Phytopathogenic organisms and mycotoxigenic fungi: Why do we control one and neglect the other? A biological control perspective in Malaysia

Siti Nur Ezzati Yazid; Selamat Jinap; Siti Izera Ismail; Naresh Magan; Nik Iskandar Putra Samsudin

This is an open access article & it is licensed under a Creative Commons Attribution Non-Commercial 4.0 International (https://creativecommons.org/licenses/by-nc/4.0/) Multicellular filamentous fungi grown on the surface and inside of moist food secretes toxins in the form of their secondary metabolites which are commonly called mycotoxins. The presence of mycotoxins in food has been a burning issue and a threat to food security and safety. The global population has skyrocketed and continues to be, which has created a challenge of providing quality food to the consumers. Aflatoxins, prevalent in most of the food crops in Nepal as well have posed a risk to national food security. Moreover, the consumption of food products containing mycotoxins is a cause of several health hazards like cancer, gastrointestinal problems, and neuropsychiatric effects. Mycotoxins not only has affected humans but also animals. Prevention, decontamination, and inhibition of absorption of toxins have been done to manage and mitigate the effects of mycotoxins. Recent research on mycotoxins is focused on the development of new methods to detect and analyse masked mycotoxins obtained from various sources. This review shows the contribution of mycotoxin in the global food security issue as well as its deleterious effects on human and animal health.

During the last decade, scientists have given increasingly frequent warnings about global warming, linking it to mycotoxin-producing moulds in various geographical regions across the world. In the future, more pronounced climate change could alter host resilience and host–pathogen interaction and have a significant impact on the development of toxicogenic moulds and the production of their secondary metabolites, known as mycotoxins. The current climate attracts attention and calls for novel diagnostic tools and notions about the biological features of agricultural cultivars and toxicogenic moulds. Since European climate environments offer steadily rising opportunities for Aspergillus flavus growth, an increased risk of cereal contamination with highly toxic aflatoxins shall be witnessed in the future. On top of that, the profile (representation) of certain mycotoxigenic Fusarium species is changing ever more substantially, while the rise in frequency of Fusarium graminearum contamin...

Contamination of crops with phytopathogenic genera such as Fusarium, Aspergillus, Alternaria, and Penicillium usually results in mycotoxins in the stored crops or the final products (bread, beer, etc.). To reduce the damage and suppress the fungal growth, it is common to add antifungal substances during growth in the field or storage. Many of these antifungal substances are also harmful to human health and the reduction of their concentration would be of immense importance to food safety. Many eminent researchers are seeking a way to reduce the use of synthetic antifungal compounds and to implement more eco-friendly and healthier bioweapons against fungal proliferation and mycotoxin synthesis. This paper aims to address the recent advances in the effectiveness of biological antifungal compounds application against the aforementioned fungal genera and their species to enhance the protection of ecological and environmental systems involved in crop growing (water, soil, air) and to red...

Received: 25 August 2019 Revised: 3 January 2020 Accepted: 8 January 2020 DOI: 10.1111/1541-4337.12541 COMPREHENSIVE REVIEWS IN FOOD SCIENCE AND FOOD SAFETY Phytopathogenic organisms and mycotoxigenic fungi: Why do we control one and neglect the other? A biological control perspective in Malaysia Siti Nur Ezzati Yazid1 Selamat Jinap1,2 Siti Izera Ismail3,4 Naresh Magan5 Nik Iskandar Putra Samsudin1,2 1 Laboratory of Food Safety and Food Integrity, Institute of Tropical Agriculture and Food Security, Universiti Putra Malaysia, Serdang, Malaysia 2 Department Abstract In this review, we present the current information on development and applications of biological control against phytopathogenic organisms as well as mycotoxigenic fungi of Food Science, Faculty of Food Science and Technology, Universiti Putra Malaysia, Serdang, Malaysia in Malaysia as part of the integrated pest management (IPM) programs in a collective effort to achieve food security. Although the biological control of phytopathogenic 3 Laboratory organisms of economically important crops is well established and widely practiced in Malaysia with considerable success, the same cannot be said for mycotoxigenic of Climate-Smart Food Crop Production, Institute of Tropical Agriculture and Food Security, Universiti Putra Malaysia, Serdang, Malaysia 4 Department of Plant Protection, Faculty of Agriculture, Universiti Putra Malaysia, Serdang, Malaysia 5 Applied Mycology Group, Cranfield Soil and AgriFood Institute, Cranfield University, Cranfield, UK Correspondence Nik Iskandar Putra Samsudin, Laboratory of Food Safety and Food Integrity, Institute of Tropical Agriculture and Food Security, Universiti Putra Malaysia, 43400 UPM Serdang, Selangor, Malaysia. Email: nikiskandar@upm.edu.my Funding information Malaysian Ministry of Education (MOE) under the High Impact Centre of Excellence (HICoE) grant scheme, Grant/Award Number: HICoE/ITAFoS/2017/FS9 fungi. This is surprising because the year round hot and humid Malaysian tropical climate is very conducive for the colonization of mycotoxigenic fungi and the potential contamination with mycotoxins. This suggests that less focus has been made on the control of mycotoxigenic species in the genera Aspergillus, Fusarium, and Penicillium in Malaysia, despite the food security and health implications of exposure to the mycotoxins produced by these species. At present, there is limited research in Malaysia related to biological control of the key mycotoxins, especially aflatoxins, Fusarium-related mycotoxins, and ochratoxin A, in key food and feed chains. The expected threats of climate change, its impacts on both plant physiology and the proliferation of mycotoxigenic fungi, and the contamination of food and feed commodities with mycotoxins, including the discovery of masked mycotoxins, will pose significant new global challenges that will impact on mycotoxin management strategies in food and feed crops worldwide. Future research, especially in Malaysia, should urgently focus on these challenges to develop IPM strategies that include biological control for minimizing mycotoxins in economically important food and feed chains for the benefit of ensuring food safety and food security under climate change scenarios. KEYWORDS Aspergillus, biocontrol, climate change, Fusarium, mycotoxin, Penicillium 1 BIOLOGICA L CONTROL AND FOOD SECUR I T Y Feeding the global population of nearly eight billion people is indeed the most daunting challenge facing mankind in the Compr Rev Food Sci Food Saf. 2020;19:643–669. modern world. Food security, which is defined as “…when all people, at all times, have physical, social, and economic access to sufficient, safe, and nutritious food that meets their dietary needs and food preferences to support an active and healthy life…,” is the primary concern in addressing this wileyonlinelibrary.com/journal/crf3 © 2020 Institute of Food Technologists® 643 644 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… challenge (FAO, 2009). Since time immemorial, agriculture and aquaculture have been, and still are, the primary nutritional support to mankind, and act as the first pillar of food security, which is “food availability” (that is, supply of food through production, distribution, and exchange). The other three pillars according to Gregory, Ingram, and Brklacich (2005) are “food accessibility” (that is, affordability and allocation of food, as well as the preferences of individuals and households), “food utilization” (that is, consumption of safe, quality, and sufficient food for human health and metabolism), and “food stability” (that is, ability to obtain food over time). However, the continual growth in the human population and competition for land, water, and energy will eventually and inevitably impair our agricultural ability and capacity to produce sufficient food to feed this increasing global demand (Godfray et al., 2010). What is more, the global agricultural sector is also constantly left at the mercy of its old arch enemy—pests and pathogens! It was recently estimated that 40% to 50% of crop yields in developing countries are wastefully lost to pests and crop diseases pre- and postharvest. For developed countries such as the United States, the estimate stands at 20% to 25% (NIFA, 2015). Apart from pests and crop diseases, weeds can also thrive in monoculture systems, thus significantly impacting on the agricultural yields (Poggio, 2005). Pest is a broad term frequently used and defined as any species or strain of living organism (for example, animals, plants, and microorganisms), usually occurring in high density in a specific niche, which results in detrimental and injurious effects on humans, animals, or plants. In agriculture, pests are regarded as the key biotic component that can adversely impact on agricultural yields and productivity along the supply chain, with the major abiotic component being the climate. To mitigate this, the management or eradication of crop pests of economic importance has been practiced worldwide by using various control measures. There are even documents citing that pest control is as old as agriculture itself because it has always been necessary to keep the crops pest and disease free (Oerke, 2006). The various control measures of pests are generally divided into three—physical/mechanical, chemical, and biological, and have been employed either individually or collectively before, during, or after crop cultivation as part of an Integrated Pest Management (IPM) strategy and Good Agricultural Practice (GAP). Physical or mechanical pest control involves the use of simple equipment and devices as well as hands-on techniques that form a protective barrier between the crops and the pests (Khan & Ram, 2014; Sorensen, Mohankumar, & Thangaraj, 2016; Vincent, Hallman, Panneton, & Fleurat-Lessard, 2003). These include controlled irrigation to minimize stresses on the crops, crop rotation, crop monitoring, growing pest-resistant crop varieties/cultivars through selective breeding, intercropping, seed selection, soil ploughing, the use of physical bar- riers, nettings or trappings, timely harvesting, and trap cropping (that is, companion crops planted nearby the main crops [companion planting] to divert pests away from infesting the main crops; Holden, Ellner, Lee, Nyrop, & Sanderson, 2012). Chemical pest control, on the other hand, involves the use of chemically synthesized pesticides, which can be employed either as solid (for example, by luring the pests to eat chemically poisoned baits), liquid (for example, by spraying formulated pesticide suspensions onto the crops or soil), or gaseous (for example, by suffocating pests with chemically poisoned fumigants). Apart from chemicals, there are also plant-derived or plant-based control agents known as phytochemicals that pose lesser threat to the environments and human health as compared to their more recalcitrant chemically synthesized counterparts. As critically reviewed by Koul (2008), these phytochemicals, which include alkaloids, chromenes, cucurbitacins, cyclopropanoid acids, phenolics, polyacetylenes, quassinoids, saponins, terpenes, and their various derivatives, are usually applied as antifeedants and/or deterrents. Additionally, there are also semiochemicals (for example, pheromones and allelochemicals) that are defined as substances used in plant–insect or insect–insect interactions as complementary components to insecticide approaches in IPM. Semiochemicals are used to effectively modify insect pests’ behavior by affecting their survival and/or reproduction in the effort to control their infestations on crops (El-Ghany, 2019). Another alternative is biological control, or simply biocontrol, which is a technique to significantly reduce or permanently suppress the population density of pests to a level that has no significant effect on the crop yield by interfering with the pests’ ecophysiology using living organisms (Wyckhuys et al., 2013). If properly applied, biological control could be a long-term and self-sustaining treatment method for managing pests (Sahayaraj, 2014) that will consequently contribute to lower agro-operational costs. At present, there are four widely accepted fundamental strategies in biological control (van Lenteren, Bolckmans, Köhl, Ravensberg, & Urbaneja, 2018): (a) natural, in which natural enemies of the pests are not frequently noticed and consciously manipulated by humans but their presence actually helps to regulate the population density of their host or prey; (b) importation (classical), in which foreign/exotic natural enemies of the pests are intentionally reared and released into new habitats or ecosystems to regulate the population density of their host or prey; (c) augmentation, in which natural enemies of the pests, which are unable to survive and/or persist in the new habitats or ecosystems, are reared in large population and mass-released (inundative augmentation) or periodically released in smaller numbers (inoculative augmentation) to regulate the population density of their host or prey; and (d) conservation, in which natural enemies of the pests are aided (for example, BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… 645 FIGURE 1 The principles and approaches of Integrated Plant Protection (IPP) otherwise known as Integrated Pest Management (IPM) Note. Adapted from Frische et al. (2018) providing them with shelters and reducing the use of chemicals that might kill them) to maintain and optimize their survival and/or effectiveness in regulating the population density of their host or prey. Figure 1 summarizes the principles and approaches of Integrated Plant Protection (IPP) otherwise also frequently referred to as IPM (Frische, Egerer, Matezki, Pickl, & Wogram, 2018). The IPP or IPM pyramid in Figure 1 ideally works from the bottom up, in which preventive measures (for example, the use of tolerant/resistant crop cultivars; crop care such as farm irrigation and crop rotation; promoting natural antagonists against crop pests) always take priority and precedent before the use of chemicals. Only when the economic threshold is exceeded, as assessed by the risk analysis and monitoring, would the plant protection approach shift to biotechnological measures (for example, biological, physical, and chemical controls). An “economic threshold” is the pest’s population level infesting the farm or the extent of crop damage inflicted by the pests at which the value of the crop destroyed exceeds the cost of controlling the pest (Zalom, 2010). Of course as climate is in a state of flux and the pests rapidly evolve, these control measures will often have flaws and drawbacks and have to be appropriately modified. For example, the physical control on a much larger scale requires enormous time and manpower, which will most likely contribute to higher operational costs, whereas the chemical control has long been associated with high production and application costs, human health hazards, restriction by domestic and international regulatory limits, trade bans, residual effects, environmental pollution, resistance development in pests, and potential elimination of beneficial natural enemies of the targeted pests (Bommarco, Miranda, Bylund, & Björkman, 2011). Also, as illustrated in Figure 1, a good and sustainable IPM strategy always uses chemicals as the last resort. As for biological control, several drawbacks have also been highlighted such as they do not function as rapidly as chemical pesticides, their application requires special skills/knowledge, or even worse, they prey on beneficial organism thus disrupting the natural food chain. To mitigate the latter, care and consideration should be taken into account when developing an efficient and successful biological control system. Simberloff in 2012 outlined four potential risks when introducing a biological control agent: (a) direct attack on nontargets; (b) indirect effects on nontargets (for example, competition); (c) dispersal of the introduced biological control agent to other area; and (d) changed relationships between the introduced biological control agent and the native species (for example, when the introduced biological control agent multiply uncontrollably and becomes invasive) (Simberloff, 2012). Of course the risk of nontarget is always there (also known as the “nontarget