphere of plants.

### **7.1 Competition**

Competition is defined as the unequal behavior of two or more organisms before the same requirement (substrate, nutrients), if the use of this substrate by one of the organisms reduces the amount or space available to others. This type of antagonism is favored by the characteristics of the biological control agent as ecological plasticity, growth rate primarily as chlamydospores [49], speed of development, and external factors such as soil type, pH, temperature, and humidity [50]. Nutrient competition can occur for nitrogen nonstructural carbohydrates (sugars and polysaccharides such as starch, cellulose, chitin, laminarin, and pectin, among others) and microelements.

### **7.2 Mycoparasitism**

Mycoparasitism is defined as an antagonistic symbiosis between organisms, generally involving extracellular enzymes such as chitinases, cellulases, and which correspond to the composition and structure of the cell walls of parasitized fungi [51]. *Trichoderma* species' mycoparasitism during chemotropical process grows toward the host; hyphae adhere to it, wound on them frequently, and sometimes penetrate. Degradation of the cell walls of the host is observed in the late stages of the parasitic process [52], which leads to almost total phytopathogen weakening. This process is explained in four stages, within which it is recognized [53]:


**69**

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma*

*Trichoderma* can excrete metabolites like cellulases, glucanases, lipases, proteases, and chitinases in order to facilitate the insertion of hyphae for nutrient uptake of the pathogen, ending with the loss of cytoplasmic contents of the host cell [54]. The remaining cytoplasm is mainly surrounding the invading hyphae, showing

Antibiosis is the inhibition of pathogen development by metabolized products and small toxic molecules, volatile and lytic enzymes, which operate structural polymers, such as chitin and β-1-3-glucans of the cell wall in most pathogenic fungi, producing an adverse effect on development and differentiation [55]. Given the above, it is said that the greater the amount of metabolic products, the antagonistic power increases; additionally, some authors mention that this mechanism is not the principle, due to the risk of emergence of the antibiotic-resistant pathogens [55].

The production of enzymes such as chitinase and/or glucanases produced by the fungus of *Trichoderma* is involved in the control of pathogenic fungi. These hydrolytic enzymes can degrade the cell wall polysaccharides (chitin and β glucans)

The ability to induce resistance in a wide range of diseases caused by various classes of pathogens (including fungi, bacteria, and viruses) in a wide variety of plants may be an important characteristic of *Trichoderma* [56]. Currently, three classes of compounds are known that are produced by *Trichoderma* strains and that induce resistance in the plant. These are proteins with enzymatic functions, homo-

Fungi possess characteristics that define their potential as biocontrol agents. *Trichoderma* species are cosmopolitan microorganims, inhabitant natural of soil such as organic matter, decaying wood as well as in crops waste. *Trichoderma* has several advantages as a biological control agent, such as a rapid growth and development as well as good production of a large number of enzymes inducible with the presence of phytopathogenic fungi [57]. In soil, Trichoderma has the ability to assimilate nutrients faster than pathogens, favoring its establishment and development, thus controlling pathogen infection and dissemination. Its use as a biocontrol agent can provide excellent advantages from the economic, environmental, and biological point of view, since they do not cause deterioration to the environment, do not affect the development of the plants, their production is cheaper, and its use does not entail the emergence of new pests or secondary pests [58]. However, its production on an industrial scale has some drawbacks that have limited the development of these organisms with wide possibilities as antagonist [59]. Even when, *Trichoderma* already exists in commercial form, its storage life is short, several investigations have been carried out finding that the best methods of fungus conservation [60]. In this sense, the incorporation in the formulations of different additives can improve microbial viability [2]. Other important limitations in the use

logs of proteins encoded for avirulence (Avr), and oligosaccharides.

**8. Advantages and limitations on the use of** *Trichoderma*

*DOI: http://dx.doi.org/10.5772/intechopen.82784*

signs of disintegration.

**7.4 Secretion of enzymes**

affecting its stability and integrity [5].

**7.5 Production of inhibitor compounds**

**7.3 Antibiosis**

### *A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

*Trichoderma* can excrete metabolites like cellulases, glucanases, lipases, proteases, and chitinases in order to facilitate the insertion of hyphae for nutrient uptake of the pathogen, ending with the loss of cytoplasmic contents of the host cell [54]. The remaining cytoplasm is mainly surrounding the invading hyphae, showing signs of disintegration.

### **7.3 Antibiosis**

*Trichoderma - The Most Widely Used Fungicide*

enzymes against cellular attack [7].

**7.1 Competition**

others) and microelements.

chemical stimulus.

their extracellular environment.

present in the wall of the pathogen.

and allow the penetration of antagonist hyphae.

**7.2 Mycoparasitism**

genes dependent on the defense mechanism of plants. Some of the expressed genes were chit36, chit42, agn13.1, and gluc78, which correspond to defense

The biocontrol mechanisms attributed to *Trichoderma* spp. are: competition for nutrients, parasitism, antibiosis, secretion of enzymes, and the production of inhibitor compounds [46, 47]. This biocontrol agent attacks and penetrates fungal cells, causing an alteration with the consequent degradation of the cell wall, causing retraction of the plasma membrane and disorganization of the cytoplasm [48]. These mechanisms are

Competition is defined as the unequal behavior of two or more organisms before the same requirement (substrate, nutrients), if the use of this substrate by one of the organisms reduces the amount or space available to others. This type of antagonism is favored by the characteristics of the biological control agent as ecological plasticity, growth rate primarily as chlamydospores [49], speed of development, and external factors such as soil type, pH, temperature, and humidity [50]. Nutrient competition can occur for nitrogen nonstructural carbohydrates (sugars and polysaccharides such as starch, cellulose, chitin, laminarin, and pectin, among

Mycoparasitism is defined as an antagonistic symbiosis between organisms, generally involving extracellular enzymes such as chitinases, cellulases, and which correspond to the composition and structure of the cell walls of parasitized fungi [51]. *Trichoderma* species' mycoparasitism during chemotropical process grows toward the host; hyphae adhere to it, wound on them frequently, and sometimes penetrate. Degradation of the cell walls of the host is observed in the late stages of the parasitic process [52], which leads to almost total phytopathogen weakening. This process is explained in four stages, within which it is recognized [53]:

1.*Chemotrophic growth*: the positive chemotropism directs growth toward a

2.*Recognition:* it is based on lectin-carbohydrate interactions. The lectins are proteins linked to sugars or glycoproteins, which agglutinate cells and are involved in the interactions between the components of the cell surface and

3.*Adhesion and curl:* occurs when the acknowledgment response is positive, *Trichoderma* hyphae adhere to host-mediated enzymatic processes. Hyphae adhesion occurs through association of a sugar antagonist wall with a lectin

4.*Lytic activity:* this stage is the production of extracellular lytic enzymes, mainly chitinases, glucanases, and proteases, which degrade the cell walls of the host

**7.** *Trichoderma***: mechanisms for controlling pathogens**

favored by the ability of *Trichoderma* to colonize the rhizosphere of plants.

**68**

Antibiosis is the inhibition of pathogen development by metabolized products and small toxic molecules, volatile and lytic enzymes, which operate structural polymers, such as chitin and β-1-3-glucans of the cell wall in most pathogenic fungi, producing an adverse effect on development and differentiation [55]. Given the above, it is said that the greater the amount of metabolic products, the antagonistic power increases; additionally, some authors mention that this mechanism is not the principle, due to the risk of emergence of the antibiotic-resistant pathogens [55].

