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Entomopathogenic fungi C. militaris and Cordyceps sp. from western Siberia (Novosibirsk oblast): C. militaris stromata (14), Cordyceps sp. stromata (5-8), Lecanicillium sp. (from C. militaris stromata) (9), Isaria farinosa (from C. militaris stromata) (10), and the Paecilomyces like anamorph isolated from the Cordyceps sp. stromata (11). Scale bar: (1-8), 1 cm; (9-11), 20 µm.
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Unique epizootic loci were revealed in the summer-autumn complex of the multispecies communities of forest lepidopterans and
sawflies in two regions of Novosibirsk oblast and in the Baikal region at the border of the Buryat Republic and Irkutsk oblast.
Mass insect mortality was caused by two species of cordycipitoid fungi: Cordyceps militaris and C...
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... simultaneously, the mass death of the regular comembers of one consortium, lepii dopterans and sawflies of the summer-autumn ecoo logical group, but not simply different species of phyy tophagous dissociated in their ecological niches, occurred. Second, simultaneously, two teleomorphic species, C. militaris with typical orangeered stromata ( Fig. 1; 1-4) and 'Cordyceps' sp. with a strawwcolored or light yellow tint of the fruittbearing part of the stroo mata, on average, slightly smaller ( Fig. 1; 5-8), appeared to be the causative agents of the epizootic- but to which of the families and which genus this spee cies should be assigned according to the most recent system of ...
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... not simply different species of phyy tophagous dissociated in their ecological niches, occurred. Second, simultaneously, two teleomorphic species, C. militaris with typical orangeered stromata ( Fig. 1; 1-4) and 'Cordyceps' sp. with a strawwcolored or light yellow tint of the fruittbearing part of the stroo mata, on average, slightly smaller ( Fig. 1; 5-8), appeared to be the causative agents of the epizootic- but to which of the families and which genus this spee cies should be assigned according to the most recent system of cordycipitoid fungi [1] remains to be clarii fied (further molecular genetic studies are expected to be carried ...
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... from the C. miltaris samples coll lected in Novosibirsk oblast and in the Baikal region, ten anamorph isolates with morphological characterr istics completely identical to those indicated in the litt erature [4,[32][33][34] were isolated in culture. Interestt ingly, plating of the stromas of this fungus, along with the typical anamorph ( Fig. 1; 9), often revealed also another anamorph whose characteristics clearly fit in with the diagnosis of the species Isaria farinosa ( Fig. 1; 10). However, it is known that the latter has a Cordyceps memorabilis Cesati teleomorph, which rarely occurs in nature [40]. It should be noted that no unanimity of opinion exists about the generic ...
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... characterr istics completely identical to those indicated in the litt erature [4,[32][33][34] were isolated in culture. Interestt ingly, plating of the stromas of this fungus, along with the typical anamorph ( Fig. 1; 9), often revealed also another anamorph whose characteristics clearly fit in with the diagnosis of the species Isaria farinosa ( Fig. 1; 10). However, it is known that the latter has a Cordyceps memorabilis Cesati teleomorph, which rarely occurs in nature [40]. It should be noted that no unanimity of opinion exists about the generic identity of this fungus: in some sources it is referred to as Lecanicillium sp. [33,34]; in others, as a representative of the genus ...
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... microscopic study of the colonies of the Cordyceps sp. anamorph forming short conidial chains on the phyalides swollen in the basal part ( Fig. 1; 11) Notes: * Assessment in the period of mass appearance of stromata (August). ** Assessment in the period of the peak population (the end of July to August) *** Assessment in the period of nutrition of older age larvae (the end of August). **** Assessment of the wintering stock (September to ...
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... growth of the greater wax moth caterpillars infected with the ascospores and blastospores of both fungal species was insignificantly stunted, and the mortality did not exceed 20%. However, when they were treated with C. militaris conidia, the delay of the caterpillar growth and pupation was markedly proo nounced ( Fig. 1; 12). Pupation was delayed 15- 25 days, the total mortality attained 45%, and the share of cadavers the surfaces of which were overgrown with mycelium in the humid chamber was 1%. When ten pupae killed as result of conidial infection were placed into moist soil, one C. militaris stroma with perr ithecia was formed one and a half months ...
