Extracellular hydrolytic enzymes produced by entomopathogenic fungi--role in an infection process

Entomopathogenic fungi have a great potential in biological control of insect pest population. Fungal pathogens are promising source of insecticides and notable alterative to chemical pesticides. These fungi possess a unique mechanism of insects paralysis. As natural enemies of insects they attack direct host cuticle via a combination of mechanical pressure and cuticle-degrading enzymes. Entomopathogenic fungi produce several proteo-, chitino- and lipolytic enzymes, which are accepted as key factors in insect mycosis. The role of extracellular enzymes in pathogenesis is still not well understood. Profound understanding the mechanisms of insect paralysis by entomopathogenic fungi will help in the production of safer for environment and more efficiency mycoinsecticides.     

Role of entomopathogenic fungi in the control of Tetranychus evansi and Tetranychus urticae (Acari: Tetranychidae), pests of horticultural crops

The spider mites Tetranychus urticae Koch and Tetranychus evansi Baker and Pritchard are important pests of horticultural crops. They are infected by entomopathogenic fungi naturally or experimentally. Fungal pathogens known to cause high infection in spider mite populations belong to the order Entomophthorales and include Neozygites spp. Studies are being carried out to develop some of these fungi as mycoacaricides, as stand-alone control measures in an inundative strategy to replace the synthetic acaricides currently in use or as a component of integrated mite management. Although emphasis has been put on inundative releases, entomopathogenic fungi can also be used in classical, conservation and augmentative biological control. Permanent establishment of an exotic agent in a new area of introduction may be possible in the case of spider mites. Conservation biological control can be achieved by identifying strategies to promote any natural enemies already present within crop ecosystems, based on a thorough understanding of their biology, ecology and behaviour. Further research should focus on development of efficient mass production systems, formulation, and delivery systems of fungal pathogens.     

虫生真菌研究与应用的发展与机遇 ——2012年虫生真菌专刊序言

我国从事虫生真菌基础及应用研究的人员众多,各方面均取得了重要的研究成果。2012年“虫生真菌专刊”收集了国内关于虫生真菌资源、基因克隆及活性成分代谢等方面的最新研究进展。进一步的基础研究,应加强致病基因功能和真菌-昆虫分子相互作用等研究,以提高真菌杀虫剂或药用虫生真菌的应用效率。     

Entomopathogenic Fusarium species: a review of their potential for the biological control of insects,implications and prospects

The genus Fusarium is noted for including important plant pathogens and mycotoxin producers. Furthermore, many Fusarium lineages have been reported to be efficient in controlling insects and to exhibit promising characteristics for agricultural pest control such as causing high mortality rates and having fast action and abundant sporulation. In this review we present a survey of peer-reviewed papers published from 2000 to 2019, demonstrating the widespread pathogenicity of Fusarium to insects. This survey was made using search strings in a number of databases. We list the major complexes and species of Fusarium reported as entomopathogenic and highlight the features of interest for insect control as well as the advantages and implications of this practice. Out of a total of forty papers, at least 30 species and 273 isolates of Fusarium were reported as pathogenic to at least one species of insect. Ten complexes of Fusarium species harbor entomopathogenic fungi, of which F. incarnatum-equiseti, F. fujikuroi, F. oxysporum and F. solani (= Neocosmospora solani) species complexes represented the most abundant number of entomopathogenic strains. The entomopathogenic interactions of these fungi have received greater attention in recent years, but much remains to be explored, especially regarding the specificity of these fungi for the host insect, the production of undesirable secondary metabolites, and side-effect and safety tests organisms not targets. Detailed investigations in this regard will be crucial if we are to fully exploit the potential of Fusarium for controlling insect pests.     

Distribution,occurrence and natural invertebrate hosts of indigenous entomopathogenic fungi of Central India

We report here that, during periodical surveys of insects inhabiting diverse habitats for the collection of entomopathogenic fungi; a large number of isolates were recovered belonging to seven species, from various regions of Madhya Pradesh and Chhattisgarh forest areas and agricultural fields. The most common entomopathogenic fungi such as Beauveria bassiana, Nomuraea rileyi, Paecilomyces farinosus and Paecilomyces fumosoroseus were found to infect various insect hosts species naturally viz. Hyblaea puera, Eutectona machaeralis, Diachrysia orichalcea, Spodoptera litura, and few new insect hosts of these fungal pathogens among Indian insect population were collected for the first time from Central India, such as beetles of Agrilus species, hairy caterpillars of Lymantria species. The isolation, identification, maintenance and pathogenicity assay of these isolates was performed prior to deposition in culture collection center.     