### **7.4 Secretion of enzymes**

The production of enzymes such as chitinase and/or glucanases produced by the fungus of *Trichoderma* is involved in the control of pathogenic fungi. These hydrolytic enzymes can degrade the cell wall polysaccharides (chitin and β glucans) affecting its stability and integrity [5].

### **7.5 Production of inhibitor compounds**

The ability to induce resistance in a wide range of diseases caused by various classes of pathogens (including fungi, bacteria, and viruses) in a wide variety of plants may be an important characteristic of *Trichoderma* [56]. Currently, three classes of compounds are known that are produced by *Trichoderma* strains and that induce resistance in the plant. These are proteins with enzymatic functions, homologs of proteins encoded for avirulence (Avr), and oligosaccharides.

### **8. Advantages and limitations on the use of** *Trichoderma*

Fungi possess characteristics that define their potential as biocontrol agents. *Trichoderma* species are cosmopolitan microorganims, inhabitant natural of soil such as organic matter, decaying wood as well as in crops waste. *Trichoderma* has several advantages as a biological control agent, such as a rapid growth and development as well as good production of a large number of enzymes inducible with the presence of phytopathogenic fungi [57]. In soil, Trichoderma has the ability to assimilate nutrients faster than pathogens, favoring its establishment and development, thus controlling pathogen infection and dissemination. Its use as a biocontrol agent can provide excellent advantages from the economic, environmental, and biological point of view, since they do not cause deterioration to the environment, do not affect the development of the plants, their production is cheaper, and its use does not entail the emergence of new pests or secondary pests [58]. However, its production on an industrial scale has some drawbacks that have limited the development of these organisms with wide possibilities as antagonist [59]. Even when, *Trichoderma* already exists in commercial form, its storage life is short, several investigations have been carried out finding that the best methods of fungus conservation [60]. In this sense, the incorporation in the formulations of different additives can improve microbial viability [2]. Other important limitations in the use of *Trichoderma* involve the lack of precise information to farmers, the little training about how to use the antagonistic fungi, as well as the lack of government economic support [61].

### **9.** *Trichoderma* **in combination with other control systems at postharvest management**

In the management of postharvest fruit diseases, the use of antagonistic fungi is an alternative with great potential; specifically, the application of *Trichoderma* and its various species has obtained very promising results leading to the fabrication of commercial biocontrol products [62]. In addition, considering their mechanisms of action, such as competition for nutrients, production of lytic enzymes, parasitism, antibiosis, and induction of resistance, ensures the control and destruction of the main postharvest fungi. Even when *Trichoderma* has several mechanisms of action reported [63–65], the sole application of antagonists for controlling postharvest pathogens does not ensure 100% of disease control, that is why the development of new control technologies like combined system with other substances and methods could offer economical treatments and enhance biocontrol activity [66–68]. The combination of strains of *Pseudomonas*-*Trichoderma* and *Trichoderma viride*-*Bacillus subtilis* was effective controlling *in vitro* and *in vivo* tests such as *Penicillium digitatum* and *Penicillium expansum* and *Fusarium moniliforme,* respectively, in citrus and grapes. These studies confirm the synergistic effect of the combination of biological control agents [69, 70]. The addition of *Trichoderma harzianum* in polymeric matrices like chitosan has been evaluated against *Fusarium oxysporum* with good results, inducing defense mechanisms and the production of lytic enzymes as well as parasitism. In strawberry fruits, the incorporation of *Trichoderma* in chitosan and the application of a physical treatment (hot air) were effective to reduce microorganisms and maintain the fruit quality [71, 72]. The efficacy of the combination of bacterial (*P. syringae*) and fungal (*Trichoderma*) antagonists has been evaluated against *Botrytis cinerea* and *Fusarium oxysporum in vitro* tests, a synergistic effect was observed controlling successfully both fungus by the alteration and, consequently, the degradation of cell wall [73]. It has been reported that the different species of *Trichoderma* present different degrees of pathogen control at postharvest stage applied individually, but if the strains are combining, the potential of biocontrol increases, as previously reported in combinations with *T. viride*, *T. harzianum,* and *T. koningii* in the control of *Colletotrichum musae* reducing the incidence of crown rot [68]. The use of GRAS substances like sodium bicarbonate with *Trichoderma harzianum* was effective to control crown rot in banana fruits (*Colletotrichum musae* and *Fusarium verticillioides*) [74]. Recently, silver nanoparticles with *Trichoderma* have been synthesized and their antifungal capacity against postharvest pathogenic fungi such as *Alternaria*, *Penicillium*, and *Fusarium* has been evaluated; a good control on the mycelial growth and development of the pathogenic fungi was reported at low concentrations of *Trichoderma* spp. [75].

### **10. Conclusions**

*Trichoderma* is a biocontrol agent with several benefits for its application on different commodities; this fungus represents a suitable eco-friendly alternative to fungicides by reducing postharvest losses.

**71**

provided the original work is properly cited.

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

\*, Francisco Blancas-Benítez1

1 National Technology of Mexico/Technological Institute of Tepic, Tepic, Nayarit,

, Anelsy Ramos-Guerrero1

3 Technological University of Nayarit, Tepic, Nayarit, Mexico

\*Address all correspondence to: ramgonzalez@ittepic.edu.mx

2 University of Sonora, Hermosillo, Sonora, Mexico

, Luz Del Carmen Romero-Islas1

, Beatriz Montaño-Leyva<sup>2</sup>

, Angel Fonseca-Cantabrana1

, Jovita Romero-Islas1

,

,

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma*

*DOI: http://dx.doi.org/10.5772/intechopen.82784*

Authors declare no conflict of interest.

**Conflict of interest**

**Author details**

Rosa Avila-Peña3

Mexico

Ramsés González-Estrada1

Cristina Moreno-Hernández1

and Porfirio Gutierrez-Martinez1

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

### **Conflict of interest**

*Trichoderma - The Most Widely Used Fungicide*

support [61].

**management**

of *Trichoderma* involve the lack of precise information to farmers, the little training about how to use the antagonistic fungi, as well as the lack of government economic

**9.** *Trichoderma* **in combination with other control systems at postharvest** 

In the management of postharvest fruit diseases, the use of antagonistic fungi is an alternative with great potential; specifically, the application of *Trichoderma* and its various species has obtained very promising results leading to the fabrication of commercial biocontrol products [62]. In addition, considering their mechanisms of action, such as competition for nutrients, production of lytic enzymes, parasitism, antibiosis, and induction of resistance, ensures the control and destruction of the main postharvest fungi. Even when *Trichoderma* has several mechanisms of action reported [63–65], the sole application of antagonists for controlling postharvest pathogens does not ensure 100% of disease control, that is why the development of new control technologies like combined system with other substances and methods could offer economical treatments and enhance biocontrol activity [66–68]. The combination of strains of *Pseudomonas*-*Trichoderma* and *Trichoderma viride*-*Bacillus subtilis* was effective controlling *in vitro* and *in vivo* tests such as *Penicillium digitatum* and *Penicillium expansum* and *Fusarium moniliforme,* respectively, in citrus and grapes. These studies confirm the synergistic effect of the combination of biological control agents [69, 70]. The addition of *Trichoderma harzianum* in polymeric matrices like chitosan has been evaluated against *Fusarium oxysporum* with good results, inducing defense mechanisms and the production of lytic enzymes as well as parasitism. In strawberry fruits, the incorporation of *Trichoderma* in chitosan and the application of a physical treatment (hot air) were effective to reduce microorganisms and maintain the fruit quality [71, 72]. The efficacy of the combination of bacterial (*P. syringae*) and fungal (*Trichoderma*) antagonists has been evaluated against *Botrytis cinerea* and *Fusarium oxysporum in vitro* tests, a synergistic effect was observed controlling successfully both fungus by the alteration and, consequently, the degradation of cell wall [73]. It has been reported that the different species of *Trichoderma* present different degrees of pathogen control at postharvest stage applied individually, but if the strains are combining, the potential of

biocontrol increases, as previously reported in combinations with

*T. viride*, *T. harzianum,* and *T. koningii* in the control of *Colletotrichum musae* reducing the incidence of crown rot [68]. The use of GRAS substances like sodium bicarbonate with *Trichoderma harzianum* was effective to control crown rot in banana fruits (*Colletotrichum musae* and *Fusarium verticillioides*) [74]. Recently, silver nanoparticles with *Trichoderma* have been synthesized and their antifungal capacity against postharvest pathogenic fungi such as *Alternaria*, *Penicillium*, and *Fusarium* has been evaluated; a good control on the mycelial growth and development of the pathogenic fungi was reported at low concentra-

