Subasinghe and Hettiarachchige /Journal of Tropical Forestry and Environment Vol. 11 No. 02 (2021) 1-9 1 Feature Article Endophytic Fungal Species in Tropical Trees: A Review S.M.C.U.P. Subasinghe and R.P. Hettiarachchige Centre for Forestry and Environment Department of Forestry and Environmental Science University of Sri Jayewardenepura, Sri Lanka Abstract Pathogenic fungi are common in forest ecosystems which cause diseases and sometimes death of plants, while some fungi live inside trees harmlessly without causing issues. Sometimes, plants benefit from the presence of those endophytic fungi, such as gaining resistance to environmental stresses, protection from harmful pathogens etc. Numerous studies have been conducted on such relationships between endophytic fungi and short-term agricultural crops. However, such studies are rare in the literature on tropical tree species which bear timber and non-timber values. This study illustrates the studies conducted on endophytic fungi in tropical trees and explores the potential use of such fungi for obtaining benefits. Keywords: Endophytic fungi, interactions, tropical trees, abundance 1. Introduction Fungi are essential components in every ecosystem, intimately associated with crucial processes like the decomposition, recycling, and transportation of nutrients in different environments. Endophytic fungi are a group that colonize living, internal tissues of plants without causing immediate or overt adverse effects (Bacon and White, 2000). However, they may be pathogenic during host senescence (Rodriguez and Redman, 2008). They are a taxonomically and ecologically heterogeneous group of organisms; mainly belonging to Ascomycota, coelomycetes, and hyphomycetes (Rajagopal et al., 2012). Most endophytes are horizontally transmitted to their host plants through airborne spores. In contrast, some endophytes can vertically transmit to the next plant generations via seeds (Hartley and Gange, 2009). Though endophytic fungi were known for over one hundred years, they did not receive much attention until the recent recognition of their pharmaceutical and ecological significance (Gunatilaka, 2006). Various relationships exist between fungal endophytes and their host plants, ranging from mutualistic or symbiotic to slightly pathogenic (Schulz and Boyle, 2005). Symbiotic associations between fungi and photosynthetic organisms are both ancient and ubiquitous. The relationship of endophytes with hosts is described in different terms of host specificity, host recurrence, host selectivity etc. Host specificity is the relationship where a fungus is restricted to a single plant species or a group of related species and not found in other unrelated plants in the same habitat (Ananda and Sridhar, 2002). Host recurrence is the predominant occurrence of endophytic fungus on a particular host or a range of plant hosts. Still, the fungus can also occur infrequently on other host plants in the same habitat. If a single fungal endophyte form relationship with two related plant hosts, but demonstrate more likely to one host, it is described as host selectivity (Cohen, 2006). Endophytic fungi commonly demonstrate host specificity at the plant species level, but environmental conditions can influence this specificity (Cohen, 2004). At the same time, differences in endophytic fungal assemblage have been found in different tissues of the same plant species and even in other tissues of a single plant. This phenomenon reflects the tissues specificity (Ganley and Newcome, 2006). 