effect”; Louda, Pemberton, Johnson, & Follett, 2003) and has been heavily debated; that is why risk analysis and assessment (for example, host specificity test; Kaufman & Wright, 2017) must be critically performed before a biological control agent is introduced in the crop agro-ecosystem. Köhl, Postma, Nicot, Ruocco, and Blum (2011) proposed BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Production (metric tonne) 646 73.8% 20,000,000 15,000,000 10,000,000 5,000,000 10.2% 5.5% 0.8% 0.2% 0.1% 3.7% 0.1% 2.7% 2.9% 0 Crop commodities FIGURE 2 Malaysian agricultural production for the year 2017 Note. Adapted from DOA (2019). a 2015 data. b “Major industrial crops,” as managed by the Malaysian Ministry of Primary Industries. Data for other “major industrial crops” such as timber, kenaf (Hibiscus cannabinus), and tobacco (Nicotiana tabacum) are not publicly available. “Other industrial crops” and other crop commodities are managed by the Malaysian Ministry of Agriculture and Agro-based Industry. “Cash crops” include cassava (Manihot esculenta), groundnut (Arachis hypogaea), maize (Zea mays), sugar cane (Saccharum officinarum), sweet potato (Ipomoea batatas), taro (Colocasia esculenta), and yam bean (Pachyrhizus erosus). “Other industrial crops” include areca nut (Areca catechu), coconut (Cocos nucifera), coffee (Coffea arabica), mushroom (various species), nipa palm (Nypa fruticans), roselle (Hibiscus sabdariffa), sago (Metroxylon sagu), and tea (Camellia sinensis) nine sequential steps in selecting, developing, and introducing a biological control agent of plant pathogens: (a) assessment of targeted crop, disease, and markets, (b) isolation of candidate antagonists, (c) rapid-throughput screening in terms of production, safety, and ecology, (d) database mining in terms of intellectual property protection, safety, ecology, environmental risks, and marketing, (e) efficacy testing in bioassays, (f) preliminary assessment of mass production, (g) pilot formulation, (h) upscaling mass production and full field testing, and (i) integration into cropping systems with constant monitoring of its efficacy and environmental risks. These are critically performed to prevent the disadvantages of introducing a biological control agent outweighing its advantages. Furthermore, as previously mentioned, biological control is never applied individually but collectively with other measures as part of IPM or GAP programs. In Malaysia, agriculture is the third engine that drives and sustains the economic growth after services and manufacturing in terms of gross domestic product (GDP), with oil palm (Elaeis guineensis) and rice (Oryza sativa) dominating the production scene (Fahmi, Abu-Samah, & Abdullah, 2013). Although the industrial commodity subsector (for example, cocoa, palm oil, rubber, and timber) serves as the primary contributor to the sector’s GDP, the growth rate of the food commodity subsector (for example, fruits, rice, and vegetables) is gradually increasing, which indicates a positive improve- ment of the domestic food production while reducing import dependency (Ahmad & Suntharaligam, 2009). Moreover, the approach recently undertaken by the government to further increase the domestic food production to feed the 32 million population has shifted from the expansion of land for animal rearing and crop cultivation to sustainable agricultural intensification in which farmers are encouraged to use fewer inputs (for example, energy, water, nutrients, and pesticides) and achieve higher productivity within similar acreage of land. Figure 2 illustrates the agricultural production in Malaysia in 2017. Like many other countries, Malaysian agricultural productivity is also impacted by pest infestation, against which the country has launched numerous measures in a collective effort to combat further deterioration. Among such measures is biological control. Biological control has a long history in Malaysia. The earliest available online resource dates back to 40 years ago in 1978 on the use of the beetle Metrogaleruca obscura and the wasp Eurytoma attiva to control the invasive weed Cordia curassavica (tropical black sage) through defoliation and seed destruction, respectively (Simmonds, 1980). Since then, Malaysian agricultural authorities, particularly the Department of Agriculture (DOA) and Malaysian Agricultural Research and Development Institute (MARDI), both under the jurisdiction of the Ministry of Agriculture and Agro-based Industry, have been conducting numerous research initiatives BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… to combat pest infestation on economically important crops such as oil palm and rice paddy. Although it is understandable why the focus has always been, and still is, on the control of phytopathogenic organisms, it must not be forgotten that pests come in different forms. Since the discovery of T-2 toxin that caused alimentary toxic aleukia in Russia in the 1940s (Joffe, 1971; Williams, 1989), and aflatoxins that killed nearly 100,000 turkey poults in England in the 1960s (Blount, 1961; Wannop, 1961), the global scientific community has been alarmed by the disease- and death-causing capability of fungal secondary metabolites, which can affect both humans and animals (Pitt & Miller, 2017). These secondary metabolites were henceforth termed mycotoxins (from the Greek mykes, which is fungus, and toxikon, which is poison), and their fungal producers are known as mycotoxigenic fungi. Being produced by several genera of microfungi, mycotoxin contamination is in fact the secondary outcome of fungal infestation of crop commodities, with the primary ones being the damages and diseases such as blights, rusts, rots, and wilts. Although just a secondary outcome, the clinical effects of mycotoxins on humans and animals are certain with several types of mycotoxins being classified as carcinogens (International Agency for Research on Cancer [IARC]: https://www.iarc.fr/; Ostry, Malir, Toman, & Grosse, 2017). The ecophysiological conditions influencing mycotoxigenic fungal growth and mycotoxin production as well as major mycotoxins and their associated health risks on humans and animals have been extensively reviewed elsewhere (Bhat, Rai, & Karim, 2010). Nevertheless, despite Malaysian climatic conditions (that is, tropical; warm and humid all year round) being highly favorable for fungal proliferation, including that of mycotoxigenic fungi, on staple crop commodities pre- and postharvest, to the best of our knowledge thus far, there has been very little, if any, research related to the use of biological control agents to control mycotoxigenic fungi and their associated mycotoxin contamination in Malaysia. The present review therefore discusses (a) the application of biological control agents in regulating the infestation of phytopathogenic organisms in Malaysia, (b) the current status of biological control against mycotoxigenic fungi in Malaysia and the potential reasons for their lack of use, and (c) the future directions of biological control research in Malaysia, with a focus on mycotoxigenic fungi and mycotoxin control. 2 BIOLOGICA L CONTROL AGAINST PH Y TOPAT H O G E N I C ORGANISMS Although regarded as a newly industrialized country (that is, transitional phase between developing and developed countries), Malaysia still relies on the agriculture sector, which 647 is the nation’s third economic engine. The First (1984 to 1991), Second (1992 to 1997), and Third National Agricultural (1998 to 2010) Policies helped to shape the sector to be productive, sustainable, and competitive in the face of agricultural challenges in a rapidly changing world. Under the various long-term and far-sighted initiatives undertaken by the Ministry of Agriculture and Agro-based Industry, crops are being cultivated on a larger scale and are usually well managed. Approaches such as IPM and GAP—which among others include regular monitoring, timely harvesting, supplementation and utilization of natural control agents, and the prudent use of chemicals—are being put into practice in order to improve yield, safety, and quality of the agricultural products. As part of a good IPM, biological control of phytopathogenic organisms has been widely applied in Malaysia with all the four main biological control strategies (that is, natural, importation, augmentation, and conservation) have been documented. The role of natural enemies in controlling native pests has been recognized in economically important crops especially oil palm, cocoa, coconut, and rice. The introduction of parasitoid wasps Diadegma semiclausum and Diadromus collaris in the 1970s to control Plutella xylostella (diamondback moth whose larvae destructively infest cruciferous vegetables especially cabbages) had suppressed the drawbacks of the reduction in chemical pesticide usage (Ooi, 1992; Ooi & Lim, 1989). The introduction of beneficial plants such as Cassia cobanensis (cassia), Turnera subulata (white buttercup), and Antigonon leptopus (Mexican creeper) has provided shelter and supplementary food such as nectar, and encouraged the proliferation of predators and parasites (Jamian, 2017). The conservation of natural or introduced enemies has also been important in insect pest management in Malaysia. Table 1 summarizes almost 40 years of research on the biological control agents, their targeted pests/pathogens/diseases, and their potential mechanisms of action against phytopathogenic organisms in Malaysia. Figure 3 further illustrates the distribution of the targeted crop commodities on which research on potential biological control agents against phytopathogenic organisms has been focused. From Figure 3, it is apparent that oil palm—being the country’s foremost crop in the agriculture sector—evidently receives appreciable attention and emphasis in terms of protection and prevention against diseases. The major pests of oil palm in Malaysia include bagworm moths (Metisa plana and Pteroma pendula), bunch moths (Tirathaba rufivena), rats, rhinoceros beetles (Oryctes rhinoceros), and termites (Coptotermes curvignathus), whereas the most important diseases are Ganoderma basal stem rot (caused by Ganoderma boninense) and Marasmius bunch rot (caused by Marasmius palmivorus) (SALCRA, 2019). To mitigate the Ganoderma basal stem rot, the Malaysian Palm Oil Board (MPOB), a government agency responsible for the research and development of palm oil industry in Malaysia, has introduced the Integrated BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… 648 TABLE 1 Available publications on the use of biological control agents against phytopathogenic organisms in Malaysia (1980–2019) Targeted pest/pathogen/disease Application/potential mechanism of action Beauveria bassiana Atteva sciodoxa (tiger moth) whose larvae infest long jack Spraying of conidial suspension targeting larvae Abood et al., 2010 Beauveria bassiana and Paecilomyces fumosoroseus Plutella xylostella (diamondback moth) whose larvae infest cabbages Spraying of conidial suspension Ibrahim & Low, 1993 Exserohilum longirostratum Echinochloa crus-galli (barnyard grass), weed infesting rice paddy fields Spraying of conidial suspension Ng et al., 2012 Exserohilum monoceras Echinochloa crus-galli (barnyard grass), weed infesting rice paddy fields Spraying of conidial suspension Kadir et al., 2008 Exserohilum prolatum Rottboellia cochinchinensis (itch grass), a noxious weed impacting crops Soil treatment (spraying) with conidial suspension Alloub et al., 2009 Exserohilum rostratum Cyperus iria (flat sedge), weed infesting rice paddy fields Spraying of conidial suspension Kundat et al., 2010 Fusarium oxysporum (non-pathogenic) Fusarium oxysporum f. sp. cubense (ascomycete) causing Fusarium wilt in bananas As an endophyte Ting et al., 2009 Isaria fumosorosea Coptotermes curvignathus and Coptotermes gestroi (termites) infesting trees Conidia germinate and penetrate into the termite cuticle Jessica et al., 2019 Isaria fumosorosea and Metarhizium anisopliae Atteva sciodoxa (tiger moth) whose larvae infest long jack Spraying of conidial suspension Sajap et al., 2014 Metarhizium anisopliae var. anisopliae Coptotermes curvignathus (termite) infesting oil palms Conidia germinate and penetrate into the termite cuticle Hoe et al., 2009 Metarhizium anisopliae var. major Oryctes rhinoceros (rhinoceros beetle) attacking oil palm fronds Granules of mycelia and spores attacking beetle larvae Moslim, Kamarudin, Wahid, 2011; Moslim et al., 2009 Paecilomyces carneus and Paecilomyces farinosus Pteroma pendula (bagworm moth) whose larvae infest oil palms Spraying of conidial suspension targeting larvae Bakeri et al., 2009 Paecilomyces lilacinus Meloidogyne incognita (nematode) causing root-knot in black peppers Spraying of conidial suspension Pau et al., 2012 Trichoderma harzianum Fusarium proliferatum and F. verticillioides (ascomycete) causing Fusarium ear rot in maize Production of volatile and nonvolatile inhibitory compounds Suhaida & NurAinIzzati, 2013 Trichoderma harzianum Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Production of volatile and nonvolatile inhibitory compounds Siddiquee et al., 2009; Nur Ain Izzati & Abdullah, 2008; Sundram et al., 2008 Trichoderma harzianum Macrophomina phaseolina (ascomycete) causing charcoal rot disease in soybeans Production of volatile and nonvolatile inhibitory compounds Khalili et al., 2016 Trichoderma virens Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Production of volatile and nonvolatile inhibitory compounds Angel et al., 2016 Trichoderma sp. Botryodiplodia theobromae (ascomycete) causing stem-end rot in mangos Fruit treatment (spraying) with conidial suspension Suhanna et al., 2013 Trichoderma spp. Fusarium fujikuroi (ascomycete) causing bakanae disease in rice Production of volatile and nonvolatile inhibitory compounds Ng et al., 2015 Zoophthora radicans Plutella xylostella (diamondback moth) whose larvae infest cabbages Spraying of conidial suspension Furlong et al., 1995 Biological control agent Reference (a) Ascomycete (Continues) BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… TABLE 1 649 (Continued) Targeted pest/pathogen/disease Application/potential mechanism of action Clitopilus prunulus, Grammothele fuligo, Lentinus tigrinus, Pycnoporus sanguineus, and Trametes lactinea Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Root treatment of oil palm seedlings Naidu et al., 2018 Ganoderma orbiforme, Neonothopanus nambi, and Schizophyllum commune Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Competition for substrate and space, and production of nonvolatile inhibitory compounds Naidu et al., 2016 Schizophyllum commune Wood-degrading fungi on rubber wood Production of inhibitory compounds Teoh et al., 2017; Peng & Don, 2013; Teoh et al., 2012 Colletotrichum gloeosporioides (ascomycete) causing anthracnose in papayas Production of hydrolytic enzymes and antimicrobial compounds Hamizah et al., 2013 Actinomycetes Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Production of inhibitory compounds Shariffah-Muzaimah et al., 2018; Nur Azura et al., 2016; ShariffahMuzaimah et al., 2015; Tan et al., 2002 Bacillus amyloliquefaciens and Pseudomonas pachastrellae Phytophthora capsici (oomycete) and Fusarium solani (ascomycete) infesting black peppers Production of extracellular hydrolytic enzymes Kota et al., 2015 Bacillus thuringiensis subsp. Kurstaki Metisa plana (bagworm moth) whose larvae infest oil palms Soil treatment (spraying) with bacterial suspension Salim et al., 2015 Bacillus spp. and Enterobacter spp. Fusarium solani f. sp. piperis and F. oxysporum f. sp. piperis (ascomycetes) causing foot rot and stem blight in black peppers Production of volatile and diffusible bioactive antifungal compounds Edward et al., 2013 Bacillus spp., Burkholderia spp., and Pseudomonas spp. Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms As an endophyte Ramli et al., 2016; Seung et al., 2015 Burkholderia cepacia Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Production of proteolytic enzymes Azadeh et al., 2010 Burkholderia cepacia and Pseudomonas aeruginosa Colletotrichum gloeosporioides (ascomycete) causing anthracnose in papayas Production of volatile and nonvolatile inhibitory compounds Rahman et al., 2007 Burkholderia sp. Phytophthora capsici (oomycete) causing foot rot and stem blight in black peppers Production of hydrolytic enzymes and antimicrobial compounds Ahmad & Ahmadu, 2017 Burkholderia sp. and Pseudomonas sp. Fusarium oxysporum f. sp. cubense (ascomycete) causing Fusarium wilt in bananas As an endophytes which induces host defense enzymes Mohd Fishal et al., 2010 Enterobacter asburiae, Enterobacter cancerogenus, and Enterobacter cloacae Phytophthora capsici (oomycete) causing foot rot and stem blight in black peppers Production of volatile inhibitory compounds Toh et al., 2016 Biological control agent Reference (b) Basidiomycete (c) Yeast 110 species were screened, of which 29 species showed antagonism (d) Bacterium (Continues) BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… 650 TABLE 1 (Continued) Application/potential mechanism of action Reference Colletotrichum capsici (ascomycete) causing anthracnose in chili Seed treatment of chili El-Mabrok et al., 2012 Pseudomonas aeruginosa Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms As an endophyte Sathyapriya et al., 2012; Bivi et al., 2010 Pseudomonas fluorescens Fusarium oxysporum f. sp. cubense (ascomycete) causing Fusarium wilt in bananas Production of inhibitory compounds Mohammed et al., 2011 Pseudomonas spp. Rhizoctonia solani (basidiomycete) causing sheath blight of rice Soil treatment (spraying) with bacterial suspension Akter et al., 2016, 2014 Serratia marcescens Fusarium oxysporum f. sp. cubense (ascomycete) causing Fusarium wilt in bananas Soil treatment with inoculum which induces host defense enzymes Ting et al., 2010 Streptomyces glauciniger Phytopathogenic fungi Production of chitinase Awad et al., 2014 Streptomyces griseus Fusarium oxysporum f. sp. cubense (ascomycete) causing Fusarium wilt in bananas Soil treatment (spraying) with bacterial suspension Zacky & Ting, 2015, 2013 Streptomyces violaceusniger Fusarium oxysporum f. sp. cubense (ascomycete) causing Fusarium wilt in bananas Production of inhibitory compounds Getha & Vikineswary, 2002 Streptomyces spp. Colletotrichum capsici, Colletotrichum acutatum, and Colletotrichum gloeosporioides (ascomycete) causing anthracnose in chili Soil treatment (spraying) with bacterial suspension Shahbazi et al., 2014, 2013 Streptomyces spp. Pyricularia oryzae (rice blast fungus) infesting rice Production of inhibitory compounds Awla et al., 2017; Law et al., 2017 Acerophagus papayae (wasp) Paracoccus marginatus (papaya mealybug) infesting papayas As an endoparasitoid Mastoi et al., 2018 Apanteles metesae (wasp) Metisa plana (bagworm moth) whose larvae infest oil palms As a parasitoid Salmah et al., 2012 Calycomyza lantanae and Ophiomyia lantanae (flies) Lantana camara (big sage), invasive weed Larvae of flies are leaf-eaters Ooi, 1987 Cotesia vestalis (wasp) Plutella xylostella (diamondback moth) whose larvae infest cabbages As a parasitoid Kermani et al., 2014 Cosmolestes picticeps and Sycanus dichotomus (bugs) Metisa plana (bagworm moth) whose larvae infest oil palms As predatory natural enemies Jamian et al., 2017 Diachasmimorpha longicaudata, Fopius arisanus, Fopius vandenboschi, Psyttalia fletcheri, and Psyttalia incisi (wasps) Bactrocera carambolae, Bactrocera papayae, and Bactrocera cucurbitae (fruit flies) infesting fruits As a parasitoid Yaakop et al., 2015; Shariff et al., 2014; Ibrahim et al., 2013; Chinajariyawong et al., 2000 Dolichoderus thoracicus (ant) Conopomorpha cramerella (cocoa pod borer moth) whose larvae infest cocoa Active and dispersive predatory behavior of ants Adnan et al., 2018 Dolichoderus thoracicus and Oecophylla smaragdina (ants) Helopeltis theobromae (mosquito bug) causing leaf tattering and fruit blemishes in cocoa Active and dispersive predatory behavior of ants Way & Khoo, 1991 Menochilus sexmaculatus (ladybird beetle) Rhopalosiphum maidis (aphid) infesting maize Predatory behavior of beetles Ibrahim & Kueh, 2013 Micromus tasmaniae (lacewings) Aphids infesting potatoes Plant treatment (spraying) with egg suspension Hussein, 1984 Biological control agent Targeted pest/pathogen/disease Lactic acid bacteria (e) Insect (Continues) BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… TABLE 1 651 (Continued) Biological control agent Targeted pest/pathogen/disease Application/potential mechanism of action Oecophylla smaragdina (weaver ant) Hypsipyla robusta (moth) whose larvae feed on timber shoots, flowers, and barks Active and dispersive predatory behavior of ants Lim et al., 2008 Oecophylla smaragdina (weaver ant) Pteroma pendula (bagworm moth) whose larvae infest oil palms Active and dispersive predatory behavior of ants Pierre & Idris, 2013 Tamarixia radiata (wasp) Diaphorina citri (Asian citrus psyllid) infesting citrus fruits As a parasitoid Sule et al., 2014 Plutella xylostella (diamondback moth), Spodoptera exigua (beet armyworm moth), and Spodoptera litura (tobacco cutworm moth or cotton leafworm moth) whose larvae infest fruits and vegetables Spraying of spore suspension targeting larvae Ghani et al., 2013; Kermani et al., 2013; Ramli et al., 2011 Echinochloa crus-galli (barnyard grass), weed infesting rice paddy fields Production of volatile inhibitory compounds Aslani et al., 2015 Limnocharis flava (yellow velvetleaf), weed infesting rice paddy fields As herbivore Gilal et al., 2016 Baculovirus Spodoptera litura (tobacco cutworm moth or cotton leafworm moth) whose larvae infest star fruits and vegetables Spraying of viral occlusion bodies Zabedah et al., 2010 Baculovirus Plutella xylostella (diamondback moth) whose larvae infest cabbages Spraying of viral occlusion bodies Abdul Kadir et al., 1999 Cucumber Mosaic Virus Bemisia tabaci (silverleaf whitefly) infesting chili Induction of host resistance through artificial viral inoculation onto the plant Saad et al., 2019 Nudivirus Oryctes rhinoceros (rhinoceros beetle) infesting oil palm fronds Spraying of viral occlusion bodies Moslim, Kamarudin, Ghani, et al., 2011; Ramle et al., 2005 A consortium of functional microbiota in microbial-enriched compost tea Golovinomyces cichoracearum (ascomycete) causing powdery mildew on melons Spraying of melon seedlings with the compost tea Naidu et al., 2012 Bacillus thuringiensis (bacteria) and Metarhizium anisopliae (ascomycete) Tirathaba rufivena (bunch moth) whose larvae infest oil palms Spraying with fungal and bacterial suspension Mohamad et al., 2017 Bacillus spp. (bacteria) and Trichoderma spp. (ascomycetes) Ganoderma boninense (basidiomycete) causing basal stem rot in oil palms Production of volatile and nonvolatile inhibitory compounds Angel et al., 2018; Alexander et al., 2017 Bacillus subtilis (bacterium) and Trichoderma spp. (ascomycetes) Rhizoctonia solani (basidiomycete) causing various commercially significant plant diseases Soil treatment (spraying) with fungal and bacterial suspension Ali & Nadarajah, 2013 Cataenococcus hispidus (mealybug) and Dolichoderus thoracicus (ant) Helopeltis theobromae (mosquito bug) causing leaf tattering and fruit blemishes in cocoa Active and dispersive predatory behavior of ant and bug Ho & Khoo, 1997 Reference (f) Microsporidia Nosema bombycis (g) Shrub Tinospora tuberculate (h) Snail Pomacea canaliculata and Pomacea maculata (i) Virus (j) Combination (Continues) BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… 652 TABLE 1 (Continued) Biological control agent Targeted pest/pathogen/disease Application/potential mechanism of action Eurytoma attiva (wasp) and Metrogaleruca obscura (beetle) Cordia curassavica (tropical black sage or wild sage), invasive weed Defoliation by beetle and seed-destruction by wasp Simmonds, 1980 Pseudomonas aeruginosa (bacterium) Trichoderma harzianum and Trichoderma virens (ascomycetes) Colletotrichum truncatum (ascomycete) causing damping-off in soybeans Seed treatment with conidial suspension Begum et al., 2010 Reference Number of available publications (n) Note. Publications were obtained based on a literature search on SCOPUS® . Search keywords: biological control, Malaysia. Biological control agents are sorted according to types of organisms. FIGURE 3 Distribution of targeted crop commodities on which scientific investigations on potential biological control agents against phytopathogenic organisms have been conducted in Malaysia (1980–2019) Note. Publications were obtained based on a literature search on SCOPUS® , as similarly listed in Table 1. Search keywords: biological control, Malaysia 40 32 30 28 20 11 10 10 8 3 0 Cocoa Deciduous Fruits & trees vegetables Herbs & spices Oil palm Rice paddy Targeted crop commodities Ganoderma Management (IGM), which incorporates several elements including sanitation, biological control, chemical control, and the use of fertilizers with beneficial trace elements. The integration of biological control in the system is based on the understanding that low incidence of disease has been achieved in the presence of natural competitors/predators that keep the parasitic fungi under control (Bivi, Farhana, & Khairulmazmi, 2010). In an effort to further strengthen this approach, MPOB has also developed and commercialized their biological control formulations, namely, GanoEF biofertilizer (contains endophytic Hendersonia GanoEF1 fungus; mechanism of action: production of inhibitory bioactive compounds; Nurrashyeda et al., 2015) and EmbioTM actinoPLUS (contains Streptomyces sp.; mechanism of action: production of inhibitory bioactive compounds; Shariffah-Muzaimah et al., 2015). For rice paddy, the rice blast disease caused by Pyricularia oryzae is considered a major endemic in rice paddy planting areas in Malaysia and across the world. Currently, the cultivation of resistant varieties and the application of fungicides are the most commonly used methods in controlling the disease in rice paddy planting areas in Malaysia (Abed-Ashtiani et al., 2018). However, due to the fact that both practices fail to provide a long-term solutions for the disease control, growing amounts of research have been conducted to incorporate biological control approach as an alternative. Recently, an endophytic actinomycete, Streptomyces sp. strain UPMRS4, isolated from the rhizosphere of irrigated rice paddy fields in Malaysia, has been shown to successfully suppress the rice blast disease through the production of inhibitory bioactive compounds, thus inducing the systemic resistance of rice paddy against P. oryzae (Awla et al., 2017). Figure 4 depicts the types of potential biological control agents against phytopathogenic organisms in Malaysia, which are dominated by bacteria, ascomycetes, and insects. The use of bacteria as biological control agents, especially Bacillus spp., Burkholderia spp., and Pseudomonas spp., has shown considerable success in Malaysia (Table 1). Ubiquitous in nature, these bacteria exist as both nonparasitic and parasitic pathogenic species; the latter as potent producers of various enzymes and antimicrobial substances that inhibit important phytopathogens while at the same time promote plant growth by the production of volatile substances (Beneduzi, Ambrosini, & Passaglia, 2012). BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Number of available publications on potential biological control agents against phytopathogenic organisms from Malaysia (1980–2019) Note. Publications were obtained based on a literature search on SCOPUS® , as similarly listed in Table 1. Search keywords: biological control, Malaysia Number of available publications (n) FIGURE 4 653 40 35 30 29 18 20 10 5 5 1 1 1 1 0 Types of biological control agents Furthermore, considering that the majority of crop pests are insects, the use of entomopathogenic fungi, especially from the Division Ascomycota, has proven efficient on many crop species (Strasser, Vey, & Butt, 2010). Species such as Beauveria bassiana, Isaria fumosorosea, Metarhizium anisopliae, Paecilomyces spp., and Trichoderma spp. have been efficacious against beetles, moths, and termites infesting a multitude of crop commodities in Malaysia (Table 1). The tropical climate of Malaysia enhances the propagation and proliferation of these entomopathogenic fungi. In lethally parasitizing the target insect pests, or severely disabling them, entomopathogenic ascomycetes usually initiate the infection in the form of airborne conidia that naturally or artificially (that is, through human intervention) land onto the insects’ exoskeleton. Under suitable ecophysiological conditions, these spores germinate and grow into hyphae, thereby colonizing the insect’s cuticle. These hyphae are capable of producing hydrolyzing enzymes, which bore a tunnel through which the hyphae could penetrate into the insects’ body cavity, and there proliferate further, thus killing the host (Fernandes, Valério, Feltrin, & Van Der Sand, 2012). Entomopathogenic fungi can grow over a wide range of temperatures, and some strains have the appropriate ecophysiological growth parameters for use in climates such as Malaysia (Borisade & Magan, 2015). Being environmentally safe, the use of entomopathogenic ascomycetes draws worldwide interest for biological control of insect pests including in Malaysia. Apart from insects, these entomopathogenic fungi have also been shown to parasitize other phytopathogenic ascomycetes as well as nematodes (Table 1). For insect species that are used to control insect pests, the biological control agents come either as insectivorous predators or insect parasitoids. Insectivorous predators are mainly free-living species that aggressively prey on a large number of insects during their lifetime, thus controlling the pest density (Eggleton & Belshaw, 1992). Given that many major crop pests are insects, majority of the biological control agents used are insectivorous species (Table 1). The ladybird beetles (Menochilus sexmaculatus) are voracious predators of the aphids (Rhopalosiphum maidis) infesting maize. Various predatory ant species such as Dolichoderus thoracicus have been successful in controlling the cocoa pod borer moths (Conopomorpha cramerella) whose larvae infest cocoa; mosquito bugs (Helopeltis theobromae) that cause leaf tattering and fruit blemishes in cocoa; Oecophylla smaragdina that prey on bagworm moths (Pteroma pendula) whose larvae infest oil palms; and Hypsipyla robusta whose larvae feed on timber shoots, flowers, and barks. For insect parasitoids, their mechanism of action is generally by laying their eggs (oviposition) on (or in) the body of an insect host, on (or in) which their developing larvae will feed, consequently decapitating the host (Kapranas, Tena, & Luck, 2012). Parasitoids are almost always endopterygote insects that have complete metamorphosis stages during their lifecycle, such as parasitoid larvae and free-living adults. Parasitoids are generally classified into endoparasitoids, which live within their host’s body, or ectoparasitoids, which feed on their host’s body from outside (Colmenarez, Corniani, Jahnke, Sampaio, & Vásquez, 2018). The majority of insect parasitoids that have been used in Malaysia are from the wasp family such as Acerophagus papayae, Apanteles metesae, Cotesia vestalis, Diachasmimorpha longicaudata, Fopius arisanus, F. vandenboschi, Psyttalia incisi, P. fletcheri, and Tamarixia radiata, which have shown considerable success in reducing the density of moth caterpillars, bugs, fruit flies, and psyllids infesting various economically important fruits and vegetables such as cabbages, carambolas (star fruit), melons, oranges, and papayas (Table 1). Wasps have been shown to exhibit both endo– and ectoparasitoid behaviors. 654 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Although the majority of insect parasitoids are wasps (Hymenoptera) and flies (Diptera) (Eggleton & Belshaw, 1992), to the best of our knowledge thus far, there has been no available record on the use of the latter as a biological control agent in Malaysia. This could either be due to the fact that this particular application is totally new in Malaysia, therefore suitable for future investigation, or that ongoing trials are not yet conclusive and published. 