*Trichoderma* is a biocontrol agent with several benefits for its application on different commodities; this fungus represents a suitable eco-friendly alternative to

**70**

tions of *Trichoderma* spp. [75].

fungicides by reducing postharvest losses.

**10. Conclusions**

Authors declare no conflict of interest.

### **Author details**

Ramsés González-Estrada1 \*, Francisco Blancas-Benítez1 , Beatriz Montaño-Leyva<sup>2</sup> , Cristina Moreno-Hernández1 , Luz Del Carmen Romero-Islas1 , Jovita Romero-Islas1 , Rosa Avila-Peña3 , Anelsy Ramos-Guerrero1 , Angel Fonseca-Cantabrana1 and Porfirio Gutierrez-Martinez1

1 National Technology of Mexico/Technological Institute of Tepic, Tepic, Nayarit, Mexico

2 University of Sonora, Hermosillo, Sonora, Mexico

3 Technological University of Nayarit, Tepic, Nayarit, Mexico

\*Address all correspondence to: ramgonzalez@ittepic.edu.mx

© 2018 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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[6] Valenzuela N, Angel D, Ortiz D, Rosas R, García C, Santos M. Biological control of anthracnose by postharvest application of *Trichoderma* spp. on maradol papaya fruit. Biological Control. 2015;**91**:88-93. DOI: 0.1016/j. biocontrol.2015.08.002

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[10] Barbosa-Cánovas G. Handling and Preservation of Fruits and Vegetables by Combined Methods for Rural Areas: Technical Manual, No. 149. Food and Agriculture Organization; 2003. Available from: https://books.google.com.mx/ books?hl=es&lr=&id=IKsaRY9sb\_AC& oi=fnd&pg=PA1&dq=Handling+an d+P reservation+of+Fruits+and+Veg etabl es+by+Combined+Methods+fo r+Rural+Areas&ots=8RHV2OIVqt& sig=0dHL3i44Vkk2mK5gycYDAQN wA3E#v=onepage&q=Handling%20 and%20Preservation%20of%20 Fruits%20and%20Vegetables%20 by%20Combined%20Methods%20 for%20Rural%20Areas&f=false. [Review date:November 2018]

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[20] Han J. Edible films and coatings: A review. In: Han J, editor. Innovations in Food Packaging. 2nd ed. Academic press, Elsevier; 2014. pp. 213-255. DOI: 0.1016/B978-0-12-394601-0.00009-6

[21] Falguera V, Quintero J, Jiménez A, Muñoz J, Ibarz A. Edible films and coatings: Structures, active functions and trends in their use. Trends in Food Science and Technology. 2011;**22**: 292-303. DOI: 0.1016/j.tifs.2011.02.004

[22] Otoni C, Avena-Bustillos R, Azeredo H, Lorevice M, Moura M, Mattoso L, et al. Recent advances on edible films based on fruits and vegetables—A review. Comprehensive Reviews in Food Science and Food Safety. 2017;**16**: 1151-1169. DOI: 0.1111/1541-4337.12281

[23] González-Estrada R, Carvajal-Millán E, Ragazzo-Sánchez J, Bautista-Rosales P, Calderón-Santoyo M. Control of blue mold decay on Persian lime: Application of covalently cross-linked arabinoxylans bioactive coatings with antagonistic yeast entrapped. LWT- Food Science and Technology. 2017;**85**:187-196. DOI: 0.1016/j.

[24] Ponce A, Roura S, Del Valle C, Moreira M. Antimicrobial and antioxidant activities of edible coatings enriched with natural plant extracts: in vitro and in vivo studies. Postharvest Biology and Technology. 2008;**49**:294-300. DOI: 0.1016/j. postharvbio.2008.02.013

[25] Chávez-Magdaleno M, González-Estrada R, Plascencia-Jatomea M, Gutiérrez-Martínez P. Effect of pepper tree (Schinus molle) essential

lwt.2017.07.019

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[13] Janisiewicz W, Korsten L. Biological control of postharvest diseases of fruits. Annual Review of Phytopathology. 2002;**40**:411-441. DOI: 0.1146/annurev.

[14] Droby S, Wisniewski M, El-Ghaouth A, Wilson C. Biological control of postharvest diseases of fruit and vegetables: Current achievements and future challenges. In: XXVI International Horticultural Congress: Issues and Advances in Postharvest Horticulture. 2002. pp. 703-713. DOI: 10.17660/ActaHortic.2003.628.89

[15] Sui Y, Wisniewski M, Droby S, Norelli J, Liu J. Recent advances and current status of the use of heat treatments in postharvest disease management systems: Is it time to turn up the heat? Trends in Food Science and Technology. 2016;**51**:34-40. DOI:

[16] Wisniewski M, Droby S, Norelli J, Liu J, Schena L. Alternative management technologies for postharvest disease control: The journey from simplicity to complexity. Postharvest Biology and Technology. 2016;**122**:3-10. DOI: 0.1016/j.postharvbio.2016.05.012

[17] Sivakumar D, Bautista-Baños S. A review on the use of essential oils for postharvest decay control and maintenance of fruit quality during storage. Crop Protection. 2014;**64**:27-37.

DOI: 0.1016/j.cropro.2014.05.012

[18] González-Estrada R, Calderón-Santoyo M, Ragazzo-Sánchez J, Peyron S, Chalier P. Antimicrobial soy protein isolate-based films: Physical characterisation, active agent retention and antifungal properties against *Penicillium italicum*. International Journal of Food Science and Technology. 2018;**53**:921-929. DOI:

[19] González-Estrada R, Chalier P, Ragazzo-Sánchez J, Konuk D,

0.1016/j.tifs.2016.03.004

phyto.40.120401.130158

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

[13] Janisiewicz W, Korsten L. Biological control of postharvest diseases of fruits. Annual Review of Phytopathology. 2002;**40**:411-441. DOI: 0.1146/annurev. phyto.40.120401.130158

[14] Droby S, Wisniewski M, El-Ghaouth A, Wilson C. Biological control of postharvest diseases of fruit and vegetables: Current achievements and future challenges. In: XXVI International Horticultural Congress: Issues and Advances in Postharvest Horticulture. 2002. pp. 703-713. DOI: 10.17660/ActaHortic.2003.628.89

[15] Sui Y, Wisniewski M, Droby S, Norelli J, Liu J. Recent advances and current status of the use of heat treatments in postharvest disease management systems: Is it time to turn up the heat? Trends in Food Science and Technology. 2016;**51**:34-40. DOI: 0.1016/j.tifs.2016.03.004