Insect pathology and fungal endophytes

Fungi that occur inside asymptomatic plant tissues are known as fungal endophytes. Different genera of fungal entomopathogens have been reported as naturally occurring fungal endophytes, and it has been shown that it is possible to inoculate plants with fungal entomopathogens, making them endophytic. Their mode of action against insects appears to be due to antibiosis or feeding deterrence. Research aimed at understanding the fungal ecology of entomopathogenic fungi, and their role as fungal endophytes, could lead to a new paradigm on how to successfully use these organisms in biological control programs.     

Interactions Between Entomopathogenic Fungi and Other Natural Enemies: Implications for Biological Control

Pathogens and arthropod natural enemies may contribute to the suppression of insect pest populations either as individual species or as species complexes. However, because natural enemies of insects have evolved and function in a multitrophic context it is important to assess interactions within complexes of natural enemies if they are to be exploited effectively in pest management. Natural enemies can interact either synergistically/additively (e.g. enhanced transmission and dispersal of insect pathogens) or antagonistically (e.g. parasitism/infection, predation and competition). In this paper, studies assessing the potential interactions between insect and fungal natural enemies are reviewed. In general, these studies indicate the positive nature of the interactions between arthropod natural enemies and fungal pathogens with respect to the control of insect populations. More work is required to investigate further the many ways in which the natural enemy community interacts in the agroecosystem     

Fungal pathogens as classical biological control agents against arthropods

Fungal entomopathogens have been used more frequently than other types of pathogens for classical biological control. Among 136 programs using different groups of arthropod pathogens, 49.3% have introduced fungal pathogens (including both the traditional fungi and microsporidia). The most commonly introduced species was Metarhizium anisopliae (Metschnikoff) Sorokin, with 13 introductions, followed by Entomophaga maimaiga Humber, Shimazu & Soper, which was released seven times. The majority of introduction programs have focused on controlling invasive species of insects or mites (70.7%) rather than on native hosts (29.4%). Almost half of the introductions of traditional fungi targeted species of Hemiptera and 75% of the microsporidia introduced have been introduced against lepidopteran species. The United States was the country where most introductions of fungi took place (n = 24). From 1993 to 2007, no arthropod pathogens were released in the US due to the rigorous regulatory structure, but in 2008 two species of microsporidia were introduced against the gypsy moth, Lymantria dispar (L.). Establishment of entomopathogenic fungi in programs introducing traditional fungi was 32.1% and establishment was 50.0% for programs introducing microsporidia. In some programs, releases have resulted in permanent successful establishment with no non-target effects. In summary, classical biological control using fungal entomopathogens can provide a successful and environmentally friendly avenue for controlling arthropod pests, including the increasing numbers of invasive non-native species.     

Extracellular peptidases of insect-associated fungi and their possible use in biological control programs and as pathogenicity markers

This review deals with characteristics of peptidases of fungi whose life cycles are associated with insects to varying degrees. The review examines the characteristic features of the extracellular peptidases of entomopathogenic fungi, the dependence of the specificity of these peptidases on the ecological characteristics of the fungi, and the role of peptidases in the development of the pathogenesis. Data on the properties and physiological role of hydrolytic enzymes of symbiotic fungi in “fungal gardens” are also considered in detail. For the development of representations about mechanisms of control over populations of insect pests, special attention is given to analysis of possibilities of genetic engineering for the creation of entomopathogens with enhanced virulence. Clarification of the role of fungi and their secreted enzymes and careful environmental studies are still required to explain their significance in the composition of the biota and to ensure widespread adoption of these organisms as effective biological control agents. The systematization and comparative analysis of the existing data on extracellular peptidases of insect-associated fungi will help in the planning of further work and the search for markers of pathogenesis and symbiosis.     

Pathogenesis-related genes of entomopathogenic fungi

All living things on Earth experience various diseases such as those caused by viruses, bacteria, and fungi. Insects are no exception to this rule, and fungi that cause disease in insects are called entomopathogenic fungi. These fungi have been developed as microbial insecticides and are used to control various pests. Generally, the mode of action of entomopathogenic fungi is divided into the attachment of conidia, germination, penetration, growth, and generation of secondary infectious conidia. In each of these steps, that entomopathogenic fungi use genes in a complex manner (specific or diverse) has been shown by gene knock-out and RNA-sequencing analysis. In this review, the information mechanism of entomopathogenic fungi was divided into six steps: (1) attachment of conidia to host, (2) germination and appressorium, (3) penetration, (4) fungal growth in hemolymph, (5) conidia production on host, and (6) transmission and dispersal. The strategy used by the fungi in each step was described at the genetic level. In addition, an approach for studying the mode of action of the fungi is presented.     