2 2. Positive and Negative Interactions between Endophytic Fungi and Trees Beneficial interactions between fungi and relevant hosts could be presented in different aspects. Endophytic fungi produce many substances inside the host plants that have potential use in modern medicine, agriculture and industry such as antibiotics, anticancer compounds (Michell et al., 2008). Some endophytic fungi could produce different plant hormones to enhance the growth of their host plants (Waqas et al., 2012). For example, the growth of wheat (Triticumaestivum L.) could be enhanced by Azospirillum sp. under drought stresses (Dingle and McGee, 2003). Some produce different bioactive compounds, such as alkaloids, diterpenes, flavonoids, and isoflavonoids to increase their host plants' resistance to biotic and abiotic stresses (Rodriguez et al., 2009). Certain endophytic fungal species could also promote the accumulation of secondary metabolites (including important medicinal components or drugs) originally produced by plants. According to the references, these metabolites may be produced by both host plants and/or endophytic fungi (Shwab and Keller, 2008). Endophytic fungi can protect their host plants from pathogens and pests (Arnold et al., 2003). The foliar endophytes can reduce herbivory by producing alkaloids that are toxic to insects and vertebrates (Schardl, 2001). The endophytic fungus Acremonium coenophialum exhibited insecticidal activity against aphids (Rhopalosiphum padi, Schizaphis graminum) and milkweed bugs. Cladosporium herbarum, A. alternata, Rhodotorula rubra, Epicoccum nigrum, Cryptococcus sp., Penicillium sp. and Fusarium graminearum act as protectants to plants from herbivores (Larran et al., 2002). Senthilkumar et al., (2014) reported that phytochemicals such as dodecanoic acid, ethyl ester, phthalic acid, octyl 2-pentyl ester isolated from Phomopsis sp. can be used as an insecticide. Further, Cladosporium oxysporum showed insecticidal activity against Aphis fabae (Bensaci et al., 2015). Extracts of Emericella nidulans, A. oryzae, A. tamarii and A. versicolor showed the insecticidal ability on Spodoptera litura larvae with the maximum activity of A. versicolor (Abraham et al., 2015). Endophytes can actively or passively promote plant growth through different mechanisms. Endophytic metabolites provide a variety of fitness to host plants enhanced by increasing plant resistance to biotic and abiotic stresses, as well as enhancing plant growth. Many endophytes are capable of solubilization of phosphate, enhance uptake of phosphorus, nitrogen fixation and plant hormones such as auxin, abscisic, ethylene, gibberellins, and indole acetic acid (e.g., Fusarium tricinctum alternate), which are important for plant growth regulation (Khan et al., 2015; Sandhiya et al., 2005). Fungal endophytes the form mycorrhizal associations assists to distribute nutrients within the surrounding plants (Franklin et al., 2014). The mechanisms of certain endophytic fungi induced resistance related to the nutritional status of the host, and thereby to increasing the fitness of plants by enhancing their tolerance to abiotic stress (Lu et al., 2000). Symbiotic fungi play an important role in the ecological community by decreasing the spoilage of land and water caused by excessive toxic organic insecticide, environmental degradation, industrial sewage, poisonous gases and loss of biodiversity (Guo et al., 2008). Due to the potential of microorganisms for bioaccumulation of heavy metals and other pollutants from the environment, it can enhance and increase plant growth (Ma et al., 2011). Due to that, endophytes may also play a direct or indirect role in the phytoremediation process and degradation of environmental toxins, indirectly through enhancing plant growth having the ability of phytoremediation and this accelerate phytoremediation process, or directly through degradation and/or accumulating pollutants (Oses et al., 2008). For example, Penicillium funiculosum acts against the copper stress and enhances the plant growth. Therefore, it can be used