3 BIOLOGICA L CONTROL AGA I N ST M YCOTOX I G E N I C FU NG I Mycotoxins are low-molecular-weight and heat stable secondary metabolites produced by microfungi, and capable of causing diseases and death in vertebrates (humans and animals; Bennett & Klich, 2003). Members of the genera Aspergillus, Fusarium, and Penicillium are often regarded as the major producers of mycotoxins with health and economic impacts. Among the major challenges in controlling mycotoxigenic fungi and the subsequent mycotoxin contamination in crop commodities is that they are not very sensitive/susceptible to fungicides commonly used in the field to control fungal pathogens. At present, almost 500 different mycotoxins and fungal metabolites have been identified as being produced by more than 300 microfungal species (Nielsen & Smedsgaard, 2003). Although mycotoxins are not necessary for growth of their fungal producers, but because mycotoxins can weaken their competitors (usually other microorganisms competing for the same substrate on crop commodities; competitive exclusion), the fungal producers are thought to use mycotoxins as a strategy to ensure their survival and proliferation (Hussein & Brasel, 2001). In terms of their carcinogenicity toward humans and animals, the IARC, which is under the auspices of the World Health Organization (WHO) of the United Nations (UN) and serves as the authoritative source of information on causes of cancers (IARC: https://www.iarc.fr/), has classified mycotoxins into several groups (Ostry et al., 2017) as in Table 2. Although the biological control against phytopathogenic organisms has a long history in Malaysia, as mentioned earlier, the same is not entirely true for mycotoxigenic fungi. Surprisingly, the application of biological control against mycotoxigenic fungi is almost nonexistent in Malaysia with only a handful of publications available online on the subject matter (Samsudin & Magan, 2016; Samsudin, Medina, & Magan, 2016; Samsudin, Rodriguez, Medina, & Magan, 2017). These studies examined the use of Clonostachys rosea and Streptomyces sp. to control the growth of Fusarium verticillioides and the contamination of maize with fumonisins. The only other study of similar nature was published slightly earlier on the use of the mycoparasite Trichoderma harzianum against TABLE 2 The International Agency for Research on Cancer (IARC) Monographs evaluations of the carcinogenic hazard of selected mycotoxins Mycotoxin Group Aflatoxins B1 , B2 , G1 , G2 , and M1 1 Citrinin 3 Cyclochlorotine 3 Deoxynivalenol 3 Fumonisin B1 2B Fumonisin B2 2B Fusarenone X 3 Fusarin C 2B Kojic acid 3 Luteoskyrin 3 Nivalenol 3 Ochratoxin A 2B Patulin 3 Penicillic acid 3 Rugulosin 3 Sterigmatocystin 2B T-2 toxin 3 Zearalenone 3 Note. Adapted from Ostry et al. (2017). Group 1, carcinogenic to humans (sufficient evidence); Group 2A, probably carcinogenic to humans (limited evidence of carcinogenicity in humans but sufficient evidence in experimental animals); Group 2B, possibly carcinogenic to humans (limited evidence of carcinogenicity in humans and less than sufficient evidence in experimental animals); Group 3, not classifiable as to its carcinogenicity to humans (evidence is inadequate in humans and inadequate or limited in experimental animals); Group 4, probably not carcinogenic to humans (evidence suggesting lack of carcinogenicity in humans and in experimental animals). At present, no known mycotoxin has been classified in groups 2A and 4. The complete listing of the LD50 of each mycotoxin can be obtained from the Toxicology Data Network, National Library of Medicine, National Institutes of Health, USA (https://toxnet.nlm.nih.gov/). LD50 = median lethal dose/semi-lethal dose/sublethal dose, which is the dose required to kill half the members of a tested population after a specified test duration, and is frequently used as a general indicator of a substance’s acute toxicity. F. proliferatum and F. verticillioides in controlling Fusarium ear rot in maize (Suhaida & NurAinIzzati, 2013). However, they did not specifically examine or quantify the impact on control of any mycotoxins. Because these fungal pathogens, under Malaysian climatic conditions, would certainly contaminate maize with fumonisins during silking, this would have been useful additional information. It is possible that this has not been recognized as an important research area in Malaysia so far. However, as Malaysia does have legislations for mycotoxin contamination of some commodities (Table 3; Malaysian Food Regulation, 1985), this could be a major driver for practically related research for the reduction of mycotoxins using biological control strategies in the very near future. Table 4 lists the occurrence data of mycotoxins on Malaysian foods and feedstuffs, from which it is apparent that the concern is more on aflatoxins. Malaysia’s tropical climate, BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… TABLE 3 655 Maximum permitted proportion (𝜇g/kg or ppb) of mycotoxins in foods Maximum permitted proportion (𝝁g/kg or ppb) Food Total B1 M1 Aflatoxins Groundnuts, almonds, hazelnuts, pistachios, and Brazil nuts (shelled, for further processing) 15 Groundnuts, almonds, hazelnuts, pistachios, and Brazil nuts (shelled, ready-to-eat) 10 Cereal-based food for infants and children 0.1 Milk 0.5 Infant formula and follow-up formula (ready-to-drink) Others 0.025 5 Ochratoxin A Cereal-based food for infants and children 0.5 Coffee or ground coffee or coffee powder 5 Instant coffee or soluble coffee and decaffeinated coffee 10 Patulin Apple juice (includes apple juices as ingredients in other beverages) 50 Note. Adapted from Malaysian Food Regulations (1985). which is favorable for Aspergillus spp. proliferation and the subsequent mycotoxin production, poses a constant threat to its agriculture sector in producing safe and quality harvest. A more comprehensive review on mycotoxin contamination of Malaysian food and agricultural produce has been published elsewhere (Afsah-Hejri, Jinap, Hajeb, Radu, & Shakibazadeh, 2013). Given the fact that peanuts and maize are the two most contaminated crop commodities by members of Aspergillus section Flavi (Pitt & Hocking, 2009; Samson, Houbraken, Thrane, Frisvad, & Andersen, 2010), which almost invariably produce the Group 1 carcinogenic aflatoxins, and that these two are relevant in Malaysian agricultural sector (listed as cash crops; Figure 2), further investigation on the development of an efficient control against aflatoxigenic fungal colonization on these crops is highly warranted. Similar threat on peanuts and maize is also observed in the Philippines (Balendres, Karlovsky, & Cumagun, 2019), Ghana (Agbetiameh et al., 2018), and Zambia (Kachapulula, Akello, Bandyopadhyay, & Cotty, 2017). As described in Table 1, several biological control agents have been investigated on their effects against phytopathogenic Fusarium spp. More detailed examination shows that some of these Fusarium spp. are in fact also mycotoxigenic (for example, F. proliferatum: beauvericins, fumonisins, and moniliformin; F. solani: moniliformin and trichothecenes; F. verticillioides: fumonisins, fusaric acid, and moniliformin) according to Desjardins (2006). Therefore, opportunities do exist for replicating such studies with biological control agents and investigate their effects in controlling mycotoxin contamination. In addition, as most studies have focused on the use of effective biological control agents against plant diseases, it may be ben- eficial to also screen these for mycotoxin control (Medina, Mohale, et al., 2017). Nevertheless, previous studies have also suggested that several potential bacterial antagonists may, under certain abiotic conditions, stimulate mycotoxin production instead of inhibiting it (Al-Saad, Al-Badran, Al-Jumayli, Magan, & Rodríguez, 2016). Thus, care is needed when screening for mycotoxin control. There are also some cases in which the fungal pathogens could “disarm” the applied antagonist. For example, it was shown that deoxynivalenol-producing F. culmorum and F. graminearum strains could repress the expression of genes that regulate the chitinase enzyme production in the biological control agent Trichoderma atroviride (Lutz, Feichtinger, Défago, & Duffy, 2003). This is not unusual because pathogens also possess diverse responses to counteract antagonists, which include detoxification and active efflux of antibiotics, and repression of biosynthetic genes involved in biological control (Duffy, Schouten, & Raaijmakers, 2003). In other parts of the world, however, the application of biological control agents to combat mycotoxigenic fungi is more widely practiced. Among the earliest and excellent reviews that discussed the potential use of nontoxigenic strains of fungi, yeasts, and bacteria in controlling mycotoxigenic fungi was published by Cleveland, Dowd, Desjardins, Bhatnagar, and Cotty (2003). The review also discussed the indirect controls of mycotoxigenic fungi by controlling their vectoring insects or developing insect-resistance plant cultivars (e.g., Bt maize) because insect damage provides entry points for mycotoxigenic fungi to colonize the crop, thus resulting in increased mycotoxin contamination. Recently, a review on the dynamics and mechanisms of action of biological control agents against mycotoxigenic fungi was critically discussed BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… 656 TABLE 4 Occurrence of mycotoxins on Malaysian food and feed Food/Feed (number of sample, n) Mycotoxin n positive, n above limit Reference Spices and spice-based products (n = 90) Aflatoxins and ochratoxin A AFs n positive: 19/90 n above limit: 19/90 OTA n positive: 37/90 n above limit: 25/90 Ahmad-Zaidi et al., 2020 Commercial food and beverage (n = 120); apple, grape, orange, and pomegranate juices; wheat and barley flour; dried figs, raisins, chili powder, and spices; nonroasted peanut; roasted pistachio Aflatoxins and ochratoxin A AFs n positive: 19/120 n above limit: 14/120 OTA n positive: 3/120 n above limit: None Alsharif et al., 2019 Raw peanuts (n = 87) and peanut-based products (n = 91) Aflatoxins Raw peanuts n positive: 36/87 n above limit: 20/87 Peanut-based products n positive: 50/91 n above limit: 9/91 Norlia et al., 2018 Milk and dairy products (n = 53) Aflatoxin M1 n positive: 19/53 n above limit: 4/53 Nadira et al., 2017 Cow’s, goat’s, and human’s milk (n = 33) Aflatoxin M1 n positive: 2/33 n above limit: None Shuib et al., 2017a Cow’s milk (n = 102) Aflatoxin M1 n positive: 4/102 n above limit: 3/102 Shuib et al., 2017b Commercial vegetable oil (n = 25) Aflatoxins, ochratoxin A, deoxynivalenol, and zearalenone AFs n positive: None n above limit: None OTA n positive: None n above limit: None DON n positive: None n above limit: None ZEN n positive: 15/25 n above limit: None Sharmili et al., 2016 Commercial spice (n = 58) Aflatoxins and ochratoxin A AFs n positive: 50/58 n above limit: 15/58 OTA n positive: 27/34 n above limit: 1/34 Ali et al., 2015 Red yeast rice (traditional Chinese medicine; n = 50) Aflatoxins, ochratoxin A, and citrinin AFs n positive: 46/50 n above limit: 35/50 OTA n positive: 50/50 n above limit: None CIT n positive: 50/50 n above limit: 50/50 Samsudin and Abdullah, 2013 Commercial dried chili (n = 80) Aflatoxins and ochratoxin A AFs n positive: 52/80 n above limit: 9/80 OTA n positive: 65/80 n above limit: 13/80 Jalili and Jinap, 2012 (Continues) BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… TABLE 4 657 (Continued) Food/Feed (number of sample, n) Mycotoxin n positive, n above limit Reference Commercial rice (n = 5), peanut (n = 9), and chili (n = 10) Aflatoxins Rice n positive: 4/5 n above limit: 3/5 Peanut n positive: 4/9 n above limit: 2/9 Chili n positive: 9/10 n above limit: 3/10 Khayoon et al., 2012 Commercial coffee and cereal (n = 45) Ochratoxin A Commercial cereal (n = 100) Aflatoxins B1 , B2 , G1 , and G2 , ochratoxin A, zearalenone, deoxynivalenol, and fumonisins B1 , B2 , T-2, and HT-2 AFs n positive: 60/100 n above limit: 2/100 OTA n positive: 40/100 n above limit: 2/100 ZEN n positive: 25/100 n above limit: None DON n positive: 36/100 n above limit: None FUM n positive: 19/100 n above limit: None T-2 n positive: 16/100 n above limit: None HT-2 n positive: 16/100 n above limit: None Soleimany, Jinap, & Abas, 2012 Commercial cereal (n = 80) Aflatoxins B1 , B2 , G1 , and G2 , ochratoxin A, zearalenone, deoxynivalenol, and fumonisins B1 , B2 , T-2, and HT-2 AFs n positive: 40/80 n above limit: 2/80 OTA n positive: 24/80 n above limit: 1/80 ZEN n positive: 15/80 n above limit: None DON n positive: 24/80 n above limit: None FUM n positive: 13/80 n above limit: None T-2 n positive: 11/80 n above limit: None HT-2 n positive: 9/80 n above limit: None Soleimany, Jinap, Faridah, & Khatib, 2012 Commercial food products (n = 95) Aflatoxin B1 Grain corn for animal feed (n = 80) Aflatoxin B1 and fumonisins n positive: 25/45 n above limit: 1/45 n positive: 69/95 n above limit: None AFB1 n positive: 65/80 n above limit: 18/80 FUM n positive: 80/80 n above limit: None Lee et al., 2012 Reddy et al., 2011 Reddy & Salleh, 2011 (Continues) BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… 658 TABLE 4 (Continued) Food/Feed (number of sample, n) Mycotoxin n positive, n above limit Commercial food and feed (n = 39) Fumonisins n positive: 5/39 n above limit: None Reference Khayoon, Saad, Salleh, et al., 2010 Animal feed (n = 42) Aflatoxins n positive: 8/42 n above limit: 3/42 Khayoon, Saad, Yan, et al., 2010 Commercial wheat (n = 15) and barley (n = 15) Aflatoxin B1 Wheat n positive: 3/15 n above limit: None Barley n positive: 1/15 n above limit: None Reddy & Salleh, 2010 Commercial cereal (n = 60) Aflatoxins, ochratoxin A, and zearalenone AFs n positive: 24/60 n above limit: 1/42 OTA n positive: 9/60 n above limit: 2/42 ZEN n positive: 11/60 n above limit: None Rahmani et al., 2010 Commercial noodle (n = 135) Deoxynivalenol n positive: 110/135 n above limit: None Moazami & Jinap, 2009 Raw shelled peanut (n = 145) Aflatoxins n positive: 73/145 n above limit: 33/145 Sulaiman et al., 2007 Abbreviations: AFs, aflatoxins; CIT, citrinin; DON, deoxynivalenol; FUM, fumonisin; HT-2, HT-2 toxin; OTA, ochratoxin A; T-2, T-2 toxin; ZEN, zearalenone. by Medina, Mohale, et al. (2017). They suggested that the mechanism of action and the inoculum dose required for effective control of mycotoxigenic fungi may be different from that for phytopathogenic fungi. Therefore, economical production/formulation is critical for successful biological control of mycotoxins. Nevertheless, they concur that the primary mechanisms of action for the control of mycotoxigenic fungi or phytopathogenic organisms usually include niche/habitat exclusion, competition for nutrients, and production of inhibitory volatiles. In fact, Köhl, Kolnarr, and Ravensberg (2019) have suggested that biological control agents utilize a cascade of different interacting approaches for efficacy. Based on the available literature, we could summarize the biological control agents into three major groups: (a) nontoxigenic strains of known mycotoxin producers, (b) mycoparasitic microfungi, and (c) phyllosphere or soilborne bacteria. The use of nontoxigenic Aspergillus flavus and A. parasiticus strains to control their toxigenic counterparts began in the early 1990s in the United States (Brown, Cotty, & Cleveland, 1991). Since then, this application has found success in other parts of the world where aflatoxin contamination is significant and has had significant impacts on the economies, such as Kenya (Marchera & Ndwiga, 2015), Italy (Mauro, Garcia-Cela, Pietri, Cotty, & Battilani, 2018), Argentina (Alaniz Zanon, Clemente, & Chulze, 2018), and Thailand (Tran-Dinh, Pitt, & Markwell, 2018). Its modes of action are either through competitive exclusion (that is, toxigenic strain displacement) (Damann, 2015) or through thigmoregulation (that is, touch inhibition) (Huang, Jha, Sweany, DeRobertis, & Damann, 2011). However, in securing an efficient biological control agent, it should be noted that lack of mycotoxin biosynthesis because of the deletion of key biosynthetic gene clusters may not be the only criterion, rather the nontoxigenic candidate must also be an effective colonizer of the crop commodity onto which it would be applied (to competitively exclude its competitors), and that it must also be able to reduce the mycotoxin contamination (Horn & Dorner, 2011). Recent investigation proposed yet another mechanism for the antagonism of the nontoxigenic strains, which is via active extrolites (that is, uncharacterized compounds) secretion (Moore, Lebar, & Carter-Wientjes, 2019). Another successful area for research and applications is the use of mycoparasitic microfungi. As described in Table 1, Trichoderma spp. feature predominantly as a competent biological control agent against a wide variety of phytopathogenic organisms. Similar traits are also exhibited against mycotoxigenic fungi and subsequent mycotoxin production (Braun et al., 2018; Saravanakumar et al., 2018; Veenstra, Rafudeen, & Murray, 2019). Prevalently found in the rhizo- and phyllosphere, the free-living and fast-growing Trichoderma spp. often exist in mutualistic endophytic relationships with plants. As a biological control agent, Trichoderma spp. demonstrate antagonism through antibiotic production, parasitism (for example, penetration and killing), inducing host-plant resistance (that is, Trichoderma spp. are also known to have plant growth promoting characteristics), and habitat/nutrient competition (that is, Trichoderma spp. are effective decomposers and aggressive colonizers, BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… thus leaving scarce macro- and micronutrients from the soil for other competitors). Trichoderma spp. also produce a wide array of enzymes, chief among which is the fungal-cell-walldegrading chitinase (Harman, 2006). The ability of Trichoderma spp. to produce chemically diverse secondary metabolites that could modify the environmental conditions into that which is inhospitable for their competitors has also been extensively reviewed by Keswani, Mishra, Sarma, Singh, and Singh (2014) and Reino, Guerrero, Hernández-Galán, and Collado (2008). Finally is the use of soilborne bacteria that produce various antibiotics and fungal-cell-wall-degrading enzymes. As in the control of phytopathogenic organisms, Bacillus spp. also exhibit potency and prowess in the control of mycotoxigenic fungi. This is mainly attributed to the production of cyclic lipopeptides that display amphiphilic properties that could disrupt the fungal cell wall and cell membrane (Siahmoshteh et al., 2017). Other chemical metabolites produced by Bacillus spp. could also inhibit fungal nutrient acquisition, mycelial growth, and conidial germination (Cao et al., 2012; Li et al., 2009; Volpon, Besson, & Lancelin, 2000). Recent investigation also elucidates the ability of enzymes produced by Bacillus spp. in degrading mycotoxins (Afsharmanesh, PerezGarcia, Zeriouh, Ahmadzadeh, & Romero, 2018). Besides Bacillus spp., lactic acid bacteria have also been shown to possess antimycotoxigenic properties, including the production of organic acids, which will acidify environmental pH, thus inhibiting fungal growth and mycotoxin production; antifungal compounds (for example, propionate and cyclic dipeptide); and competitive exclusion of fungal pathogens (Guimarães, Santiago, Teixeira, Venâncio, & Abrunhosa, 2018; Oliveira, Zannini, & Arendt, 2014). The efficiency of Streptomyces spp. in combatting mycotoxigenic fungi is also of note. The production of various bioactive molecules such as antibiotics (e.g., candicidin, phosphlactomycin, tubercidin, and pyrrole-2-carboxylic acid), lytic enzymes, and volatile organic compounds (e.g., phenylethyl alcohol and (+)-epi-bicyclesesquiphellandrene) has greatly contributed to the antagonism (Hoster, Schmitz, & Daniel, 2005; Mari, Bautista-Baños, & Sivakumar, 2016; Nguyen et al., 2018). The growth inhibition of mycotoxigenic fungi and the subsequent mycotoxin production by Streptomyces spp. in wheat (Nourozian, Etebarian, & Khodakaramian, 2006; Palazzini, Ramirez, Torres, & Chulze, 2007; Palazzini, Yerkovich, Alberione, Chiotta, & Chulze, 2017) and in peanuts (Sultan & Magan, 2011) have been demonstrated. 