[16] Wisniewski M, Droby S, Norelli J, Liu J, Schena L. Alternative management technologies for postharvest disease control: The journey from simplicity to complexity. Postharvest Biology and Technology. 2016;**122**:3-10. DOI: 0.1016/j.postharvbio.2016.05.012

[17] Sivakumar D, Bautista-Baños S. A review on the use of essential oils for postharvest decay control and maintenance of fruit quality during storage. Crop Protection. 2014;**64**:27-37. DOI: 0.1016/j.cropro.2014.05.012

[18] González-Estrada R, Calderón-Santoyo M, Ragazzo-Sánchez J, Peyron S, Chalier P. Antimicrobial soy protein isolate-based films: Physical characterisation, active agent retention and antifungal properties against *Penicillium italicum*. International Journal of Food Science and Technology. 2018;**53**:921-929. DOI: 0.1111/ijfs.13664

[19] González-Estrada R, Chalier P, Ragazzo-Sánchez J, Konuk D,

Calderón-Santoyo M. Antimicrobial soy protein based coatings: Application to Persian lime (*Citrus latifolia* Tanaka) for protection and preservation. Postharvest Biology and Technology. 2017;**132**:138-144. DOI: 0.1016/j. postharvbio.2017.06.005

[20] Han J. Edible films and coatings: A review. In: Han J, editor. Innovations in Food Packaging. 2nd ed. Academic press, Elsevier; 2014. pp. 213-255. DOI: 0.1016/B978-0-12-394601-0.00009-6

[21] Falguera V, Quintero J, Jiménez A, Muñoz J, Ibarz A. Edible films and coatings: Structures, active functions and trends in their use. Trends in Food Science and Technology. 2011;**22**: 292-303. DOI: 0.1016/j.tifs.2011.02.004

[22] Otoni C, Avena-Bustillos R, Azeredo H, Lorevice M, Moura M, Mattoso L, et al. Recent advances on edible films based on fruits and vegetables—A review. Comprehensive Reviews in Food Science and Food Safety. 2017;**16**: 1151-1169. DOI: 0.1111/1541-4337.12281

[23] González-Estrada R, Carvajal-Millán E, Ragazzo-Sánchez J, Bautista-Rosales P, Calderón-Santoyo M. Control of blue mold decay on Persian lime: Application of covalently cross-linked arabinoxylans bioactive coatings with antagonistic yeast entrapped. LWT- Food Science and Technology. 2017;**85**:187-196. DOI: 0.1016/j. lwt.2017.07.019

[24] Ponce A, Roura S, Del Valle C, Moreira M. Antimicrobial and antioxidant activities of edible coatings enriched with natural plant extracts: in vitro and in vivo studies. Postharvest Biology and Technology. 2008;**49**:294-300. DOI: 0.1016/j. postharvbio.2008.02.013

[25] Chávez-Magdaleno M, González-Estrada R, Plascencia-Jatomea M, Gutiérrez-Martínez P. Effect of pepper tree (Schinus molle) essential

**72**

*Trichoderma - The Most Widely Used Fungicide*

International Journal of Current Microbiology and Applied Sciences. 2018;**7**:2382-2397. DOI: 10.20546/

[8] W. H. Organization. Fruit and Vegetable Promotion Initiative: A Meeting Report, 25-27/08/03. Geneva: World Health Organization; 2003

[9] W. H. Organization. Diet, Nutrition, and the Prevention of Chronic Diseases: Report of a Joint WHO/FAO Expert Consultation. Vol. 916. World Health Organization; 2003. Available from: http://apps.who.int/ iris/bitstream/handle/10665/42665/? sequence=1. [Review date: November

[10] Barbosa-Cánovas G. Handling and Preservation of Fruits and Vegetables by Combined Methods for Rural Areas: Technical Manual, No. 149. Food and Agriculture Organization; 2003. Available from: https://books.google.com.mx/

books?hl=es&lr=&id=IKsaRY9sb\_AC& oi=fnd&pg=PA1&dq=Handling+an d+P reservation+of+Fruits+and+Veg etabl es+by+Combined+Methods+fo r+Rural+Areas&ots=8RHV2OIVqt& sig=0dHL3i44Vkk2mK5gycYDAQN wA3E#v=onepage&q=Handling%20 and%20Preservation%20of%20 Fruits%20and%20Vegetables%20 by%20Combined%20Methods%20 for%20Rural%20Areas&f=false. [Review date:November 2018]

[11] FAOSTAT. "Agriculture Organization of the United Nations Statistics Division (2014)," Prod. 2016. Available from: http//faostat3.fao.org/browse/Q/QC/S

[12] FAO. Losses and food waste in Latin America and the Caribbean, 2018. Available from: http://www.fao.org/ americas/noticias/ver/en/c/239392/. [Review date: November 2018]

[Review date: April 2015]

ijcmas.2018.702.291

2018]

[1] Nunes C. Biological control of postharvest diseases of fruit. European Journal of Plant Pathology. 2012;**133**:181-196. DOI: 10.1007/

[2] Sharma R, Singh D, Singh R. Biological control of postharvest diseases of fruits and vegetables by microbial antagonists: A review. Biological Control. 2009;**50**:205-221. DOI: 10.1016/j.biocontrol.2009.05.001

[3] Gutiérrez-Martínez P, Ramos-Guerrero A, Rodríguez-Pereida C, Coronado-Partida L, Angulo-Parra J, González-Estrada R. Chitosan for postharvest disinfection of fruits and vegetables. In: Wasim Siddiqui M, editor. Postharvest Disinfection of Fruits and Vegetables. 1st ed. Academic Press: Elsevier; 2018. pp. 231-241. DOI: 10.1016/ B978-0-12-812698-1.00012-1

[4] Medina-Cordova N, Rosales-Mendoza S, Hernández-Montiel L, Angulo C. The potential use of

*Debaryomyces hansenii* for the biological control of pathogenic fungi in food. Biological Control. 2018;**121**:216-222. DOI: 0.1016/j.biocontrol.2018.03.002

[5] Howell C. Mechanisms employed by *Trichoderma* species in the biological control of plant diseases: The history and evolution of current concepts. Plant Disease. 2003;**87**:4-10. DOI: 0.1094/

[6] Valenzuela N, Angel D, Ortiz D, Rosas R, García C, Santos M. Biological control of anthracnose by postharvest application of *Trichoderma* spp. on maradol papaya fruit. Biological Control. 2015;**91**:88-93. DOI: 0.1016/j.

[7] Singh A, Shukla N, Kabadwal B, Tewari A, Kumar J. Review on plant-*Trichoderma*-pathogen interaction.