in bioremediation of pollution in the cultivated area by stress mediating endophytes (Khan and Lee, 2013). Subasinghe and Hettiarachchige /Journal of Tropical Forestry and Environment Vol. 11 No. 02 (2021) 1-9 3 3. Potential for the Use of Positive Impacts Endophytes generated an interest among the scientists of microbial chemistry due to their potential to contribute to the discovery of new bioactive compounds. It has been suggested that the close biological association between endophytes and their plant hosts results in the production of a large number and diversity of biologically active molecules compared to epiphytes or soil-related microbes (Strobel, 2003). Moreover, the symbiotic nature of this relationship indicates that endophytic bioactive compounds are less toxic to the animal cells, as these chemicals do not kill the eukaryotic host system. This is particularly important to the researchers in the medical field as potential drugs may not adversely affect human cells (Alvin et al., 2014). 4. The Abundance of Fungi in Tropical Trees Endophytic fungi have been found in all plant species examined to date, including algae, mosses, conifers, angiosperms, palms and a variety of dicotyledonous shrubs and trees. Studies conducted on the presence of endophytic fungi in commercially valuable tropical timber trees such as Dipterocarpus tuberculatus, Guarea guidonia, Hevea brasiliensis, Khaya anthotheca, and Tectona grandis. In addition, similar studies were conducted on commercially valuable non-timber species in food and confectionary (Mangifera indica, Musa acuminate, Theobroma cacao), medicine, (Azadirachta Indica, Camptotheca acuminate, Crocus sativus, Ginkgo biloba, Mesua ferrea, Samanea saman, Terminalia sp., Vitex negundo) and enzyme production (Calophyllum inophyllum). Several authors have suggested that endophytes are especially diverse in tropical forests (Table 01). Table 01. Fungal species identified in tropical tree species Tree species No. of fungal species Dominant species Location and abundance Reference B L S T F R Aegle marmelos 10 Alternaria sp. Aspergillus sp. Bipolaris crotonis Cladosporium sp. Colletotrichum sp. Curvularia sp. Daldinia sp. Fusarium sp. Nigrospora sp. Phomopsis sp. Schizophyllum sp. Trichoderma sp. Xylaria sp. L H L H H L H L L L L L L - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Gond et al., 2007 Artocarpus heterophyllus 1 Colletotrichum sp. - H - - - - Sunkar et al., 2017 Azadirachta Indica 29 Alternaria sp. Aspergillus sp. Colletotrichum sp. Diaporthe sp. Fusarium sp. Penicillium sp. Trichoderma sp. Xylaria sp. - L - H L - H - H L L H L - H - - L - H L - H H - - - - - - - - H L L - L L H - H L - - L L H - Chutulo et al., 2018 Calophyllum inophyllum 8 Cladosporium sp. Discosia sp. - - H H - - - - - - - - Hedge et al., 2011 4 Fusarium sp. Isaria sp. Penicillium sp. Pestalotiopsis sp. Xylaria sp. - - - - - H H H L L - - - - - - - - - - - - - - - - - - - - Camptotheca acuminata >5 Alternaria sp. Fusarium sp. Paecilomyces sp. Phomopsis sp. Rhizoctonia sp. H - - - - - - - - - - - - - - H - - H T - L L - - H - - H - Lin et al., 2007 Coffea arabica 1 Glomerella cingulata - H - H - - Sette et al., 2006 Coffea robusta 2 Glomerella cingulate Phomopsis sp. - H - - - - L - - - - - Sette et al., 2006 Crocus sativus 12 Alternaria sp. Fusarium sp. Penicillium sp. Phytophthora sp. Rhizoctonia sp. Trichoderma sp. - - - - - - - - - - - - L H L H H L - - - - - - - - - - - - - - - - - - Raj et al., 2013 Dillenia indica 25 Alternaria sp. Aspergillus sp. Bipolaris crotonis Cladosporium sp. Colletotrichum sp. Daldinia sp. Fusarium sp. Nigrospora sp. Phomopsis sp. Schizophyllum sp. Trichoderma sp. Xylaria sp. - - - - - - - - - - - - L H H H H L L L H L L L - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - Prasher and Kumar, 