4 EMERGING CHALLENGES I N M YCOTOX I N M A NAGE M E N T At present, the world is threatened by climate change and the potential dangers it brings. Climate change is an alteration 659 of the average weather distributions over prolonged period of time (that is, decades to centuries), which is caused by several factors such as biotic processes, solar radiation, and volcanic eruptions. The worldwide increase of atmospheric carbon dioxide (CO2 ; second-most abundant greenhouse gas) and consequently the rise of temperature (that is, global warming) constitute the major manifestations of climate change. The average temperature of the lower troposphere (that is, lowest layer of Earth’s atmosphere) has increased between 0.15 and 0.25 ◦ C per decade since 1980 according to satellite temperature measurements, whereas the atmospheric concentration of CO2 has increased from 280 ppm in 1750 to 400 ppm in 2015 (ESRL, 2019). What is more alarming is that the increment trends of both parameters have been accelerating decade by decade, and are actually made worse by human activities. For instance, CO2 has approximately increased from 280 ppm in 1750 (beginning of Industrial Revolution) to 330 ppm in 1970, which saw an increase of 50 ppm in around 220 years. However, the next 50 ppm steadily rose within just 30 years to 380 ppm (ca. 1970 to 2000), and the next 20 ppm in 15 years (ca. 2000 to 2015) to the present concentration of 411 ppm (ESRL, 2019). Worldwide, as a result of climate change, humans will face substantial threats in terms of health, food security due to decreasing crop yields, and the desertification of populated areas due to rising sea levels (Battisti & Naylor, 2009). Furthermore, there has also been an increasing concern that climate change permits for the increased propagation of crop pests and pathogens and their movement to other previously unsuitable regions (Bebber, Ramotowski, & Gurr, 2013) due to the alteration in the natural environments, which creates new opportunities for evolution (Fisher et al., 2012). The impacts of climate change on the propagation of microfungi, especially that of the mycotoxigenic fungi and the subsequent mycotoxin production, have been previously discussed in depth (Botana & Sainz, 2015; Ksenija, 2018; Medina, Akbar, Baazeem, Rodriguez, & Magan, 2017, 2015; Paterson & Lima, 2011, 2010). As if the threats of climate change are not severe enough, we are also burdened by the recently discovered “masked mycotoxins.” Masked mycotoxins are biologically modified mycotoxins that occur as a result of plant defense mechanism against xenobiotic compounds that alters the chemical structure of the mycotoxins through conjugation with sugars, amino acids, or sulfate groups, and compartmentalize them in plant cell vacuoles or conjugated to biopolymers such as cell wall components. These conjugated mycotoxins are nontoxic. However, they could be converted back into their parent form by hydrolysis during food and feed processing or in the digestive tract of humans and animals (Berthiller, Schuhmacher, Adam, & Krska, 2009a; 2016; Galaverna, Dallsta, Mangia, Dossena, & Marchelli, 2009). The issue of masked mycotoxins began attracting scientific interest after several mysterious cases of mycotoxicoses during the mid-1980s, 660 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… in which symptoms in severely affected animals did not correlate with the low mycotoxin content detected in their feed (Berthiller et al., 2009b; 2016). Furthermore, masked mycotoxins generally elude the established quantification analyses (for example, HPLC), thus leading to an underestimation of their exposure (hence the term “masked”), and masked mycotoxin standards, reference materials, and interlaboratory-validated quantification methods are not yet commercially/widely available. The terms “masked,” “hidden,” “conjugated” (covalently linked), and “bound” (noncovalently linked) are frequently used in the literature, with “masked” being the most popular (Berthiller et al., 2009a). More recently, a new term, “modified mycotoxin,” has been proposed that encompasses other possible forms of mycotoxins and their modifications including masked mycotoxins in the effort “to harmonize future scientific wording and subsequent legislation” (Rychlik et al., 2014). The looming threats of climate change and its effects on the proliferation of mycotoxigenic fungi and the subsequent contamination of staple food and feed chains with mycotoxins, and the emergence of masked mycotoxins are seen as new global challenges that further impact on the management strategies for mycotoxins. 5 CONC LU SI O N S A N D T H E WAY FORWA R D As part of a good IPM, biological control of phytopathogenic organisms has been widely applied in Malaysia with all the four main biological control strategies (that is, natural, importation, augmentation, and conservation) have been documented. The role of natural enemies in controlling native pests has been recognized in economically important crops especially oil palm, cocoa, coconut, and rice. However, the same is not entirely true for mycotoxigenic fungi, against which the application of biological control is almost nonexistent. Based on the severely limited data, we conclude that this could either be due to the fact that (a) this particular application is totally new in Malaysia and is not yet recognized as an important research area, (b) ongoing trials are not yet conclusive and published, or (c) the safety consideration and risk analysis on whether or not to introduce biological control in Malaysian agro-ecosystem is taking longer than anticipated. This is rather perplexing because (a) Malaysian tropical climate is conducive for fungal proliferation including that of the mycotoxigenic fungi, (b) reports on mycotoxin contamination of Malaysian foods and crop commodities are abound, (c) Malaysia does have legislations for mycotoxin contamination, (d) in other parts of the world, the application of biological control agents to combat mycotoxigenic fungi is more widely practiced and accepted, and (e) the application of biological control agents against plant diseases has long and successful history in Malaysia. To fully utilize the abundant potentials of biological control approaches against mycotoxigenic fungi in the effort to minimize the impact and exposure of humans and animals to these toxic compounds, the followings are henceforth suggested: 1. Further and deeper investigation on the development and application of biological control against mycotoxigenic fungi and mycotoxin contamination using indigenous isolates; 2. As most studies have focused on the use of effective biological control agents against plant diseases, it may be beneficial to also screen these for mycotoxin control; 3. Since fungal growth and mycotoxin production are not always parallel, care and consideration when selecting and developing a biological control agent should be taken to avoid mycotoxin stimulation (instead of mycotoxin inhibition). ACKNOW LEDGMENTS The authors would like to acknowledge the financial support received from the Malaysian Ministry of Education (MOE) under the High Impact Centre of Excellence (HICoE) grant scheme (HICoE/ITAFoS/2017/FS9). The first author also thanks the School of Graduate Studies, Universiti Putra Malaysia for her PhD studentship under the Graduate Research Fund (GRF) scheme 2017–2020. CO N F L I C T O F I N T E R E ST The authors declare no conflict of interest. AU T H O R CO N T R I B U T I O N S S. N. E. Yazid was involved in drafting the entire work. S. Jinap, N. Magan, and N. I. P. Samsudin were involved in critically revising the “mycotoxigenic fungi” contents. S. I. Ismail was involved in critically revising the “phytopathogenic organisms” contents. O RC I D https://orcid.org/0000-0002-8369-9536 Selamat Jinap Naresh Magan https://orcid.org/0000-0002-5002-3564 Nik Iskandar Putra Samsudin https://orcid.org/0000-0002-2756-0142 REFERENCES Abdul Kadir, H. B., Payne, C. C., Crook, N. E., Fenlon, J. S., & Winstanley, D. (1999). The comparative susceptibility of the diamondback moth Plutella xylostella and some other major lepidopteran pests of BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… brassica crops to a range of baculoviruses. Biocontrol Science and Technology, 9, 421–433. https://doi.org/10.1080/09583159929686 Abed-Ashtiani, F., Arzanlou, M., Nasehi, A., Kadir, J., Vadamalai, G., & Azadmard-Damirchi, S. (2018). Plant tonic, a plant-derived bioactive natural product, exhibits antifungal activity against rice blast disease. Industrial Crops and Products, 112, 105–112. https://doi.org/10.1016/j.indcrop.2017.11.013 Abood, F., Bajwa, G. A., Ibrahim, Y. B., & Sajap, A. S. (2010). Pathogenicity of Beauveria bassiana against the tiger moth, Atteva sciodoxa (Lepidoptera: Yponomeutidae). Journal of Entomology, 7, 19–32. https://doi.org/10.3923/je.2010.19.32 Adnan, N. A., Bakar, S., Mazlan, A. H., Yusoff, Z. M., & Rasam, A. R. A. (2018). Comparative study of cocoa black ants temporal population distribution utilizing geospatial analysis. IOP Conference Series: Earth and Environmental Science, 117, 012041. https://doi.org/10.1088/1755-1315/117/1/012041 Afsah-Hejri, L., Jinap, S., Hajeb, P., Radu, S., & Shakibazadeh, S. (2013). A review on mycotoxins in food and feed: Malaysia case study. Comprehensive Reviews in Food Science and Food Safety, 12, 629–651. https://doi.org/10.1111/1541-4337.12029 Afsharmanesh, H., Perez-Garcia, A., Zeriouh, H., Ahmadzadeh, M., & Romero, D. (2018). Aflatoxin degradation by Bacillus subtilis UTB1 is based on production of an oxidoreductase involved in bacilysin biosynthesis. Food Control, 94, 48–55. https://doi.org/ 10.1016/j.foodcont.2018.03.002 Agbetiameh, D., Ortega-Beltran, A., Awuah, R. T., Atehnkeng, J., Cotty, P. J., & Bandyopadhyay, R. (2018). Prevalence of aflatoxin contamination in maize and groundnut in Ghana: Population structure, distribution, and toxigenicity of the causal agents. Plant Disease, 102, 764–772. https://doi.org/10.1094/PDIS-05-17-0749-RE Ahmad, K., & Ahmadu, T. (2017). Prospect and potential of Burkholderia sp. against Phytophthora capsici Leonian: A causative agent for foot rot disease of black pepper. Agriculturally Important Microbes for Sustainable Agriculture, 2, 343–374. https://doi.org/10.1007/978-981-10-5343-6_12 Ahmad, T. T. M. A., & Suntharalingam, C. (2009). Transformation and economic growth of the Malaysian agricultural sector. Economic and Technology Management Review, 4, 1–10 Ahmad-Zaidi, A. I., Ghazali, M. A. A., Nik-Muhammad, N. A., Sazali, N. S., Mahror, N., Yazid, S. N. E., … Samsudin, N. I. P. (2020). Does manufacturers’ size affect the prevalence of mycobiota and occurrence of mycotoxins in spices and spice-based products? World Mycotoxin Journal. https://doi.org/10.3920/WMJ2019.2487 Akter, S., Kadir, J., Juraimi, A. S., & Saud, H. M. (2016). In vitro evaluation of Pseudomonas bacterial isolates from rice phylloplane for biocontrol of Rhizoctonia solani and plant growth promoting traits. Journal of Environmental Biology, 37, 597–602. Akter, S., Kadir, J., Juraimi, A. S., Saud, H. M., & Elmahdi, S. (2014). Isolation and identification of antagonistic bacteria from phylloplane of rice as biocontrol agents for sheath blight. Journal of Environmental Biology, 35, 1095–1100. Alaniz Zanon, M. S., Clemente, M. P., & Chulze, S. N. (2018). Characterization and competitive ability of non-aflatoxigenic Aspergillus flavus isolated from the maize agro-ecosystem in Argentina as potential aflatoxin biocontrol agents. International Journal of Food Microbiology, 277, 58–63. https://doi.org/10.1016/ j.ijfoodmicro.2018.04.020 Alexander, A., Abdullah, S., Rossall, S., & Chong, K. P. (2017). Evaluation of the efficacy and mode of action of biological control for 661 suppression of Ganoderma boninense in oil palm. Pakistan Journal of Botany, 49, 1193–1199. Ali, H. Z., & Nadarajah, K. (2013). Evaluating the efficacy of Trichoderma isolates and Bacillus subtilis as biological control agents against Rhizoctonia solani. Research Journal of Applied Sciences, 8, 72–81. https://doi.org/10.3923/rjasci.2013.72.81 Ali, N., Hashim, N. H., & Shuib, N. S. (2015). Natural occurrence of aflatoxins and ochratoxin A in processed spices marketed in Malaysia. Food Additives and Contaminants - Part A Chemistry, Analysis, Control, Exposure and Risk Assessment, 32, 518–532. https://doi.org/10.1080/19440049.2015.1011712 Alloub, H., Juraimi, A. S., Kadir, J., Sastroutomo, S., & Begum, M. (2009). Field efficacy of Exserohilum prolatum—A potential mycoherbicide for biological control of itch grass (Rottboellia cochinchinensis). Journal of Biological Sciences, 9, 119–127. https://doi.org/10.3923/jbs.2009.119.127 Al-Saad, L. A., Al-Badran, A. I., Al-Jumayli, S. A., Magan, N., & Rodríguez, A. (2016). Impact of bacterial biocontrol agents on aflatoxin biosynthetic genes, aflD and aflR expression, and phenotypic aflatoxin B1 production by Aspergillus flavus under different environmental and nutritional regimes. International Journal of Food Microbiology, 217, 123–129. https://doi.org/10.1016/ j.ijfoodmicro.2015.10.016 Alsharif, A. M. A., Choo, Y. M., & Tan, G. H. (2019). Detection of five mycotoxins in different food matrices in the Malaysian market by using validated liquid chromatography electrospray ionization triple quadrupole mass spectrometry. Toxins, 11, 196. https://doi.org/10.3390/toxins11040196 Angel, L. P. L., Sundram, S., Ping, B. T. Y., Yusof, M. T., & Ismail, I. S. (2018). Profiling of anti-fungal activity of Trichoderma virens 159C involved in biocontrol assay of Ganoderma boninense. Journal of Oil Palm Research, 30, 83–93. https://doi.org/10.21894/jopr.2017. 0009 Angel, L. P. L., Yusof, M. T., Ismail, I. S., Ping, B. T. Y., Mohamed Azni, I. N. A., Kamarudin, N. H., & Sundram, S. (2016). An in vitro study of the antifungal activity of Trichoderma virens 7b and a profile of its non-polar antifungal components released against Ganoderma boninense. Journal of Microbiology, 54, 732–744. https://doi.org/ 10.1007/s12275-016-6304-4 Aslani, F., Juraimi, A. S., Ahmad-Hamdani, M. S., Alam, M. A., Hashemi, F., Omar, D., & Hakim, M. A. (2015). Phytotoxic interference of volatile organic compounds and water extracts of Tinospora tuberculata Beumee on growth of weeds in rice fields. South African Journal of Botany, 100, 132–140. https://doi.org/ 10.1016/j.sajb.2015.04.011 Awad, H. M., El-Enshasy, H. A., Hanapi, S. Z., Hamed, E. R., & Rosidi, B. (2014). A new chitinase-producer strain Streptomyces glauciniger WICC-A03: Isolation and identification as a biocontrol agent for plants phytopathogenic fungi. Natural Product Research, 28, 2273– 2277. https://doi.org/10.1080/14786419.2014.939083 Awla, H. K., Kadir, J., Othman, R., Rashid, T. S., Hamid, S., & Wong, M. Y. (2017). Plant growth-promoting abilities and biocontrol efficacy of Streptomyces sp. UPMRS4 against Pyricularia oryzae. Biological Control, 112, 55–63. https://doi.org/10.1016/ j.biocontrol.2017.05.011 Azadeh, B. F., Sariah, M., & Wong, M. Y. (2010). Characterization of Burkholderia cepacia genomovar I as a potential biocontrol agent of Ganoderma boninense in oil palm. African Journal of Biotechnology, 9, 3542–3548. 