PDIS.2003.87.1.4

biocontrol.2015.08.002

**References**

s10658-011-9919-7

oil-loaded chitosan bio-nanocomposites on postharvest control of *Colletotrichum gloeosporioides* and quality evaluations in avocado (*Persea americana*) cv. Hass. Food Science and Biotechnology. 2018;**27**: 1871-1875. DOI: 0.1007/ s10068-018-0410-5

[26] Liu J, Sui Y, Wisniewski M, Droby S, Liu Y. Utilization of antagonistic yeasts to manage postharvest fungal diseases of fruit. International Journal of Food Microbiology. 2013;**167**:153-160. DOI: 0.1016/j. ijfoodmicro.2013.09.004

[27] Droby S, Wisniewski M, Macarisin D, Wilson C. Twenty years of postharvest biocontrol research: Is it time for a new paradigm? Postharvest Biology and Technology. 2009;**52**:137-145. DOI: 0.1016/j. postharvbio.2008.11.009

[28] Mari M, Bautista-Baños S, Sivakumar D. Decay control in the postharvest system: Role of microbial and plant volatile organic compounds. Postharvest Biology and Technology. 2016;**122**:70-81. DOI: 10.1016/j. postharvbio.2016.04.014

[29] Gutierrez-Martinez P, Ledezma-Morales A, Romero-Islas L, Ramos-Guerrero A, Romero-Islas J, Rodríguez-Pereida C, et al. Antifungal activity of chitosan against postharvest fungi of tropical and subtropical fruits. In: Dongre R, editor. Chitin-Chitosan-Myriad Functionalities in Science and Technology. 1st ed. Rijeka: IntechOpen; 2018. pp. 311-327. DOI: 10.5772/ intechopen.76095

[30] Gutiérrez-Martínez P, Osuna-López SG, Calderón-Santoyo M, Cruz-Hernández A, Bautista-Baños S. Influence of ethanol and heat on disease control and quality in stored mango fruits. LWT- Food Science and Technology. 2012;**45**:20-27. DOI: 0.1016/j.lwt.2011.07.033

[31] Kong Q. Marine microorganisms as biocontrol agents against fungal phytopathogens and mycotoxins. Biocontrol Science and Technology. 2018;**28**:77-93. DOI: 0.1080/09583157.2017.1419164

[32] Liu J, Wisniewski M, Artlip T, Sui Y, Droby S, Norelli J. The potential role of PR-8 gene of apple fruit in the mode of action of the yeast antagonist, *Candida oleophila*, in postharvest biocontrol of *Botrytis cinerea*. Postharvest Biology and Technology. 2013;**85**:203-209. DOI: 0.1016/j.postharvbio.2013.06.007

[33] Calvo-Garrido C, Viñas I, Usall J, Rodríguez-Romera M, Ramos M, Teixidó N. Survival of the biological control agent *Candida* sake CPA-1 on grapes under the influence of abiotic factors. Journal of Applied Microbiology. 2014;**117**:800-811. DOI: 0.1111/jam.12570

[34] Spadaro D, Droby S. Development of biocontrol products for postharvest diseases of fruit: The importance of elucidating the mechanisms of action of yeast antagonists. Trends in Food Science and Technology. 2016;**47**:39-49. DOI: 0.1016/j.tifs.2015.11.003

[35] González-Estrada R, Ascencio-Valle F, Ragazzo-Sánchez J, Calderón-Santoyo M. Use of a marine yeast as a biocontrol agent of the novel pathogen *Penicillium citrinum* on Persian lime. Emirates Journal of Food and Agriculture. 2017;**29**:114-122. DOI: 0.9755/ ejfa.2016-09-1273

[36] Batta Y. Control of postharvest diseases of fruit with an invert emulsion formulation of *Trichoderma harzianum* Rifai. Postharvest Biology and Technology. 2007;**43**:143-150. DOI: 0.1016/j.postharvbio.2006.07.010

[37] Wilson-Wijeratnam R, Marikar F, Abeyesekere M, Wijesundera R, Sivakumar D. Antagonistic effect of *Trichoderma harzianum* on post

**75**

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma*

[44] Quaglia M, Ederli L, Pasqualini S, Zazzerini A. Biological control agents and chemical inducers of resistance for postharvest control of *Penicillium expansum* Link. on apple fruit. Postharvest Biology and Technology. 2011;**59**:307-315. DOI: 0.1016/j. postharvbio.2010.09.007

[45] Dal Bello G, Lampugnani G, Abramoff C, Fusé C, Perelló A. Postharvest control of *Botrytis* gray mould in tomato by antagonists and biorational compounds. IOBC-WPRS

[46] Guédez C, Cañizález L, Castillo C, Olivar R. Efecto antagónico de *Trichoderma harzianum* sobre algunos hongos patógenos postcosecha de la fresa (Fragaria spp). Revista de la Sociedad Venezolana de Microbiología. 2009;**29**:34-38

[47] Zimand G, Elad Y, Chet I. Effect of *Trichoderma harzianum* on *Botrytis cinerea* pathogenicity. Phytopathology.

[48] Tronsmo A, Raa J. Antagonistic action of *Trichoderma pseudokoningii* against the apple pathogen *Botrytis cinerea*. Journal of Phytopathology.

[49] Hjeljord L, Tronsmo A. *Trichoderma* and *Gliocladium* in bilogical control: A review. In: Harman GE, Kubicek CP, editors. *Trichoderma* & *Gliocladium*— Enzymes, Biological Control and Commercial Applications. 1st. ed. CRC Press, Taylor & Francis; 1998.

[50] Ahmad J, Baker R. Rhizosphere competence of *Trichoderma harzianum*. Phytopathology. 1987;**77**:182-189

[51] Lorito M, Harman G, Di Prieto A, Hayes C. Extracellular chitimolytie enzymes produced by *T. harzianum*, purification, characterization and molecular cloning. Phytopathology.

Bulletin. 2015;**111**:417-425

1996;**86**:1255-1260

1977;**89**:216-220

pp. 131- 145

1990;**82**:10-77

*DOI: http://dx.doi.org/10.5772/intechopen.82784*

harvest pathogens of rambutans. In: IV International Conference on Postharvest Science. 2000. pp. 389-392. DOI: 10.17660/ ActaHortic.2001.553.88

[38] Prabakar K, Raguchander T, Saravanakumar D, Muthulakshmi P,

Management of postharvest disease of mango anthracnose incited by *Colletotrichum gloeosporioides*. Archives of Phytopathology and Plant Protection. 2008;**41**:333-339. DOI: 0.1080/03235400600793502

[39] Marín A, Atarés L, Chiralt A.

biocontrol agents incorporated in antifungal fruit coatings: A review. Biocontrol Science and Technology. 2017;**27**:1220-1241. DOI: e0.1080/09583157.2017.1390068

[40] Batta Y. Production and testing of biopesticide for control of

postharvest mold infections on fresh fruits of apple and pear. Advances in Microbiology. 2015;**5**:787. DOI: 0.4236/

[41] Batta YA. Effect of treatment with *Trichoderma harzianum* Rifai formulated in invert emulsion on postharvest decay of apple blue mold. International Journal of Food Microbiology. 2004;**96**:281-288. DOI: 0.1016/j.ijfoodmicro.2004.04.002

[42] Bordbar F, Etebarian H, Sahebani N, Rohani H. Control of postharvest decay of apple fruit with *Trichoderma virens* isolates and induction of defense responses. Journal of Plant Protection Research. 2010;**50**:146-152. DOI: 0.2478/

[43] Mortuza M, Ilag L. Potential for biocontrol of *Lasiodiplodia theobromae* (Pat.) Griff. & Maubl. in banana fruits by *Trichoderma* species. Biological Control. 1999;**15**:235-240. DOI: 0.1006/

Parthiban V, Prakasam V.