2021 Diospyros celebica 7 Aspergillus sp. Fusarium sp. Gliocladium sp. Penicillium sp. Phytophthora sp. Trichoderma sp. L L L L L L H L L - - - L L - H - L - - - - - - - - - L - - L L - L - L Mukrimin et al., 2021 Diospyros crassiflora 7 Clypeosphaeria sp. Colletotrichum sp. Penicillium sp. Phomopsis sp. Phyllosticta sp. Xylaria sp. - - - - - - L L - H H H - - - - - - - - - - - - L - L - - - - - - - - - Douanla-Meli and Langer, 2012 Dipterocarpus tuberculatus 9 Daldinia sp. Fusarium sp. Penicillium sp. Pestalotiopsis sp. Phomopsis sp. Phyllosticta sp. Xylaria sp. - - - - - - - L L L L L L L - - - - - - - - - - - - - - - - - - - - - - - - - - - - Sutjaritvorakul et al., 2011 Subasinghe and Hettiarachchige /Journal of Tropical Forestry and Environment Vol. 11 No. 02 (2021) 1-9 5 Garcinia indica 6 Aspergillus sp. Fusarium sp. Trichoderma sp. L H H - - - - - - L H H - - - - - - Tejesvi et al., 2006 Ginkgo biloba 1 Muscodor albus - - H - - - Banerjee et al., 2010 Guarea guidonia 14 Colhotrichum sp. Phomopsis sp. Rhizoctonia sp. Trichoderma sp. Xylaria sp. - - - - - - - - - - - - - - - H L H L L - - - - - - - - - - Gamboa and Bayman, 2001 Hevea brasiliensis 58 Colletotrichum sp. Fusarium sp. Penicillium sp. Trichoderma sp. Xylaria sp. - - - - - H L L L L H L L H H - - - - - - - - - - - - - - - Gazis and Chaverri, 2010 Khaya anthotheca 10 Colletotrichum sp. Fusarium sp. Phomopsis sp. Xylaria sp. - - - - H L L H - - - - - - - - - - - - - - - - Linnakoski et al., 2012 Mangifera indica 7 Aspergillus flavus Aspergillus niger Candida sp. Cladosporium sp. Penicillium sp. Moniliella sp. Trichoderma sp. H L H H L - L H L H H L H L - - - - - - - - - - - - - - - - - - - - - H L H H L - L Nayak, 2015 Mesua ferrea 1 Phomopsis sp. - L L - - - Jayanthi et al., 2011 Musa acuminata 6 Colletotrichum sp. Cordana musae Dactylaria sp. Deightoniella sp. Guignardia sp. Pyriculariopsis sp. - - - - - - H H H H H H - - - - - - - - - - - - - - - - - - - - - - - - Photita et al., 2001 Pinus attenuate 5 Leploslrotnn sp. Naemacyclus sp. - - H H - - - - - - - - Carroll and Carroll, 1978 Pinus contoria 5 Cladosporium sp. Leploslrotnn sp. Naemacyclus sp. - - - H H H - - - - - - - - - - - - Carroll and Carroll, 1978 Pinus monticola 2 Leploslrotnn sp. - H - - - - Carroll and Carroll, 1978 Samanea saman 4 Colletotrichum sp. Nigrospora sp. Penicillium sp. Phomopsis sp. - - - - H L L L - - - - - - - - - - - - - - - - Chareprasert et al, 2006 Tectona grandis 7 Alternaria sp. Colletotrichum sp. Fusarium sp. Nigrospora sp. Penicillium sp. Phomopsis sp. Schizophyllum sp. - - - - - - - H H L L L L L - - - - - - - - - - - - - - - - - - - - - - - - - - - - Chareprasert et al., 2006 6 Terminalia arjuna 6 Colletotrichum sp. Phomopsis sp. Trichoderma sp. Tubercularia sp. - H H H L H H - - - - - - - - H - - - - - - - - Kouipou and Boyom, 2019 Terminalia catappa 7 Colletotrichum sp. Diaporthe sp. Trichoderma sp. Xylaria sp. - - L L H - - - - L - - - - - - - - - - - - - - Kouipou and d Boyom, 2019 Terminalia chebula 4 Colletotrichum sp. Penicillium sp. Xylaria sp. - - - L - - - H L - L L - - - - - - Kouipou and Boyom, 2019 Terminalia crenulate 4 Colletotrichum sp. - L - - - - Kouipou and Boyom, 2019 Terminalia mantaly 6 Colletotrichum sp. Diaporthe sp. Fusarium sp. - L - L L - - L - - - - - - - - - - Kouipou and Boyom, 2019 Theoborma cacao 3 Colletotrichum sp. Fusarium sp. Xylaria sp. - - - H L L - - - - - - - - - - - - Arnold et al., 2003 Vitex negundo 9 Alternaria sp. Aspergillus flavus Colletotrichum sp. Discosia sp. Fusarium solani Lasiodiplodia sp. - - - L L - - L - L L - - - - - - - H L L L L - - - - - - - - - L - - - Sunayana et al., 2014 Location of the tree: B=bark, F=fruit, L=leaf, R=root, S=stem, T=twig Abundance of fungal species: H=high, L=low 4. 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