662 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Bakeri, S. A., Ali, S. R. A., Tajuddin, N. S., & Kamaruzzaman, N. E. (2009). Efficacy of entomopathogenic fungi, Paecilomyces spp., in controlling the oil palm bag worm, Pteroma pendula (Joannis). Journal of Oil Palm Research, 21, 693–699. Balendres, M. A. O., Karlovsky, P., & Cumagun, C. J. R. (2019). Mycotoxigenic fungi and mycotoxins in agricultural crop commodities in the Philippines: A review. Foods, 8, 249. https://doi.org/ 10.3390/foods8070249 Battisti, D. S., & Naylor, R. L. (2009). Historical warnings of future food insecurity with unprecedented seasonal heat. Science, 323, 240–244. https://doi.org/10.1126/science.1164363 Bebber, D. P., Ramotowski, M. A. T., & Gurr, S. J. (2013). Crop pests and pathogens move polewards in a warming world. Nature Climate Change, 3, 985–988. https://doi.org/10.1038/NCLIMATE1990 Begum, M. M., Sariah, M., Puteh, A. B., Zainal Abidin, M. A., Rahman, M. A., & Siddiqui, Y. (2010). Field performance of bio-primed seeds to suppress Colletotrichum truncatum causing damping-off and seedling stand of soybean. Biological Control, 53, 18–23. https://doi.org/10.1016/j.biocontrol.2009.12.001 Beneduzi, A., Ambrosini, A., & Passaglia, L. M. P. (2012). Plant growthpromoting rhizobacteria (PGPR): Their potential as antagonists and biocontrol agents. Genetics and Molecular Biology, 35, 1044–1051. http://doi.org/10.1590/S1415-47572012000600020 Bennett, J. W., & Klich, M. (2003). Mycotoxins. Clinical Microbiology Reviews, 16, 497–516. https://doi.org/10.1128/CMR.16.3.497516.2003 Berthiller, F., Maragos, C. M., & Dall’Asta, C. (2016). Chapter 1: Introduction to masked mycotoxins. In C. Dall’Asta, & F. Berthiller (Eds.), Masked mycotoxins in food: Formation, occurrence and toxicological relevance (pp. 1–13). Cambridge, UK: RSC Publishing. https://doi.org/10.1039/9781782622574-00001 Berthiller, F., Schuhmacher, R., Adam, G., & Krska, R. (2009a). Formation, determination and significance of masked and other conjugated mycotoxins. Analytical and Bioanalytical Chemistry, 395, 1243–1252. https://doi.org/10.1007/s00216-009-2874-x Berthiller, F., Dall’Asta, C., Corradini, R., Marchelli, R., Sulyok, M., Krska, R., … Schuhmacher, R. (2009b). Occurrence of deoxynivalenol and its 3-𝛽-D-glucoside in wheat and maize. Food Additives and Contaminants, 26, 507–511. https://doi.org/10.1080/ 02652030802555668 Bhat, R., Rai, R. V., & Karim, A. A. (2010). Mycotoxins in food and feed: Present status and future concerns. Comprehensive Reviews in Food Science and Food Safety, 9, 57–81. https://doi.org/10.1111/ j.1541-4337.2009.00094.x Bivi, M. R., Farhana, M. S., & Khairulmazmi, A. (2010). Control of Ganoderma boninense: A causal agent of basal stem rot disease in oil palm with endophyte bacteria in vitro. International Journal of Agriculture and Biology, 12, 833–839. Blount, W. P. (1961). Turkey “X” disease. Journal of British Turkey Federation, 9, 55–58. Bommarco, R., Miranda, F., Bylund, H., & Björkman, C. (2011). Insecticides suppress natural enemies and increase pest damage in cabbage. Journal of Economic Entomology, 104, 782–791. https://doi.org/ 10.1603/ec10444 Borisade, O. A., & Magan, N. (2015). Resilience and relative virulence of strains of entomopathogenic fungi under interactions of abiotic stress. African Journal of Microbiology Research, 9, 988–1000. https://doi.org/10.5897/AJMR2015.7416 Botana, L. M., & Sainz, M. J. (2015). Climate change and mycotoxins. Berlin, Germany: De Gruyter. Braun, H., Woitsch, L., Hetzer, B., Geisen, R., Zange, B., & SchmidtHeydt, M. (2018). Trichoderma harzianum: Inhibition of mycotoxin producing fungi and toxin biosynthesis. International Journal of Food Microbiology, 280, 10–16. https://doi.org/10.1016/ j.ijfoodmicro.2018.04.021 Brown, R. L., Cotty, P. J., & Cleveland, T. E. (1991). Reduction in aflatoxin content of maize by atoxigenic strains of Aspergillus flavus. Journal of Food Protection, 54, 623–626. https://doi.org/10.4315/ 0362-028X-54.8.623 Cao, Y., Xu, Z., Ling, N., Yuan, Y., Yang, X., Chen, L., … Shen, Q. (2012). Isolation and identification of lipopeptides produced by Bacillus subtilis SQR 9 for suppressing Fusarium wilt of cucumber. Scientia Horticulturae, 35, 32–39. https://doi.org/ 10.1016/j.scienta.2011.12.002 Chinajariyawong, A., Clarke, A. R., Jirasurat, M., Kritsaneepiboon, S., Lahey, H. A., Vijaysegaran, S., & Walter, G. H. (2000). Survey of opiine parasitoids of fruit flies (Diptera: Tephritidae) in Thailand and Malaysia. Raffles Bulletin of Zoology, 48, 71–101. Cleveland, T. E., Dowd, P. F., Desjardins, A. E., Bhatnagar, D., & Cotty, P. J. (2003). United States Department of Agriculture—agricultural research service research on pre-harvest prevention of mycotoxins and mycotoxigenic fungi in US crops. Pest Management Science, 59, 629–642. https://doi.org/10.1002/ps.724 Colmenarez, Y. C., Corniani, N., Jahnke, S. M., Sampaio, M. V., & Vásquez, C. (2018). Use of parasitoids as a biocontrol agent in the neotropical region: Challenges and potential. In H. K. Baimey, & N. Hamamouch & Y. A. Kolombia, (Eds.), Horticulture (pp. 1– 23). London, UK: IntechOpen. https://doi.org/10.5772/intechopen. 80720 Damann, K. E., Jr. (2015). Atoxigenic Aspergillus flavus biological control of aflatoxin contamination: What is the mechanism? World Mycotoxin Journal, 8, 235–244. https://doi.org/10.3920/ WMJ2014.1719 Desjardins, A. E. (2006). Fusarium mycotoxins: Chemistry, genetics, and biology. St. Paul, MN: The American Phytopathological Society. Department of Agriculture (DOA). (2019). Crop statistics report. Putrajaya, Malaysia: Ministry of Agriculture and Agro-based Industry. Retrieved from http://www.doa.gov.my/index.php/pages/view/ 622?mid=239 Duffy, B., Schouten, A., & Raaijmakers, J. M. (2003). Pathogen selfdefence: Mechanisms to counteract microbial antagonism. Annual Review of Phytopathology, 41, 501–538. https://doi.org/10.1146/ annurev.phyto.41.052002.095606 Edward, E. J., King, W. S., Teck, S. L. C., Jiwan, M., Aziz, Z. F. A., Kundat, F. R., … Majid, N. M. A. (2013). Antagonistic activities of endophytic bacteria against Fusarium wilt of black pepper (Piper nigrum). International Journal of Agriculture and Biology, 15, 291– 296. Eggleton, P., & Belshaw, R. (1992). Insect parasitoids: An evolutionary overview. Philosophical Transactions of the Royal Society B, 337, 1–20. https://doi.org/10.1098/rstb.1992.0079 El-Ghany, N. M. A. (2019). Semiochemicals for controlling insect pests. Journal of Plant Protection Research, 59, 1–11. https://doi.org/ 10.24425/jppr.2019.126036_rfseq1 El-Mabrok, A. S. W., Hassan, Z., Mokhtar, A. M., Hussain, K. M. A., & Kahar, F. K. S. B. A. (2012). Screening of lactic acid BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… bacteria as biocontrol against (Colletotrichum capsici) on chili Bangi. Research Journal of Applied Sciences, 7, 466–473. https://doi. org/10.3923/rjasci.2012.466.473 Earth System Research Laboratory (ESRL). (2019). Trends in atmospheric carbon dioxide. Retrieved form http://www.esrl.noaa.gov/ gmd/ccgg/trends/global.html Fahmi, Z., Abu-Samah, B., & Abdullah, H. (2013). Paddy industry and paddy farmers’ well-being: A success recipe for agriculture industry in Malaysia. Asian Social Science, 9, 177–181. https://doi.org/ 10.5539/ass.v9n3p177 Food and Agriculture Organization (FAO). (2009). Declaration of the world food summit on food security, Nov 16th – 18th. Rome, Italy: Author. Fernandes, E. G., Valério, H. M., Feltrin, T., & Van Der Sand, S. T. (2012). Variability in the production of extracellular enzymes by entomopathogenic fungi grown on different substrates. Brazilian Journal of Microbiology, 43, 827–833. https://doi.org/10.1590/ S1517-83822012000200049 Fisher, M. C., Henk, D. A., Briggs, C. J., Brownstein, J. S., Madoff, L. C., McCraw, S. L., & Gurr, S. J. (2012). Emerging fungal threats to animal, plant and ecosystem health. Nature, 484, 186–194. https://doi.org/10.1038/nature10947 Frische, T., Egerer, S., Matezki, S., Pickl, C., & Wogram, J. (2018). 5-point programme for sustainable plant protection. Environmental Sciences Europe, 30, 8. https://doi.org/10.1186/s12302-0180136-2 Furlong, M. J., Pell, J. K., Pek Choo, O., & Abdul Rahman, S. (1995). Field and laboratory evaluation of a sex pheromone trap for the autodissemination of the fungal entomopathogen Zoophthora radicans (Entomophthorales) by the diamondback moth, Plutella xylostella (Lepidoptera: Yponomeutidae). Bulletin of Entomological Research, 85, 331–337. https://doi.org/ 10.1017/S0007485300036051 Galaverna, G., Dallsta, C., Mangia, M. A., Dossena, A., & Marchelli, R. (2009). Masked mycotoxins: An emerging issue for food safety. Czech Journal of Food Sciences, 27(Special Issue), S89–S92. https://doi.org/10.17221/1064-CJFS Getha, K., & Vikineswary, S. (2002). Antagonistic effects of Streptomyces violaceusniger strain G10 on Fusarium oxysporum f. sp. cubense race 4: Indirect evidence for the role of antibiosis in the antagonistic process. Journal of Industrial Microbiology and Biotechnology, 28, 303–310. https://doi.org/10.1038/sj/jim/70002 47 Ghani, I. A., Dieng, H., Hassan, Z. A. A., Ramli, N., Kermani, N., Satho, T., … Ahmad, A. H. (2013). Pathogenicity of a microsporidium isolate from the diamondback moth against noctuid moths: Characterization and implications for microbiological pest management. PLoS ONE, 8, e81642. https://doi.org/10.1371/journal.pone.0081642 Gilal, A. A., Muhamad, R., Omar, D., Aziz, N. A. A., & Gnanasegaram, M. (2016). Foes can be friends: Laboratory trials on invasive apple snails, Pomacea spp. Preference to invasive weed, Limnocharis flava (L.) Buchenau compared to rice, Oryza sativa L. Pakistan Journal of Zoology, 48, 673–679. Godfray, H. C. J., Beddington, J. R., Crute, I. R., Haddad, L., Lawrence, D., Muir, J. F., … Toulmin, C. (2010). Food security: The challenge of feeding 9 billion people. Science, 327, 812–818. https://doi.org/ 10.1126/science.1185383 Gregory, P. J., Ingram, J. S. I., & Brklacich, M. (2005). Climate change and food security. Philosophical Transactions of the Royal 663 Society B: Biological Sciences, 360, 2139–2148. https://doi.org/ 10.1098/rstb.2005.1745 Guimarães, A., Santiago, A., Teixeira, J. A., Venâncio, A., & Abrunhosa, L. (2018). Anti-aflatoxigenic effect of organic acids produced by Lactobacillus plantarum. International Journal of Food Microbiology, 264, 31–38. https://doi.org/10.1016/j.ijfoodmicro.2017.10.025 Hamizah, H., Mahmud, T. M. M., Ahmad, S. H., & Kamaruzaman, S. (2013). Screening of antagonistic yeast for biological control activity against anthracnose (Colletotrichum gloeosporioides) in ‘Frangi’ papaya. Acta Horticulturae, 1012, 739–744. https://doi.org/ 10.17660/ActaHortic.2013.1012.99 Harman, G. E. (2006). Overview of mechanisms and uses of Trichoderma spp. Phytopathology, 96, 190–194. https://doi.org/10.1094/ PHYTO-96-0190 Ho, C. T., & Khoo, K. C. (1997). Partners in biological control of cocoa pests: Mutualism between Dolichoderus thoracicus (Hymenoptera: Formicidae) and Cataenococcus hispidus (Hemiptera: Pseudococcidae). Bulletin of Entomological Research, 87, 461–470. https://doi.org/10.1017/S0007485300041328 Hoe, P. K., Bong, C. F. J., Jugah, K., & Rajan, A. (2009). Evaluation of Metarhizium anisopliae var. anisopliae (Deuteromycotina: Hyphomycete) isolates and their effects on subterranean termite Coptotermes curvignathus (Isoptera: Rhinotermitidae). American Journal of Agricultural and Biological Science, 4, 289–297. Holden, M. H., Ellner, S. P., Lee, D.-H., Nyrop, J. P., & Sanderson, J. P. (2012). Designing an effective trap cropping strategy: The effects of attraction, retention and plant spatial distribution. Journal of Applied Ecology, 49, 715–722. https://doi.org/ 10.1111/j.1365-2664.2012.02137.x Horn, B. W., & Dorner, J. W. (2011). Evaluation of different genotypes of nontoxigenic Aspergillus flavus for their ability to reduce aflatoxin contamination in peanuts. Biocontrol Science and Technology, 21, 865–876. https://doi.org/10.1080/09583157.2011.559308 Hoster, F., Schmitz, J. E., & Daniel, R. (2005). Enrichment of chitinolytic microorganisms: Isolation and characterization of a chitinase exhibiting antifungal activity against phytopathogenic fungi from a novel Streptomyces strain. Applied Microbiology and Biotechnology, 66, 434–442. https://doi.org/10.1007/s00253-004-1664-9 Huang, C., Jha, A., Sweany, R., DeRobertis, C., & Damann, K. E., Jr. (2011). Intraspecific aflatoxin inhibition in Aspergillus flavus is thigmoregulated, independent of vegetative compatibility group and is strain dependent. PLoS ONE, 6, e23470. https://doi.org/10.1371/journal.pone.0023470 Hussein, H. S., & Brasel, J. M. (2001). Toxicity, metabolism, and impact of mycotoxins on humans and animals. Toxicology, 167, 101–134. https://doi.org/10.1016/S0300-483X(01)00471-1 Hussein, M. Y. (1984). A spray technique for mass release of eggs of Micromus tasmaniae Walker (neuroptera: Hemerobiidae). Crop Protection, 3, 369–378. https://doi.org/10.1016/0261-2194(84)90043-7 Ibrahim, N. J., Shariff, S., Idris, A. B., Md-Zain, B. M., Suhana, Y., Roff, M. N., & Yaakop, S. (2013). Phylogenetic tree construction in reconfirmation of parasitoid species (Braconidae: Opiinae), reared from fruit flies (Bactrocera papayae) infesting star fruit (Averrhoa carambola) based on mitochondrial 16S rRNA sequences. Pertanika Journal of Tropical Agricultural Science, 36, 345–358. Ibrahim, Y. B., & Kueh, T. F. (2013). Biological performance of Menochilus sexmaculatus fabricius (coleoptera: Coccinellidae) upon exposure to sublethal concentration of imidacloprid. Pertanika Journal of Tropical Agricultural Science, 36, 51–59. 664 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Ibrahim, Y. B., & Low, W. (1993). Potential of mass-production and field efficacy of isolates of the entomopathogenic fungi Beauveria bassiana and Paecilomyces fumosoroseus against Plutella xylostella. International Journal of Pest Management, 39, 288–292. https://doi.org/10.1080/09670879309371807 Jalili, M., & Jinap, S. (2012). Natural occurrence of aflatoxins and ochratoxin A in commercial dried chili. Food Control, 24, 160–164. https://doi.org/10.1016/j.foodcont.2011.09.020 Jamian, S. (2017). Role of beneficial plants in improving performance of predators of oil palm bagworm (PhD thesis). Universiti Putra Malaysia, Seri Kembangan, Malaysia. Jamian, S., Norhisham, A., Ghazali, A., Zakaria, A., & Azhar, B. (2017). Impacts of two species of predatory Reduviidae on bagworms in oil palm plantations. Insect Science, 24, 285–294. https://doi.org/10.1111/1744-7917.12309 Jessica, J. J., Peng, T. L., Sajap, A. S., Lee, S. H., & Syazwan, S. A. (2019). Evaluation of the virulence of entomopathogenic fungus, Isaria fumosorosea isolates against subterranean termites Coptotermes spp. (Isoptera: Rhinotermitidae). Journal of Forestry Research, 30, 213–218. https://doi.org/10.1007/s11676-018-0614-9 Joffe, A. Z. (1971). Chapter 5: Alimentary toxic aleukia. In S. Kadis, A. Ciegler, & S. J. Ajl (Eds.), Microbial toxins (Vol. 7, pp. 139–189). New York, NY: Academic Press. Kachapulula, P. W., Akello, J., Bandyopadhyay, R., & Cotty, P. J. (2017). Aflatoxin contamination of groundnut and maize in Zambia: Observed and potential concentrations. Journal of Applied Microbiology, 122, 1471–1482. https://doi.org/10.1111/jam.13448 Kadir, J., Sajili, M. H., Juraimi, A. S., & Napis, S. (2008). Effect of Exserohilum monoceras (Drechslera) Leonard & Suggs on the competitiveness of Echinocloa cruss-galli (L.) P. Beauv. Pertanika Journal of Tropical Agricultural Science, 31, 19–26. Kapranas, A., Tena, A., & Luck, R. F. (2012). Dynamic virulence in a parasitoid wasp: The influence of clutch size and sequential oviposition on egg encapsulation. Animal Behaviour, 83, 833–838. https://doi.org/10.1016/j.anbehav.2012.01.004 Kaufman, L. V., & Wright, M. G. (2017). Assessing probabilistic risk assessment approaches for insect biological control introductions. Insects, 8, 67. https://doi.org/10.3390/insects8030067 Kermani, N., Hassan Abu, A. Z., Suhaimi, A., Abuzid, I., … Attia, M., & Abd Ghani, I. (2014). Parasitism performance and fitness of Cotesia vestalis (Hymenoptera: Braconidae) infected with Nosema sp. (Microsporidia: Nosematidae): Implications in integrated pest management strategy. PLoS ONE, 9, e100671. https://doi.org/10.1371/journal.pone.0100671 Kermani, N., Abu-Hassan, Z. A., Dieng, H., Ismail, N. F., Attia, M., & Abd Ghani, I. (2013). Pathogenicity of Nosema sp. (Microsporidia) in the diamondback moth, Plutella xylostella (Lepidoptera: Plutellidae). PLoS ONE, 8, e62884. https://doi.org/10.1371/journal.pone.0062884 Keswani, C., Mishra, S., Sarma, B. K., Singh, S. P., & Singh, H. B. (2014). Unravelling the efficient applications of secondary metabolites of various Trichoderma spp. Applied Microbiology and Biotechnology, 98, 533–544. https://doi.org/10.1007/s00253-013-5344-5 Khalili, E., Javed, M. A., Huyop, F., Rayatpanah, S., Jamshidi, S., & Wahab, R. A. (2016). Evaluation of Trichoderma isolates as potential biological control agent against soybean charcoal rot disease caused by Macrophomina phaseolina. Biotechnology and Biotechnological Equipment, 30, 479–488. https://doi.org/10.1080/ 13102818.2016.1147334 Khan, M. R., & Ram, G. M. (2014). Non-pesticidal management (NPM) of pests and pesticides. Ecology, Environment and Conservation, 20, S95–S98. Khayoon, W. S., Saad, B., Lee, T. P., & Salleh, B. (2012). High performance liquid chromatographic determination of aflatoxins in chilli, peanut and rice using silica based monolithic column. Food Chemistry, 133, 489–496. https://doi.org/10.1016/j.foodchem. 