Improving function of

aim.2015.512083

v10045-010-0025-1

bcon.1999.0716

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

harvest pathogens of rambutans. In: IV International Conference on Postharvest Science. 2000. pp. 389-392. DOI: 10.17660/ ActaHortic.2001.553.88

*Trichoderma - The Most Widely Used Fungicide*

oil-loaded chitosan bio-nanocomposites

[31] Kong Q. Marine microorganisms

mycotoxins. Biocontrol Science and Technology. 2018;**28**:77-93. DOI: 0.1080/09583157.2017.1419164

[32] Liu J, Wisniewski M, Artlip T, Sui Y, Droby S, Norelli J. The potential role of PR-8 gene of apple fruit in the mode of action of the yeast antagonist, *Candida oleophila*, in postharvest biocontrol of *Botrytis cinerea*. Postharvest Biology and Technology. 2013;**85**:203-209. DOI: 0.1016/j.postharvbio.2013.06.007

[33] Calvo-Garrido C, Viñas I, Usall J, Rodríguez-Romera M, Ramos M, Teixidó N. Survival of the biological control agent *Candida* sake CPA-1 on grapes under the influence of abiotic factors. Journal of Applied Microbiology. 2014;**117**:800-811. DOI:

[34] Spadaro D, Droby S. Development of biocontrol products for postharvest diseases of fruit: The importance of elucidating the mechanisms of action of yeast antagonists. Trends in Food Science and Technology. 2016;**47**:39-49.

[35] González-Estrada R, Ascencio-Valle F, Ragazzo-Sánchez J, Calderón-Santoyo M. Use of a marine yeast as a biocontrol agent of the novel pathogen *Penicillium citrinum* on Persian lime. Emirates Journal of Food and Agriculture. 2017;**29**:114-122. DOI: 0.9755/

[36] Batta Y. Control of postharvest diseases of fruit with an invert emulsion formulation of *Trichoderma harzianum* Rifai. Postharvest Biology and Technology. 2007;**43**:143-150. DOI: 0.1016/j.postharvbio.2006.07.010

[37] Wilson-Wijeratnam R, Marikar F, Abeyesekere M, Wijesundera R, Sivakumar D. Antagonistic effect of *Trichoderma harzianum* on post

DOI: 0.1016/j.tifs.2015.11.003

0.1111/jam.12570

ejfa.2016-09-1273

as biocontrol agents against fungal phytopathogens and

[26] Liu J, Sui Y, Wisniewski M, Droby S, Liu Y. Utilization of antagonistic yeasts to manage postharvest fungal diseases of fruit. International Journal of Food Microbiology. 2013;**167**:153-160. DOI: 0.1016/j.

on postharvest control of

Biotechnology. 2018;**27**: 1871-1875. DOI: 0.1007/ s10068-018-0410-5

ijfoodmicro.2013.09.004

[27] Droby S, Wisniewski M,

[28] Mari M, Bautista-Baños S, Sivakumar D. Decay control in the postharvest system: Role of microbial and plant volatile organic compounds. Postharvest Biology and Technology. 2016;**122**:70-81. DOI: 10.1016/j. postharvbio.2016.04.014

[29] Gutierrez-Martinez P, Ledezma-

Ramos-Guerrero A, Romero-Islas J, Rodríguez-Pereida C, et al. Antifungal activity of chitosan against postharvest fungi of tropical and subtropical fruits. In: Dongre R, editor. Chitin-Chitosan-Myriad Functionalities in Science and Technology. 1st ed. Rijeka: IntechOpen;

2018. pp. 311-327. DOI: 10.5772/

[30] Gutiérrez-Martínez P, Osuna-López SG, Calderón-Santoyo M, Cruz-Hernández A, Bautista-Baños S. Influence of ethanol and heat on disease control and quality in stored mango fruits. LWT- Food Science and Technology. 2012;**45**:20-27. DOI:

intechopen.76095

0.1016/j.lwt.2011.07.033

Morales A, Romero-Islas L,

Macarisin D, Wilson C. Twenty years of postharvest biocontrol research: Is it time for a new paradigm? Postharvest Biology and Technology. 2009;**52**:137-145. DOI: 0.1016/j. postharvbio.2008.11.009

*Colletotrichum gloeosporioides* and quality evaluations in avocado (*Persea americana*) cv. Hass. Food Science and

**74**

[38] Prabakar K, Raguchander T, Saravanakumar D, Muthulakshmi P, Parthiban V, Prakasam V. Management of postharvest disease of mango anthracnose incited by *Colletotrichum gloeosporioides*. Archives of Phytopathology and Plant Protection. 2008;**41**:333-339. DOI: 0.1080/03235400600793502

[39] Marín A, Atarés L, Chiralt A. Improving function of biocontrol agents incorporated in antifungal fruit coatings: A review. Biocontrol Science and Technology. 2017;**27**:1220-1241. DOI: e0.1080/09583157.2017.1390068

[40] Batta Y. Production and testing of biopesticide for control of postharvest mold infections on fresh fruits of apple and pear. Advances in Microbiology. 2015;**5**:787. DOI: 0.4236/ aim.2015.512083

[41] Batta YA. Effect of treatment with *Trichoderma harzianum* Rifai formulated in invert emulsion on postharvest decay of apple blue mold. International Journal of Food Microbiology. 2004;**96**:281-288. DOI: 0.1016/j.ijfoodmicro.2004.04.002

[42] Bordbar F, Etebarian H, Sahebani N, Rohani H. Control of postharvest decay of apple fruit with *Trichoderma virens* isolates and induction of defense responses. Journal of Plant Protection Research. 2010;**50**:146-152. DOI: 0.2478/ v10045-010-0025-1

[43] Mortuza M, Ilag L. Potential for biocontrol of *Lasiodiplodia theobromae* (Pat.) Griff. & Maubl. in banana fruits by *Trichoderma* species. Biological Control. 1999;**15**:235-240. DOI: 0.1006/ bcon.1999.0716

[44] Quaglia M, Ederli L, Pasqualini S, Zazzerini A. Biological control agents and chemical inducers of resistance for postharvest control of *Penicillium expansum* Link. on apple fruit. Postharvest Biology and Technology. 2011;**59**:307-315. DOI: 0.1016/j. postharvbio.2010.09.007

[45] Dal Bello G, Lampugnani G, Abramoff C, Fusé C, Perelló A. Postharvest control of *Botrytis* gray mould in tomato by antagonists and biorational compounds. IOBC-WPRS Bulletin. 2015;**111**:417-425

[46] Guédez C, Cañizález L, Castillo C, Olivar R. Efecto antagónico de *Trichoderma harzianum* sobre algunos hongos patógenos postcosecha de la fresa (Fragaria spp). Revista de la Sociedad Venezolana de Microbiología. 2009;**29**:34-38

[47] Zimand G, Elad Y, Chet I. Effect of *Trichoderma harzianum* on *Botrytis cinerea* pathogenicity. Phytopathology. 1996;**86**:1255-1260

[48] Tronsmo A, Raa J. Antagonistic action of *Trichoderma pseudokoningii* against the apple pathogen *Botrytis cinerea*. Journal of Phytopathology. 1977;**89**:216-220

[49] Hjeljord L, Tronsmo A. *Trichoderma* and *Gliocladium* in bilogical control: A review. In: Harman GE, Kubicek CP, editors. *Trichoderma* & *Gliocladium*— Enzymes, Biological Control and Commercial Applications. 1st. ed. CRC Press, Taylor & Francis; 1998. pp. 131- 145

[50] Ahmad J, Baker R. Rhizosphere competence of *Trichoderma harzianum*. Phytopathology. 1987;**77**:182-189

[51] Lorito M, Harman G, Di Prieto A, Hayes C. Extracellular chitimolytie enzymes produced by *T. harzianum*, purification, characterization and molecular cloning. Phytopathology. 1990;**82**:10-77

[52] Carsolio C, Benhamou N, Haran S, Cortés C, Gutiérrez A, Chet I, et al. Role of the *Trichoderma harzianum* endochitinase gene, ech42, in mycoparasitism. Applied and Environmental Microbiology. 1999;**65**:929-935