Mycobiota of the date palm phylloplane: description and interactions

We have analysed the mycobiota of date palm (Phoenix dactylifera, L.) leaves using several techniques. Profusely sporulating fungi (Penicillium spp. and Cladosporium spp.) developed when plating leaf fragments and leaf washings. Fusarium oxysporum, was particularly abundant in leaves infested with the red scale insect Phoenicococcus marlatti Cockerell, 1899, but an undescribed Lecanicillium cf. psalliotae was also found. Dual and overlay cultures showed interactions between palm pathogens, entomopathogenic and saprotrophic fungi. The most significant was the strong inhibition of the palm pathogen Penicillium vermoesenii caused by the entomopathogen Beauveria bassiana. No symptoms developed when F. oxysporum isolated from scale insects or the entomopathogens B. bassiana or Lecanicillium dimorphum were wound-inoculated on P. dactylifera petioles.     

丛枝菌根真菌与植食性昆虫的相互作用

丛枝菌根(arbuscular mycorrhizal AM)真菌与昆虫均是陆地生态系统中的重要组分,同植物关系密切,对植物的影响和作用是巨大的。生态系统中则以AM真菌-植物-昆虫互作体系参预食物网与生态过程。早在20世纪80年代,人们已开始研究AM真菌对昆虫的影响。进入21世纪人们越来越重视AM真菌与昆虫的相互作用。总结了AM真菌对昆虫取食偏好、生长、繁殖和对植物危害等方面的影响、以及昆虫对AM真菌侵染、扩展和产孢的影响;分析了植物营养状况、昆虫性别、昆虫龄期和AM真菌种类等对AM真菌与昆虫相互作用的影响特点;探讨了AM真菌与昆虫相互作用的机制;展望了利用AM真菌抑制植食性害虫、及促进天敌昆虫和部分传粉昆虫作用的可能性,旨在丰富菌根学研究内容、促进AM真菌与昆虫互作领域的深入研究、为探索生物防控农林业害虫的新途径提供依据。     

医学昆虫的病原真菌*

昆虫病原真菌在自然环境中对害虫数量起着抑制作用,在害虫的微生物防治中也占有独特的地位。长期以来,在应用昆虫病原真菌防治人畜害虫方面研究进展较为缓慢,六十年代以来才有较大的发展。本文介绍了近十年的发展概况,并讨论了影响这个研究领域迅速发展的原因及病原真菌在生物防治中应用的潜力。认为尽管至今对这类真菌了解还较少,但通过今后加强有关方面的研究工作,其应用前景是广阔的。     

Fungus Exposed Solenopsis invicta Ants Benefit from Grooming

We investigated aspects of resistance to entomopathogenic fungi in the social insect Solenopsis invicta, the red imported fire ant (RIFA). RIFA reared individually were significantly more susceptible to the entomopathogenic fungi Metarhizium anisopliae var. anisopliae M09 than reared in groups. Fungus exposed ants performed more self-grooming behavior when isolated as individuals and received more allo-grooming when accompanied with four healthy nestmates. Using fluorescence microscopy, we counted the number of fluorescein isothiocyanate (FITC)-labeled conidia on the cuticle of fungus exposed ants reared individually or as groups. The number of conidia on the surface of grouped ants decreased more rapidly than on isolated individuals. Allo-grooming behavior appears to be important in removing the conidia on the surface of RIFA. Individuals help fungus exposed ants by performing intensive grooming behaviors, which either risk infecting themselves or get them immunized as social immunity. We show evidence that contacting with fungus exposed ants would decrease susceptibility of nestmates to the fungus. All these results indicate that RIFA benefit from grooming behavior to fight against the fungal pathogens. Future advances in biological control of RIFA with entomopathogenic fungi are also discussed.     

Agroforestry coffee soils increase the insect‐suppressive potential offered by entomopathogenic fungi over full‐sun soils: A case proposing a “bait survival technique”