2012.01.010 Khayoon, W. S., Saad, B., Salleh, B., Ismail, N. A., Manaf, N. H. A., & Latiff, A. A. (2010). A reversed phase high performance liquid chromatography method for the determination of fumonisins B1 and B2 in food and feed using monolithic column and positive confirmation by liquid chromatography/tandem mass spectrometry. Analytica Chimica Acta, 679, 91–97. https://doi.org/ 10.1016/j.aca.2010.09.008 Khayoon, W. S., Saad, B., Yan, C. B., Hashim, N. H., Ali, A. S. M., Salleh, M. I., & Salleh, B. (2010). Determination of aflatoxins in animal feeds by HPLC with multifunctional column clean-up. Food Chemistry, 118, 882–886. https://doi.org/10.1016/j.foodchem. 2009.05.082 Köhl, J., Kolnarr, R., & Ravensberg, W. J. (2019). Mode of action of biocontrol agents against plant diseases: Relevance beyond efficacy. Frontiers in Plant Science, 10, 845. https://doi.org/10.3389/ fpls.2019.00845 Köhl, J., Postma, J., Nicot, P., Ruocco, M., & Blum, B. (2011). Stepwise screening of microorganisms for commercial use in biological control of plant-pathogenic fungi and bacteria. Biological Control, 57, 1–12. https://doi.org/10.1016/j.biocontrol.2010.12.004 Kota, M. F., Husaini, A. A. S. A., Lihan, S., Hussain, M. H. M., & Roslan, H. A. (2015). In vitro antagonism of Phytophthora capsici and Fusarium solani by bacterial isolates from Sarawak. Malaysian Journal of Microbiology, 11, 137–143. https://doi.org/10.21161/mjm. 12514 Koul, O. (2008). Phytochemicals and insect control: An antifeedant approach. Critical Reviews in Plant Sciences, 27, 1–24. https://doi. org/10.1080/07352680802053908 Ksenija, N. (2018). Mycotoxins—Climate impact and steps to prevention based on prediction. Acta Veterinaria, 68, 1–15. https://doi.org/ 10.2478/acve-2018-0001 Kundat, F. R., Shen, L. H., & Ahmed, O. H. (2010). Incorporation of bentazone with Exserohilum rostratum for controlling Cyperus iria. American Journal of Agricultural and Biological Science, 5, 210– 214. https://doi.org/10.3844/ajabssp.2010.210.214 Law, J. W., Ser, H. L., Khan, T. M., Chuah, L. H., Pusparajah, P., Chan, K. G., … Lee, L. H. (2017). The potential of Streptomyces as biocontrol agents against the rice blast fungus, Magnaporthe oryzae (Pyricularia oryzae). Frontiers in Microbiology, 8, 3. https://doi.org/10.3389/fmicb.2017.00003 Lee, T. P., Saad, B., Ng, E. P., & Salleh, B. (2012). Zeolite Linde Type L as micro-solid phase extraction sorbent for the high performance liquid chromatography determination of ochratoxin A in coffee and cereal. Journal of Chromatography A, 1237, 46–54. https://doi.org/10.1016/j.chroma.2012.03.031 Li, J., Yang, Q., Zhao, L. H., Zhang, S. M., Wang, Y. X., & Zhao, X. Y. (2009). Purification and characterization of a novel antifungal protein from Bacillus subtilis strain B29. Journal of Zhejiang University - SCIENCE B, 10, 264–272. https://doi.org/10.1631/jzus. B0820341 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Lim, G. T., Kirton, L. G., Salom, S. M., Kok, L. T., Fell, R. D., & Pfeiffer, D. G. (2008). Mahogany shoot borer control in Malaysia and prospects for biocontrol using weaver ants. Journal of Tropical Forest Science, 20, 147–155. Louda, S. M., Pemberton, R. W., Johnson, M. T., & Follett, P. A. (2003). Nontarget effects—the Achilles’ heel of biological control? Retrospective analyses to reduce risk associated with biocontrol introductions. Annual Review of Entomology, 48, 365– 396. https://doi.org/10.1146/annurev.ento.48.060402.102800 Lutz, M. P., Feichtinger, G., Défago, G., & Duffy, B. (2003). Mycotoxigenic Fusarium and deoxynivalenol production repress chitinase gene expression in the biocontrol agent Trichoderma atroviride P1. Applied and Environmental Microbiology, 69, 3077–3084. https://doi.org/10.1128/AEM.69.6.3077-3084. 2003 Malaysian Food Regulations. (1985). Fifteenth schedule (Regulation 39): Microorganisms and their toxins. Table II: Mycological contaminant. Retrieved from http://fsq.moh.gov.my/v6/xs/page.php?id=72 Marchera, G., & Ndwiga, J. (2015). Estimation of the potential adoption of Aflasafe among smallholder maize farmers in lower eastern Kenya. African Journal of Agricultural and Resource Economics, 10, 72–85. Mari, M., Bautista-Baños, S., & Sivakumar, D. (2016). Decay control in the postharvest system: Role of microbial and plant volatile organic compounds. Postharvest Biology and Technology, 122, 70– 81. https://doi.org/10.1016/j.postharvbio.2016.04.014 Mastoi, M. I., Adam, N. A., Muhamad, R., Ghani, I. A., Gilal, A. A., Khan, J., … Sahito, J. G. M. (2018). Efficiency of Acerophagus papayae on different host stage combinations of papaya mealybug, Paracoccus marginatus. Pakistan Journal of Agricultural Sciences, 55, 375–379. https://doi.org/10.21162/PAKJAS/18.5600 Mauro, A., Garcia-Cela, E., Pietri, A., Cotty, P. J., & Battilani, P. (2018). Biological control products for aflatoxin prevention in Italy: Commercial field evaluation of atoxigenic Aspergillus flavus active ingredients. Toxins, 10, 30. https://doi.org/10.3390/toxins10010030 Medina, A., Mohale, S., Samsudin, N. I. P., Rodriguez-Sixtos, A., Rodriguez, A., & Magan, N. (2017). Biocontrol of mycotoxins: Dynamics and mechanisms of action; a review. Current Opinion in Food Science, 17, 41–48. https://doi.org/10.1016/j.cofs.2017.09.008 Medina, A., Akbar, A., Baazeem, A., Rodriguez, A., & Magan, N. (2017). Climate change, food security and mycotoxins: Do we know enough? Fungal Biology Reviews, 31, 143–154. https://doi.org/ 10.1016/j.fbr.2017.04.002 Medina, A., Rodríguez, A., & Magan, N. (2015). Climate change and mycotoxigenic fungi: Impacts on mycotoxin production. Current Opinion in Food Science, 5, 99–104. https://doi.org/10.1016/ j.cofs.2015.11.002 Moazami, E. F., & Jinap, S. (2009). Natural occurrence of deoxynivalenol (DON) in wheat based noodles consumed in Malaysia. Microchemical Journal, 93, 25–28. https://doi.org/10.1016/ j.microc.2009.04.003 Mohamad, S. A., Masijan, Z., Moslim, R., Sulaiman, M. R., Ming, S. C., Chuan, S. T., … Ahmad, S. N. (2017). Biological agents and insecticides to control bunch moth, Tirathaba rufivena in oil palm estates in Sarawak, Malaysia. Journal of Oil Palm Research, 29, 323–332. https://doi.org/10.21894/jopr.2017.2903.04 Mohammed, A. M., Al-Ani, L. K. T., Bekbayeva, L., & Salleh, B. (2011). Biological control of Fusarium oxysporum f. sp. cubense by Pseudomonas fluorescens and BABA in vitro. World Applied Sciences Journal, 15, 189–191. 665 Mohd Fishal, M. E., Meon, S., & Yun, W. M. (2010). Induction of tolerance to Fusarium wilt and defense-related mechanisms in the plantlets of susceptible berangan banana preinoculated with Pseudomonas sp. (UPMP3) and Burkholderia sp. (UPMB3). Agricultural Sciences in China, 9, 1140–1149. https://doi.org/10.1016/S1671-2927(09)60201-7 Moore, G. G., Lebar, M. D., & Carter-Wientjes, C. H. (2019). The role of extrolites secreted by nonaflatoxigenic Aspergillus flavus in biocontrol efficacy. Journal of Applied Microbiology, 126, 1257–1264. https://doi.org/10.1111/jam.14175 Moslim, R., Kamarudin, N., & Wahid, M. B. (2009). Pathogenicity of granule formulations of Metarhizium anisopliae against the larvae of the oil palm rhinoceros beetle, Oryctes rhinoceros (L.). Journal of Oil Palm Research, 21, 602–612. Moslim, R., Kamarudin, N., Ghani, I. A., Wahid, M. B., Jackson, T. A., Tey, C. C., & Ahdly, A. M. (2011). Molecular approaches in the assessment of Oryctes rhinoceros virus for the control of rhinoceros beetle in oil palm plantations. Journal of Oil Palm Research, 23, 1096–1109. Moslim, R., Kamarudin, N., & Wahid, M. B. (2011). Trap for the auto dissemination of Metarhizium anisopliae in the management of rhinoceros beetle, Oryctes rhinoceros. Journal of Oil Palm Research, 23, 1011–1017. Nadira, A. F., Rosita, J., Norhaizan, M. E., & Redzwan, S. M. (2017). Screening of aflatoxin M1 occurrence in selected milk and dairy products in Terengganu, Malaysia. Food Control, 73, 209–214. https://doi.org/10.1016/j.foodcont.2016.08.004 Naidu, Y., Idris, A. S., Madihah, A. Z., & Kamarudin, N. (2016). In vitro antagonistic interactions between endophytic basidiomycetes of oil palm (Elaeis guineensis) and Ganoderma boninense. Journal of Phytopathology, 164, 779–790. https://doi.org/10.1111/jph.12498 Naidu, Y., Meon, S., & Siddiqui, Y. (2012). In vitro and in vivo evaluation of microbial-enriched compost tea on the development of powdery mildew on melon. BioControl, 57, 827–836. https://doi.org/10.1007/s10526-012-9454-2 Naidu, Y., Siddiqui, Y., Rafii, M. Y., Saud, H. M., & Idris, A. S. (2018). Inoculation of oil palm seedlings in Malaysia with white-rot hymenomycetes: Assessment of pathogenicity and vegetative growth. Crop Protection, 110, 146–154. https://doi.org/ 10.1016/j.cropro.2018.02.018 Ng, L. C., Ngadin, A., Azhari, M., & Zahari, N. A. (2015). Potential of Trichoderma spp. As biological control agents against bakanae pathogen (Fusarium fujikuroi) in rice. Asian Journal of Plant Pathology, 9, 46–58. https://doi.org/10.3923/ajppaj.2015.46.58 Ng, S. C., Kadir, J., & Hailmi, M. S. (2012). Histological study of the interaction between Exserohilum longirostratum, barnyard grass, and rice var. MR219. Pertanika Journal of Tropical Agricultural Science, 35, 57–70. Nguyen, P. A., Strub, C., Durand, N., Alter, P., Fontana, A., & SchorrGalindo, S. (2018). Biocontrol of Fusarium verticillioides using organic amendments and their actinomycete isolates. Biological Control, 118, 55–66. https://doi.org/10.1016/j.biocontrol.2017.12.006 Nielsen, K. F., & Smedsgaard, J. (2003). Fungal metabolite screening: Database of 474 mycotoxins and fungal metabolites for dereplication by standardised liquid chromatography-UV-mass spectrometry methodology. Journal of Chromatography A, 1002, 111–136. https://doi.org/10.1016/S0021-9673(03)00490-4 National Institute of Food and Agriculture (NIFA). (2015). Global scientists meet for integrated pest management idea sharing. 666 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Retrieved from https://nifa.usda.gov/blog/global-scientists-meetintegrated-pest-management-idea-sharing Norlia, M., Nor-Khaizura, M. A. R., Selamat, J., Abu Bakar, F., Radu, S., & Chin, C. K. (2018). Evaluation of aflatoxin and Aspergillus sp. contamination in raw peanuts and peanut-based products along this supply chain in Malaysia. Food Additives and Contaminants - Part A Chemistry, Analysis, Control, Exposure and Risk Assessment, 35, 1787–1802. https://doi.org/10.1080/19440049.2018.1488276 Nourozian, J., Etebarian, H. R., & Khodakaramian, G. (2006). Biological control of Fusarium graminearum on wheat by antagonistic bacteria. Songklanakarin Journal of Science and Technology, 28, 29–38. Nur Ain Izzati, M. Z., & Abdullah, F. (2008). Disease suppression in Ganoderma-infected oil palm seedlings treated with Trichoderma harzianum. Plant Protection Science, 44, 101–107. https://doi.org/ 10.17221/23/2008-PPS Nur Azura, B. A., Yusoff, M., Tan, G. Y. A., Jegadeesh, R., Appleton, D. R., & Vikineswary, S. (2016). Streptomyces sanglieri which colonised and enhanced the growth of Elaeis guineensis Jacq. seedlings was antagonistic to Ganoderma boninense in in vitro studies. Journal of Industrial Microbiology and Biotechnology, 43, 485– 493. https://doi.org/10.1007/s10295-015-1724-4 Nurrashyeda, R., Idris, A. S., Norman, K., Kushairi, D., Azha, W. W. M., & Charles, T. (2015). GanoEF biofertilizer as preventive treatment of Ganoderma disease in oil palm—Nursery and field evaluation. Kuala Lumpur, Malaysia: MPOB International Palm Oil Congress and Exhibition (PIPOC). Oerke, E. C. (2006). Centenary review: Crop losses to pests. Journal of Agricultural Science, 144, 31–43. https://doi.org/10.1017/ S0021859605005708 Oliveira, P. M., Zannini, E., & Arendt, E. K. (2014). Cereal fungal infection, mycotoxins, and lactic acid bacteria mediated bioprotection: From crop farming to cereal products. Food Microbiology, 37, 78– 95. https://doi.org/10.1016/j.fm.2013.06.003 Ooi, P. A. C. (1987). A fortuitous biological control of Lantana in Malaysia. Tropical Pest Management, 33, 234–235. https://doi.org/ 10.1080/09670878709371158 Ooi, P. A. C. (1992). Role of parasitoids in managing diamondback moth in the Cameron Highlands, Malaysia. In N. S. Talekar (Ed.), Diamondback moth and other crucifer pests. Proceedings of the 2nd international workshop (pp. 255–262). Tainan, Taiwan: AVRDC. Ooi, P. A. C., & Lim, G. S. (1989). Introduction of exotic parasitoids to control the diamondback moth in Malaysia. Journal of Plant Protection in the Tropics, 6, 103–111. Ostry, V., Malir, F., Toman, J., & Grosse, Y. (2017). Mycotoxins as human carcinogens—The IARC Monographs classification. Mycotoxin Research, 33, 65–73. https://doi.org/10.1007/s12550016-0265-7 Palazzini, J. M., Ramirez, M. L., Torres, A. M., & Chulze, S. N. (2007). Potential biocontrol agents for Fusarium head blight and deoxynivalenol production in wheat. Crop Protection, 26, 1702– 1710. https://doi.org/10.1016/j.cropro.2007.03.004 Palazzini, J. M., Yerkovich, N., Alberione, E., Chiotta, M., & Chulze, S. N. (2017). An integrated dual strategy to control Fusarium graminearum sensu stricto by the biocontrol agent Streptomyces sp. RC 87B under field conditions. Plant Gene, 9, 13–18. https://doi.org/ 10.1016/j.plgene.2016.11.005 Paterson, R. R. M., & Lima, N. (2010). How will climate change affect mycotoxins in food? Food Research International, 43, 1902–1914. https://doi.org/10.1016/j.foodres.2009.07.010 Paterson, R. R. M., & Lima, N. (2011). Further mycotoxin effects from climate change. Food Research International, 44, 2555–2566. https://doi.org/10.1016/j.foodres.2011.05.038 Pau, C. G., Leong, C. T. S., Wong, S. K., Eng, L., Jiwan, M., Kundat, F. R., … Majid, N. M. (2012). Isolation of indigenous strains of Paecilomyces lilacinus with antagonistic activity against Meloidogyne incognita. International Journal of Agriculture and Biology, 14, 197– 203. Peng, T. Y., & Don, M. M. (2013). Application of biological-based antifungal agent for controlling the growth of wood decaying fungi of rubber wood. Advanced Materials Research, 626, 21–24. https://doi.org/10.4028/www.scientific.net/AMR.626.21 Pierre, E. M., & Idris, A. H. (2013). Studies on the predatory activities of Oecophylla smaragdina (Hymenoptera: Formicidae) on Pteroma pendula (Lepidoptera: Psychidae) in oil palm plantations in Teluk Intan, Perak (Malaysia). Asian Myrmecology, 5, 163–176. https://doi.org/10.20362/am.005017 Pitt, J. I., & Hocking, A. D. (2009). Fungi and food spoilage (3rd ed.) New York, NY: Springer. Pitt, J. I., & Miller, J. D. (2017). A concise history of mycotoxin research. Journal of Agricultural and Food Chemistry, 65, 7021– 7033. https://doi.org/10.1021/acs.jafc.6b04494 Poggio, S. L. (2005). Structure of weed communities occurring in monoculture and intercropping of field pea and barley. Agriculture, Ecosystems and Environment, 109, 48–58. https://doi.org/ 10.1016/j.agee.2005.02.019 Rahman, M. A., Kadir, J., Mahmud, T. M. M., Abdul Rahman, R., & Begum, M. M. (2007). Screening of antagonistic bacteria for biocontrol activities on Colletotrichum gloeosporioides in papaya. Asian Journal of Plant Sciences, 6, 12–20. https://doi.org/ 10.3923/ajps.2007.12.20 Rahmani, A., Jinap, S., & Soleimany, F. (2010). Validation of the procedure for the simultaneous determination of aflatoxins, ochratoxin A and zearalenone in cereals using HPLC-FLD. Food Additives and Contaminants - Part A Chemistry, Analysis, Control, Exposure and Risk Assessment, 27, 1683–1693. https://doi.org/ 10.1080/19440049.2010.514951 Ramle, M., Wahid, M. B., Norman, K., Glare, T. R., & Jackson, T. A. (2005). The incidence and use of Oryctes virus for control of rhinoceros beetle in oil palm plantations in Malaysia. Journal of Invertebrate Pathology, 89, 85–90. https://doi.org/10.1016/j.jip. 2005.02.009 Ramli, N. R., Mohamed, M. S., Seman, I. A., Zairun, M. A., & Mohamad, N. (2016). The potential of endophytic bacteria as a biological control agent for Ganoderma disease in oil palm. Sains Malaysiana, 45, 401– 409. Ramli, S., Zainal-Abidin, B. A. H., & Idris, A. B. (2011). In vivo production of Nosema bombycis spores and their efficacies against diamondback moth and beet armyworm larvae in laboratory conditions. Sains Malaysiana, 40, 311–316. Reddy, K. R. N., & Salleh, B. (2010). A preliminary study on the occurrence of Aspergillus spp. and aflatoxin B1 in imported wheat and barley in Penang, Malaysia. Mycotoxin Research, 26, 267–271. https://doi.org/10.1007/s12550-010-0065-4 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… Reddy, K. R. N., & Salleh, B. (2011). Co-occurrence of moulds and mycotoxins in corn grains used for animal feeds in Malaysia. Journal of Animal and Veterinary Advances, 10, 668–673. https://doi.org/ 10.3923/javaa.2011.668.673 Reddy, K. R. N., Farhana, N. I., & Salleh, B. (2011). Occurrence of Aspergillus spp. and aflatoxin B1 in Malaysian foods used for human consumption. Journal of Food Science, 76, T99–T104. https://doi.org/10.1111/j.1750-3841.2011.02133.x Reino, J. L., Guerrero, R. F., Hernández-Galán, R., & Collado, I. G. (2008). Secondary metabolites from species of the biocontrol agent Trichoderma. Phytochemistry Reviews, 7, 89–123. https://doi.org/ 10.1007/s11101-006-9032-2 Rychlik, M., Humpf, H.