[53] Mantyla A, Paloheimo M. Suominen Industrial mutants and recombinant strains of *Trichoderma* reesei. In: Harman GE, Kubicek CP, editors. *Trichoderma* & *Gliocladium*— Enzymes, Biological Control and Commercial Applications. 1st. ed. CRC Press, Taylor & Francis; 1998. pp. 291-304

[54] Demain A, Adrio J. Contributions of microorganisms to industrial biology. Molecular Biotechnology. 2008;**38**:41. DOI: 0.1007/ s12033-007-0035-z

[55] Goldman G, Hayes C, Harman G. Molecular and cellular biology of biocontrol by *Trichoderma* spp. Trends in Biotechnology. 1994;**12**:478-482

[56] Harman G, Howell C, Viterbo A, Chet I, Lorito M. *Trichoderma* species—Opportunistic, avirulent plant symbionts. Nature Reviews. Microbiology. 2004;**2**:43

[57] Chávez García M. Producción de *Trichoderma* sp. y evaluación de suefecto en cultivo de crisantemo (Dendranthema grandiflora). 2006. Available from: https:// repository.javeriana.edu.co/ bitstream/handle/10554/8312/ tesis286.pdf?sequence=1. [Review date:November 2018]

[58] Karas P, Perruchon C, Exarhou K, Ehaliotis C, Karpouzas D. Potential for bioremediation of agro-industrial effluents with high loads of pesticides by selected fungi. Biodegradation. 2011;**22**:215-228. DOI: 0.1007/s10532- 010-9389-1

[59] Fernández-Larrea O. Los microorganismos en el control biológico. Alternativas de producción en Cuba. Memorias del Curso Int. Manejo Agroecol. Plagas en el Sist. Prod. Inisav. 2006

[60] Cárdenas YG. Métodos de conservación y formulación de *Trichoderma harzianum* Rifai. Fitosanidad. 2010;**14**:189-195

[61] Usall J, Torres R, Teixido N. Biological control of postharvest diseases on fruit: A suitable alternative? Current Opinion in Food Science. 2016;**11**:51-55. DOI: 0.1016/j.cofs.2016.09.002

[62] Woo S, Scala F, Ruocco M, Lorito M. The molecular biology of the interactions between *Trichoderma* spp., phytopathogenic fungi, and plants. Phytopathology. 2006;**96**:181-185. DOI: 0.1094/PHYTO-96-0181

[63] Brunner K, Zeilinger S, Ciliento R, Woo S, Lorito M, Kubicek C, et al. Improvement of the fungal biocontrol agent *Trichoderma atroviride* to enhance both antagonism and induction of plant systemic disease resistance. Applied and Environmental Microbiology. 2005;**71**:3959-3965. DOI: 10.1128/ AEM.71.7.3959-3965.2005

[64] Whipps J, Lumsden R. Commercial use of fungi as plant disease biological control agents: Status and prospects. In: Butt TM, Jackson C, Magan N, editors. Fungi as Biocontrol Agents: Progress, Problems and Potential. Wallingford: CABI Publishing; 2001. pp. 9-22

[65] Marra R, Ambrosino P, Carbone V, Vinale F, Woo S, Ruocco M, et al. Study of the three-way interaction between *Trichoderma atroviride*, plant and fungal pathogens by using a proteomic approach. Current Genetics. 2006;**50**:307-321. DOI: 0.1007/ s00294-006-0091-0

**77**

2016;**9**:10-24

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma*

[72] Aml A, Ezzat A, Rageh M, Saber W,

[73] Lorito M, Woo S, D'ambrosio M, Harman G, Hayes C, Kubicek C, et al. Molecular Plant-Microbe Interactions.

[74] Alvindia D. Sodium bicarbonate enhances efficacy of *Trichoderma harzianum* DGA01 in controlling crown rot of banana. Journal of General Plant Pathology. 2013;**79**:136-144. DOI:

[75] Elamawi R, Al-Harbi R, Hendi A. Biosynthesis and characterization of silver nanoparticles using *Trichoderma longibrachiatum* and their effect on phytopathogenic fungi. Egyptian Journal of Biological Pest Control. 2018;**28**:28. DOI: 10.1186/

0.1007/s10327-013-0432-z

s41938-018-0028-1

1996;**9**:206-213

El-Sheikh T. Effect of chitosan, biocontrol agents and hot air to reduce postharvest decay and microbial loads of strawberries. Development. 2017;**6**:8

*DOI: http://dx.doi.org/10.5772/intechopen.82784*

[66] Ramos-Guerrero A, González-Estrada R, Hanako-Rosas G, Bautista-Baños S, Acevedo-Hernández G, Tiznado-Hernández M, et al. Use of inductors in the control of *Colletotrichum gloeosporioides* and *Rhizopus stolonifer* isolated from soursop fruits: In vitro tests. Food Science and Biotechnology. 2018;**27**:755-763. DOI:

0.1007/s10068-018-0305-5

0.1007/BF02983969

biocontrol.2015.02.001

[68] Sangeetha G, Usharani S, Muthukumar A. Biocontrol with *Trichoderma* species for the management of postharvest crown rot of banana. Phytopathologia Mediterranea. 2009;**48**:214-225. DOI: 10.14601/Phytopathol\_Mediterr-2741

[69] Panebianco S, Vitale A, Polizzi G, Scala F, Cirvilleri G. Enhanced control of postharvest citrus fruit decay by means of the combined use of compatible biocontrol agents. Biological Control. 2015;**84**:19-27. DOI: 0.1016/j.

[70] Senthil R, Prabakar K, Rajendran L, Karthikeyan G. Efficacy of different biological control agents against major postharvest pathogens of grapes under room temperature storage conditions. Phytopathologia Mediterranea. 2011;**50**:55-64. DOI: 10.14601/ Phytopathol\_Mediterr-3115

[71] Nitu N, Masum M, Jannat R, Bhuiyan M, Sultana S. Application of chitosan and *Trichoderma* against soil-borne pathogens and their effect on yield of tomato (*Solanum lycopersicum* L.). International Journal of Biosciences.

[67] Sivakumar D, Wijeratnam R, Abeyesekere M, Wijesundera R. Combined effect of generally regarded as safe (GRAS) compounds and *Trichoderma harzianum* on the control of postharvest diseases of rambutan. Phytoparasitica. 2002;**30**:43-51. DOI:

*A Review Study on the Postharvest Decay Control of Fruit by Trichoderma DOI: http://dx.doi.org/10.5772/intechopen.82784*

[66] Ramos-Guerrero A, González-Estrada R, Hanako-Rosas G, Bautista-Baños S, Acevedo-Hernández G, Tiznado-Hernández M, et al. Use of inductors in the control of *Colletotrichum gloeosporioides* and *Rhizopus stolonifer* isolated from soursop fruits: In vitro tests. Food Science and Biotechnology. 2018;**27**:755-763. DOI: 0.1007/s10068-018-0305-5

*Trichoderma - The Most Widely Used Fungicide*

[52] Carsolio C, Benhamou N, Haran S,

[59] Fernández-Larrea O. Los microorganismos en el control

[60] Cárdenas YG. Métodos de conservación y formulación de *Trichoderma harzianum* Rifai. Fitosanidad. 2010;**14**:189-195

[61] Usall J, Torres R, Teixido N.

DOI: 0.1016/j.cofs.2016.09.002

The molecular biology of the

Woo S, Lorito M, Kubicek C, et al. Improvement of the fungal biocontrol agent *Trichoderma* 

AEM.71.7.3959-3965.2005

0.1094/PHYTO-96-0181

Biological control of postharvest diseases on fruit: A suitable alternative? Current Opinion in Food Science. 2016;**11**:51-55.