Entomopathogenic fungi are important natural enemies of insects. However, there is little information on the insect‐suppressive potential of these fungi and possible effects of farming management on this. Meanwhile, changes in natural landscapes due to agricultural intensification have caused considerable biodiversity loss and consequent decay of ecosystem services. However, the adoption of practices such as agroforestry in agroecosystems can foster abiotic and biotic conditions that conserve biodiversity, consequently restoring the provision of ecosystems services. Here, we assessed the effect of management systems (agroforestry or full‐sun) on the pest‐suppressive potential of entomopathogenic fungi in Brazilian coffee plantations. We used the insect bait method coupled with survival analyses to assess the speed of kill by entomopathogenic fungi and their presence in soil samples from both farming systems. We found that insects exposed to agroforestry soils died more quickly than insects exposed to full‐sun soils. Of the fungi isolated from the bait insects, Metarhizium was found most frequently, followed by Beauveria. Meanwhile, Fusarium was frequently isolated as primary or secondary infections. We propose that the differential survival of insects is indicative of a greater suppressive potential by entomopathogenic fungi in agroforestry, and that this could be promoted by the diversified landscape, microclimatic stability, and reduced soil disturbance in agroforestry systems. Furthermore, our results provide a useful demonstration of the potential use of the insect bait method to investigate pest‐suppressive potential through bait insect mortality, and we term this the “bait survival technique.”     

医学昆虫的病原真菌

昆虫病原真菌在自然环境中对害虫数量起着抑制作用,在害虫的微生物防治中也占有独特的地位。长期以来,在应用昆虫病原真菌防治人畜害虫方面研究进展较为缓慢,六十年代以来才有较大的发展。本文介绍了近十年的发展概况,并讨论了影响这个研究领域迅速发展的原因及病原真菌在生物防治中应用的潜力。认为尽管至今对这类真菌了解还较少,但通过今后加强有关方面的研究工作,其应用前景是广阔的。     

Phylogenetic and biogeographic implications inferred by mitochondrial intergenic region analyses and ITS1-5.8S-ITS2 of the entomopathogenic fungi <Emphasis Type="Italic">Beauveria bassiana</Emphasis> and <Emphasis Type="Italic">B. brongniartii</Emphasis>

Background  

The entomopathogenic fungi of the genus Beauveria are cosmopolitan with a variety of different insect hosts. The two most important species, B. bassiana and B. brongniartii, have already been used as biological control agents of pests in agriculture and as models for the study of insect host - pathogen interactions. Mitochondrial (mt) genomes, due to their properties to evolve faster than the nuclear DNA, to contain introns and mobile elements and to exhibit extended polymorphisms, are ideal tools to examine genetic diversity within fungal populations and genetically identify a species or a particular isolate. Moreover, mt intergenic region can provide valuable phylogenetic information to study the biogeography of the fungus.     

Resporulation of Metarhizium anisopliae granules on soil and mortality of Tenebrio molitor: Implications for wireworm management in sweetpotato

In Australia, sweetpotato (Ipomoea batatas L.) is vulnerable to root feeding insect pests such as wireworms (e.g., Agrypnus spp.). The number of registered insecticides to control these insect pests is limited and often pest pressure, for example by wireworms, is severe close to harvest, further limiting what insecticides can be applied. Incorporating biological control agents such as entomopathogenic fungi (e.g., Metarhizium anisopliae) into integrated pest management programmes may be feasible in sweetpotato. M. anisopliae has been shown to be effective in controlling more than 200 insects and it is able to reside and grow in the rhizosphere and rhizoplane, suggesting that M. anisopliae could be a promising candidate against soil insect pests. In the study presented here, M. anisopliae was formulated into calcium alginate granules fortified with nutrients. The resporulation of the fungal granules was tested on four different soil types in the laboratory. The biocontrol efficacy of the resulting fungal growth was also examined using larval mealworms, Tenebrio molitor as a model insect in the laboratory and the glasshouse. Our results indicated that sterilised soil favoured optimal fungal resporulation, although different soil types did not have a significant effect on fungal resporulation. The resulting fungal resporulation and growth on sterilised soil caused high mortality (up to 76%) of larval mealworms in the glasshouse, whereas the fungal granules applied to non-sterile soil demonstrated poor resporulation that led to low mortality (13%) of larval mealworms. The result of this study indicates that the manipulation of microbial populations in field soil is required to enhance the fungal growth and potential insect control against wireworms in the field.     

The secretome and chemistry of Metarhizium; a genus of entomopathogenic fungi

The major fungal phyla have independently and repeatedly evolved the ability to overcome the insect defense system, which is usually highly effective against fungal attack. During the genomic era of the last decade, extensive progress has been made in understanding the mechanisms of interactions between insects and their fungal pathogens, particularly Metarhizium spp., the focus of this article. New models of pathogenesis have been designed, new biological phenomena have been discovered, and a plethora of new molecules and functions have been determined that are crucial for successful establishment of fungal disease by permitting: attachment to the insect surface; germination and formation of infection structures; penetration of the host; colonization of host tissue; and sporulation on cadavers. Genomic technologies in particular are producing breakthroughs in some of the more intractable aspects of this field, e.g. evolution of host specificity.