-U., Marko, D., Dänicke, S., Mally, A., Berthiller, F., … Lorenz, N. (2014). Proposal of a comprehensive definition of modified and other forms of mycotoxins including “masked” mycotoxins. Mycotoxin Research, 30, 197–205. https://doi. org/10.1007/s12550-014-0203-5 Saad, K. A., Mohamad Roff, M. N., Hallett, R. H., & Abd-Ghani, I. B. (2019). Effects of cucumber mosaic virus-infected chili plants on non-vector Bemisia tabaci (Hemiptera: Aleyrodidae). Insect Science, 26, 76–85. https://doi.org/10.1111/1744-7917.12488 Sahayaraj, K. (2014). Basic and applied aspects of biopesticides. New Delhi, India: Springer. Sajap, A. S., Rozihawati, Z., Omar, D., & Lau, W. H. (2014). Isaria fumosorosea and Metarhizium anisopliae for controlling Atteva sciodoxa (Lepidoptera: Yponomeutidae), a pest of Eurycoma longifolia. Journal of Tropical Forest Science, 26, 84–91. Sarawak Land Consolidation and Rehabilitation Authority (SALCRA). (2019). Sustainability >Pest and diseases control. Retrieved from http://www.salcra.gov.my/en/sustainable-plantation/pest-diseasescontrol.html Salim, H., Md. Rawi, C. S., Ahmad, A. H., & Al-Shami, S. A. (2015). Efficacy of insecticide and bioinsecticide ground sprays to control Metisa plana Walker (Lepidoptera: Psychidae) in oil palm plantations, Malaysia. Tropical Life Sciences Research, 26, 73–83. Salmah, M., Basri, M. W., & Idris, A. B. (2012). Effects of honey and sucrose on longevity and fecundity of Apanteles metesae (Nixon), a major parasitoid of the oil Palm Bagworm, Metisa plana (Walker). Sains Malaysiana, 41, 1543–1548. Samson, R. A., Houbraken, J., Thrane, U., Frisvad, J. C., & Andersen, B. (2010). Food and indoor fungi. CBS laboratory manual series. Utrecht, the Netherlands: CBS-KNAW-Fungal Biodiversity Centre. Samsudin, N. I. P., & Abdullah, N. (2013). A preliminary survey on the occurrence of mycotoxigenic fungi and mycotoxins contaminating red rice at consumer level in Selangor, Malaysia. Mycotoxin Research, 29, 89–96. https://doi.org/10.1007/s12550-012-0154-7 Samsudin, N. I. P., & Magan, N. (2016). Efficacy of potential biocontrol agents for control of Fusarium verticillioides and fumonisin B1 under different environmental conditions. World Mycotoxin Journal, 9, 205–213. https://doi.org/10.3920/WMJ2015.1886 Samsudin, N. I. P., Medina, A., & Magan, N. (2016). Relationship between environmental conditions, carbon utilization patterns and Niche Overlap Indices of the mycotoxigenic species Fusarium verticillioides and the biocontrol agent Clonostachys rosea. Fungal Ecology, 24, 44–52. https://doi.org/10.1016/j.funeco.2016.05.010 Samsudin, N. I. P., Rodriguez, A., Medina, A., & Magan, N. (2017). Efficacy of fungal and bacterial antagonists for controlling growth, FUM1 gene expression and fumonisin B1 production by Fusarium verticillioides on maize cobs of different ripen- 667 ing stages. International Journal of Food Microbiology, 246, 72–79. https://doi.org/10.1016/j.ijfoodmicro.2017.02.004 Saravanakumar, K., Dou, K., Lu, Z., Wang, X., Li, Y., & Chen, J. (2018). Enhanced biocontrol activity of cellulase from Trichoderma harzianum against Fusarium graminearum through activation of defense-related genes in maize. Physiological and Molecular Plant Pathology, 103, 130–136. https://doi.org/10.1016/ j.pmpp.2018.05.004 Sathyapriya, H., Sariah, M., Siti Nor Akmar, A., & Wong, M. (2012). Root colonisation of Pseudomonas aeruginosa strain UPMP3 and induction of defense-related genes in oil palm (Elaeis guineensis). Annals of Applied Biology, 160, 137–144. https://doi.org/ 10.1111/j.1744-7348.2011.00525.x Seung, C. C. F., Chyng, A. W., & Hoe, N. W. (2015). Isolation of rhizospheric and endophytic soil bacteria SPLUMS-1 and SPLUMS-2 of oil palm against Ganoderma sp. JN234427. Malaysian Journal of Microbiology, 11, 116–120. https://doi.org/10.21161/mjm.12214 Shahbazi, P., Musa, Y., Tan, G. Y. A., Avin, F. A., Teo, W. F. A., & Sabaratnam, V. (2013). Streptomyces strain P42 as a potent biological control against chili anthracnose disease caused by Colletotrichum spp. Research on Crops, 14, 935–944. Shahbazi, P., Musa, Y., Tan, G. Y. A., Avin, F. A., Teo, W. F. A., & Sabaratnam, V. (2014). In vitro and in vivo evaluation of Streptomyces suppressions against anthracnose in chili caused by Colletotrichum. Sains Malaysiana, 43, 697–705. Shariff, S., Ibrahim, N. J., Md-Zain, B. M., Idris, A. B., Suhana, Y., Roff, M. N., & Yaakop, S. (2014). Multiplex PCR in determination of opiinae parasitoids of fruit flies, Bactrocera sp., infesting star fruit and guava. Journal of Insect Science, 14, 7. https://doi.org/10.1093/jis/14.1.7 Shariffah-Muzaimah, S. A., Idris, A. S., Madihah, A. Z., Dzolkhifli, O., Kamaruzzaman, S., & Cheong, P. C. H. (2015). Isolation of actinomycetes from rhizosphere of oil palm (Elaeis guineensis Jacq.) for antagonism against Ganoderma boninense. Journal of Oil Palm Research, 27, 19–29. Shariffah-Muzaimah, S. A., Idris, A. S., Madihah, A. Z., Dzolkhifli, O., Kamaruzzaman, S., & Maizatul-Suriza, M. (2018). Characterization of Streptomyces spp. isolated from the rhizosphere of oil palm and evaluation of their ability to suppress basal stem rot disease in oil palm seedlings when applied as powder formulations in a glasshouse trial. World Journal of Microbiology and Biotechnology, 34, 15. https://doi.org/10.1007/s11274-017-2396-1 Sharmili, K., Jinap, S., & Sukor, R. (2016). Development, optimization and validation of QuEChERS based liquid chromatography tandem mass spectrometry method for determination of multimycotoxin in vegetable oil. Food Control, 70, 152–160. https://doi.org/10.1016/j.foodcont.2016.04.035 Shuib, N. S., Makahleh, A., Salhimi, S. M., & Saad, B. (2017a). Determination of aflatoxin M1 in milk and dairy products using high performance liquid chromatography-fluorescence with post column photochemical derivatization. Journal of Chromatography A, 1510, 51–56. https://doi.org/10.1016/j.chroma.2017.06.054 Shuib, N. S., Makahleh, A., Salhimi, S. M., & Saad, B. (2017b). Natural occurrence of aflatoxin M1 in fresh cow milk and human milk in Penang, Malaysia. Food Control, 73, 966–970. https://doi.org/ 10.1016/j.foodcont.2016.10.013 Siahmoshteh, F., Siciliano, I., Banani, H., Hamidi-Esfahani, Z., Razzaghi-Abyaneh, M., Gullino, M. L., & Spadaro, D. (2017). Efficacy of Bacillus subtilis and Bacillus amyloliquefaciens in the 668 BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… control of Aspergillus parasiticus growth and aflatoxins production on pistachio. International Journal of Food Microbiology, 254, 47– 53. https://doi.org/10.1016/j.ijfoodmicro.2017.05.011 Siddiquee, S., Yusuf, U. K., Hossain, K., & Jahan, S. (2009). In vitro studies on the potential Trichoderma harzianum for antagonistic properties against Ganoderma boninense. Journal of Food, Agriculture and Environment, 7, 970–976. Simberloff, D. (2012). Risks of biological control for conservation purposes. BioControl, 57, 263–276. https://doi.org/10.1007/ s10526-011-9392-4 Simmonds, F. J. (1980). Biological control of Cordia curassavica [Boraginaceae] in Malaysia. Entomophaga, 25, 363–364. https://doi.org/ 10.1007/BF02374697 Soleimany, F., Jinap, S., & Abas, F. (2012). Determination of mycotoxins in cereals by liquid chromatography tandem mass spectrometry. Food Chemistry, 130, 1055–1060. https://doi.org/ 10.1016/j.foodchem.2011.07.131 Soleimany, F., Jinap, S., Faridah, A., & Khatib, A. (2012). A UPLC-MS/MS for simultaneous determination of aflatoxins, ochratoxin A, zearalenone, DON, fumonisins, T-2 toxin and HT2 toxin, in cereals. Food Control, 25, 647–653. https://doi.org/ 10.1016/j.foodcont.2011.11.012 Sorensen, K. A., Mohankumar, S., & Thangaraj, S. R. (2016). Chapter 5: Physical, mechanical and cultural control of vegetable insects. In R. Muniappan & E. A. Heinrichs, (Eds.), Integrated pest management of tropical vegetable crops (pp. 131–148). Dordrecht, the Netherlands: Springer. https://doi.org/10.1007/978-94-024-0924-6_5 Strasser, H., Vey, A., & Butt, T. M. (2010). Are there any risks in using entomopathogenic fungi for pest control, with particular reference to the bioactive metabolites of Metarhizium, Tolypocladium and Beauveria species? Biocontrol Science and Technology, 10, 717–735. https://doi.org/10.1080/09583150020011690 Suhaida, S., & NurAinIzzati, M. Z. (2013). The efficacy of Trichoderma harzianum T73s as a biocontrol agent of Fusarium ear rot disease of maize. International Journal of Agriculture and Biology, 15, 1175– 1180. Suhanna, A., Nor Hanis Aifaa, Y., & Shazalwardi, S. (2013). Trichoderma sp. as a biological control agent in the postharvest treatment of mango stem-end rot. Acta Horticulturae, 1012, 775–782. https://doi.org/10.17660/ActaHortic.2013.1012.105 Sulaiman, M. R., Chye, F. Y., Hamid, A., & Yatim, A. M. (2007). The occurrence of aflatoxins in raw shelled peanut samples from three districts of Perak, Malaysia. Electronic Journal of Environmental, Agricultural and Food Chemistry, 6, 2045–2052. Sule, H., Muhamad, R., Omar, D., & Hee, A. K. W. (2014). Parasitism rate, host stage preference and functional response of Tamarixia radiata on Diaphorina citri. International Journal of Agriculture and Biology, 16, 783–788. Sultan, Y., & Magan, N. (2011). Impact of a Streptomyces (AS1) strain and its metabolites on control of Aspergillus flavus and aflatoxin B1 contamination in vitro and in stored peanuts. Biocontrol Science and Technology, 21, 1437–1455. https://doi.org/ 10.1080/09583157.2011.632078 Sundram, S., Abdullah, F., Ahmad, Z. A. M., & Yusuf, U. K. (2008). Efficacy of single and mixed treatments of Trichoderma harzianum as biocontrol agents of Ganoderma basal stem rot in oil palm. Journal of Oil Palm Research, 20, 470–483. Tan, C. J., How, K. C., Loh-Mia, P. P., Ismet, A., Getha, K., Seki, T., & Vikineswary, S. (2002). Bioactivity of selected actinomycetes against Ganoderma boninense. Asia-Pacific Journal of Molecular Biology and Biotechnology, 10, 119–125. Teoh, Y. P., Don, M. M., & Ujang, S. (2012). Nutrient improvement using statistical optimization for growth of Schizophyllum commune, and its antifungal activity against wood degrading fungi of rubber wood. Biotechnology Progress, 28, 232–241. https://doi.org/10.1002/btpr.714 Teoh, Y. P., Don, M. M., & Ujang, S. (2017). Production of biomass by Schizophyllum commune and its antifungal activity towards rubber wood-degrading fungi. Sains Malaysiana, 46, 123–128. https://doi. org/10.17576/jsm-2017-4601-16 Ting, A. S. Y., Meon, S., Kadir, J., Radu, S., & Singh, G. (2010). Induction of host defense enzymes by the endophytic bacterium Serratia marcescens in banana plantlets. International Journal of Pest Management, 56, 183–188. https://doi.org/10.1080/09670870903324198 Ting, A. S. Y., Sariah, M., Kadir, J., & Gurmit, S. (2009). Field evaluation of non-pathogenic Fusarium oxysporum isolates UPM31P1 and UPM39B3 for the control of Fusarium wilt in ‘pisang berangan’ (Musa, AAA). Acta Horticulturae, 828, 139–144. https://doi.org/ 10.17660/ActaHortic.2009.828.13 Toh, S. C., Samuel, L., & Awang, A. S. A. H. (2016). Screening for antifungal-producing bacteria from Piper nigrum plant against Phytophthora capsici. International Food Research Journal, 23, 2616– 2622. Tran-Dinh, N., Pitt, J. I., & Markwell, P. J. (2018). Use of microsatellite markers to assess the competitive ability of nontoxigenic Aspergillus flavus strains in studies on biocontrol of aflatoxins in maize in Thailand. Biocontrol Science and Technology, 28, 215–225. https://doi.org/10.1080/09583157.2018.1436694 van Lenteren, J. C., Bolckmans, K., Köhl, J., Ravensberg, W. J., & Urbaneja, A. (2018). Biological control using invertebrates and microorganisms: Plenty of new opportunities. BioControl, 63, 39– 59. https://doi.org/10.1007/s10526-017-9801-4 Veenstra, A., Rafudeen, M. S., & Murray, S. L. (2019). Trichoderma asperellum isolated from African maize seed directly inhibits Fusarium verticillioides growth in vitro. European Journal of Plant Pathology, 153, 279–283. https://doi.org/10.1007/s10658-018-1530-8 Vincent, C., Hallman, G., Panneton, B., & Fleurat-Lessard, F. (2003). Management of agricultural insects with physical control methods. Annual Review of Entomology, 48, 261–281. https://doi.org/ 10.1146/annurev.ento.48.091801.112639 Volpon, L., Besson, F., & Lancelin, J. M. (2000). NMR structure of antibiotics plipastatins A and B from Bacillus subtilis inhibitors of phospholipase A2 . FEBS Letters, 485, 76–80. https://doi.org/ 10.1016/S0014-5793(00)02182-7 Wannop, C. C. (1961). The histopathology of Turkey “X” Disease in Great Britain. Avian Diseases, 5, 371–381. https://doi.org/ 10.2307/1587768 Way, M. J., & Khoo, K. C. (1991). Colony dispersion and nesting habits of the ants, Dolichoderus thoracicus and Oecophylla smaragdina (Hymenoptera: Formicidae), in relation to their success as biological control agents on cocoa. Bulletin of Entomological Research, 81, 341–350. https://doi.org/10.1017/S0007485300033629 Williams, P. P. (1989). Effects of T-2 mycotoxin on gastrointestinal tissues: A review of in vivo and in vitro models. Archives of Environmental Contamination and Toxicology, 18, 374–387. https://doi.org/10.1007/BF01062362 Wyckhuys, K. A. G., Lu, Y., Morales, H., Vazquez, L. L., Legaspi, J. C., Eliopoulos, P. A., & Hernandez, L. M. (2013). Current status BIOLOGICAL CONTROL OF PHYTOPATHOGENIC ORGANISMS AND MYCOTOXIGENIC FUNGI… and potential of conservation biological control for agriculture in the developing world. Biological Control, 65, 152–167. https://doi.org/ 10.1016/j.biocontrol.2012.11.010 Yaakop, S., Shariff, S., Ibrahim, N. J., Md-Zain, B. M., Yusof, S., & Mohamad Jani, N. (2015). Dual-target detection using simultaneous amplification of PCR in clarifying interaction between Opiinae species (Hymenoptera: Braconidae) associated with Bactrocera spp. (Diptera: Tephritidae) infesting several crops. Arthropod-Plant Interactions, 9, 121–131. https://doi.org/10.1007/s11829-015-9355-2 Zabedah, M., Aini, Z., & Hussan, A. K. (2010). Effective control of army worm (Spodoptera litura) for starfruit under a netted structure with nuclear polyhedrosis virus (NPV). Acta Horticulturae, 873, 321–324. https://doi.org/10.17660/ActaHortic.2010.873.37 Zacky, F. A., & Ting, A. S. Y. (2013). Investigating the bioactivity of cells and cell-free extracts of Streptomyces griseus towards Fusarium oxysporum f. sp. cubense race 4. Biological Control, 66, 204–208. https://doi.org/10.1016/j.biocontrol.2013.06.001 Zacky, F. A., & Ting, A. S. Y. (2015). Biocontrol of Fusarium oxysporum f. sp. cubense tropical race 4 by formulated cells and cell-free 669 extracts of Streptomyces griseus in sterile soil environment. Biocontrol Science and Technology, 25, 685–696. https://doi.org/10.1080/ 09583157.2015.1007921 Zalom, F. G. (2010). Chapter 8: Pesticide use practices in integrated pest management. In R. Krieger (Ed.), Hayes’ handbook of pesticide toxicology. (3rd ed., pp. 303–313). Cambridge, MA: Academic Press. https://doi.org/10.1016/B978-0-12-374367-1.00008-2 How to cite this article: Yazid SNE, Jinap S, Ismail SI, Magan N, Samsudin NIP. Phytopathogenic organisms and mycotoxigenic fungi: Why do we control one and neglect the other? A biological control perspective in Malaysia. Compr Rev Food Sci Food Saf. 2020;19:643– 669. https://doi.org/10.1111/1541-4337.12541

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Phytopathogenic organisms and mycotoxigenic fungi: Why do we control one and neglect the other? A biological control perspective in Malaysia

Phytopathogenic organisms and mycotoxigenic fungi: Why do we control one and neglect the other? A biological control perspective in Malaysia

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