[62] Woo S, Scala F, Ruocco M, Lorito M.

interactions between *Trichoderma* spp., phytopathogenic fungi, and plants. Phytopathology. 2006;**96**:181-185. DOI:

[63] Brunner K, Zeilinger S, Ciliento R,

*atroviride* to enhance both antagonism and induction of plant systemic disease resistance. Applied and Environmental Microbiology. 2005;**71**:3959-3965. DOI: 10.1128/

[64] Whipps J, Lumsden R. Commercial use of fungi as plant disease biological control agents: Status and prospects. In: Butt TM, Jackson C, Magan N, editors. Fungi as Biocontrol Agents: Progress, Problems and Potential. Wallingford: CABI Publishing; 2001. pp. 9-22

[65] Marra R, Ambrosino P, Carbone V, Vinale F, Woo S, Ruocco M, et al. Study of the three-way interaction between *Trichoderma atroviride*, plant and fungal pathogens by using a proteomic approach. Current Genetics.

2006;**50**:307-321. DOI: 0.1007/

s00294-006-0091-0

2006

biológico. Alternativas de producción en Cuba. Memorias del Curso Int. Manejo Agroecol. Plagas en el Sist. Prod. Inisav.

Cortés C, Gutiérrez A, Chet I, et al. Role of the *Trichoderma harzianum* endochitinase gene, ech42, in mycoparasitism. Applied and Environmental Microbiology.

[53] Mantyla A, Paloheimo M. Suominen Industrial mutants and recombinant strains of *Trichoderma* reesei. In: Harman GE, Kubicek CP, editors. *Trichoderma* & *Gliocladium*— Enzymes, Biological Control and Commercial Applications. 1st. ed. CRC Press, Taylor & Francis; 1998.

[54] Demain A, Adrio J. Contributions of

[55] Goldman G, Hayes C, Harman G. Molecular and cellular biology of biocontrol by *Trichoderma* spp. Trends in Biotechnology. 1994;**12**:478-482

[56] Harman G, Howell C, Viterbo A, Chet I, Lorito M. *Trichoderma* species—Opportunistic, avirulent plant symbionts. Nature Reviews.

[57] Chávez García M. Producción de *Trichoderma* sp. y evaluación de suefecto en cultivo de crisantemo (Dendranthema grandiflora). 2006. Available from: https:// repository.javeriana.edu.co/ bitstream/handle/10554/8312/ tesis286.pdf?sequence=1. [Review

[58] Karas P, Perruchon C, Exarhou K, Ehaliotis C, Karpouzas D. Potential for bioremediation of agro-industrial effluents with high loads of pesticides by selected fungi. Biodegradation. 2011;**22**:215-228. DOI: 0.1007/s10532-

microorganisms to industrial biology. Molecular Biotechnology.

2008;**38**:41. DOI: 0.1007/ s12033-007-0035-z

Microbiology. 2004;**2**:43

date:November 2018]

1999;**65**:929-935

pp. 291-304

**76**

010-9389-1

[67] Sivakumar D, Wijeratnam R, Abeyesekere M, Wijesundera R. Combined effect of generally regarded as safe (GRAS) compounds and *Trichoderma harzianum* on the control of postharvest diseases of rambutan. Phytoparasitica. 2002;**30**:43-51. DOI: 0.1007/BF02983969

[68] Sangeetha G, Usharani S, Muthukumar A. Biocontrol with *Trichoderma* species for the management of postharvest crown rot of banana. Phytopathologia Mediterranea. 2009;**48**:214-225. DOI: 10.14601/Phytopathol\_Mediterr-2741

[69] Panebianco S, Vitale A, Polizzi G, Scala F, Cirvilleri G. Enhanced control of postharvest citrus fruit decay by means of the combined use of compatible biocontrol agents. Biological Control. 2015;**84**:19-27. DOI: 0.1016/j. biocontrol.2015.02.001

[70] Senthil R, Prabakar K, Rajendran L, Karthikeyan G. Efficacy of different biological control agents against major postharvest pathogens of grapes under room temperature storage conditions. Phytopathologia Mediterranea. 2011;**50**:55-64. DOI: 10.14601/ Phytopathol\_Mediterr-3115

[71] Nitu N, Masum M, Jannat R, Bhuiyan M, Sultana S. Application of chitosan and *Trichoderma* against soil-borne pathogens and their effect on yield of tomato (*Solanum lycopersicum* L.). International Journal of Biosciences. 2016;**9**:10-24

[72] Aml A, Ezzat A, Rageh M, Saber W, El-Sheikh T. Effect of chitosan, biocontrol agents and hot air to reduce postharvest decay and microbial loads of strawberries. Development. 2017;**6**:8

[73] Lorito M, Woo S, D'ambrosio M, Harman G, Hayes C, Kubicek C, et al. Molecular Plant-Microbe Interactions. 1996;**9**:206-213

[74] Alvindia D. Sodium bicarbonate enhances efficacy of *Trichoderma harzianum* DGA01 in controlling crown rot of banana. Journal of General Plant Pathology. 2013;**79**:136-144. DOI: 0.1007/s10327-013-0432-z

[75] Elamawi R, Al-Harbi R, Hendi A. Biosynthesis and characterization of silver nanoparticles using *Trichoderma longibrachiatum* and their effect on phytopathogenic fungi. Egyptian Journal of Biological Pest Control. 2018;**28**:28. DOI: 10.1186/ s41938-018-0028-1

**79**

**Chapter 5**

**Abstract**

secondary metabolite

**1. Introduction**

*boninense*

A Review Report on the

*Elaeis guineensis*

*Syed Ali Nusaibah and Habu Musa*

Mechanism of *Trichoderma* spp.

as Biological Control Agent of the

Basal Stem Rot (BSR) Disease of

*Trichoderma* spp. have been the most common fungi applied as biological control agents (BCA) as an effort to combat a wide range of plant diseases. Its uses have recorded good success rate in controlling major plant diseases. Knowledge on the mechanisms employed by *Trichoderma* spp. could be further studied to improve its ability as an efficient biocontrol agent. The *Trichoderma* ability to curb plant diseases were mainly based on the activation of single or multiple control mechanisms. It is known that the *Trichoderma*-based biocontrol mechanisms mainly rely on mycoparasitism, production of antibiotic and/or hydrolytic enzymes, competition for nutrients, as well as induced plant resistance; numerous secondary metabolites produced by *Trichoderma* species could directly inhibit the growth of several plant pathogens. These mechanisms may act directly or indirectly against the targeted plant pathogen. This chapter reviews the recent updates on published research findings on mechanisms used by *Trichoderma* as biological control of plant diseases

particularly on basal stem rot disease of oil palm caused by *Ganoderma* spp.

**1.1 Biological control of oil palm basal stem rot disease caused by** *Ganoderma* 

Many promising biological antagonists, mainly from *Trichoderma*, *Aspergillus*, *Penicillium*, *Pseudomonas*, and *Bacillus*, have been reported as effective antagonists against *Ganoderma boninense* in coconut [1] and oil palm [2–4]. In 1990, [5] evaluated the incorporation of *Trichoderma* spp. grown on dried palm oil mill effluent into planting holes as a prophylactic measure. Later, [6] reported delays in infection in the field following treatment with *Trichoderma*, but eventually, the disease incidence was similar to untreated controls. Thus, the possible explanation for this could be due to a low natural occurrence of *Trichoderma*

**Keywords:** antibiosis, competition, induced resistance, mycoparasitism,

### **Chapter 5**