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The earliest pre-Linnaean fungal genera are briefly discussed here with special emphasis on the nomenclatural connection with the genus Cordyceps Fr. Since its valid publication under the basidiomycetous genus Clavaria Vaill. ex L. (Clavaria militaris L. Sp. Pl. 2:1182, 1753), the genus Cordyceps has undergone nomenclatural changes in the post-Linn...
Citations
... Последний встречается в Европе, Северной и Южной Америке, Азии, Африке. В России этот гриб отмечен во всех природно-климатических зонах: от тундры до южных областей [6,7,8,9,10]. Плодовое тело и искусственно культивируемый мицелий C. militaris содержат биоактивные соединения -кордицепиновую кислоту и кордицепин, которые оказывают антибактериальное, противовирусное действие, ингибируют развитие многих видов опухолей, что обеспечивает широкую терапевтическую и профилактическую эффективность кордицепса в медицине и ветеринарии [11,12,13]. Экспериментально показана способность кордицепина влиять на физиологию растений, регулируя накопление лектинов в клетках пшеницы при солевом стрессе [14]. ...
... Результаты и их обсуждение. Обычно C. militaris обнаруживается в лесной подстилке на куколках и гусеницах чешуекрылых из различных семейств, преимущественно бражников (Sphingidae) и близких к ним групп [7]. На куколках бабочек гриб образует булавовидные оранжево-красные плодовые тела -стромы, одиночные или по нескольку штук (рис., а, б). ...
... Например, из-за такой особенности распространения C. militaris был внесён в Красную книгу Республики Бурятии с пометкой: «Численность очень мала, тенденции её изменения не выяснены, лимитирующие факторы не установлены» [24, с. 321]. А спустя несколько лет, при изучении микозов насекомых, на склоне хребта Хамар-Дабан в южном Прибайкалье был зафиксирован всплеск численности C. militaris -до 5000 экземпляров на 1 га [7]. Из литературы известно, что подобная стратегия существования популяции присуща многим видам зоопаразитических грибов: когда широко распространенный вид с низкой численностью характеризуется взрывным ее увели- чением в отдельные годы с благоприятными условиями для развития и размножения насекомых-хозяев [22]. ...
Fungi of the genus Cordyceps Fr. (1818) (Ascomycota: Hypocreales: Cordycipitaceae) are widely distributed throughout the world, many species are cosmopolitan. Cordyceps are known as entomopathogenic fungi that infect caterpillars and pupae of insects. The fruit bodies of Chinese cordyceps Ophiocordyceps sinensis (synonym Cordyceps sinensis (Berk.) Sacc) are also well known for their pharmacological effects and are widely used in traditional Chinese medicine for the treatment of a wide range of diseases. This species has a limited distribution and, due to the huge demand, its resources are rapidly depleted in the wild due to uncontrolled fees. In addition, O. sinensis is difficult to cultivate artificially. Another entomopathogenic species of fungus, Cordyceps militaris, has a chemical composition and properties similar to those of O. sinensis, but, unlike O. sinensis, it is easier to be grown artificially in mycelial culture. In this regard, the study of the distribution and occurrence of the fungus Cordyceps militaris has now acquired exceptional relevance. The article presents data on the first sporadic records of Cordyceps militaris fungus, which has unique biological properties, in the Kirov region. The ecological strategy of the population of this species, the peculiarities of its distribution and population fluctuations in Russia are discussed. The question is considered why, with the wide distribution of this species, it is considered rare and even included in the Red Books of individual regions. C. militaris found in the northern part of the Kirov region was isolated into pure culture (strain T1) for artificial cultivation and study of physiological and biochemical properties. The expediency of including natural cordyceps isolates obtained by introduction into mycelial culture in mycological collection funds is substantiated, since artificial cultivation of the fungus is of obvious interest for the development of biotechnologies in medicine, veterinary medicine, crop production and plant protection.
... Commonly known as the orange caterpillar club fungus, Cordyceps militaris (Clavicipitaceae, Hypocreales, Sordariomycetes, Pezizomycotina, Ascomycota) [ figure 1.1] is a cosmopolitan entomopathogen, with a broad range of insect hosts globally [Kryukov et al. 2011;Shrestha et al. 2012]. Hosts are predominantly lepidopteran larvae (butterfly and moth caterpillars) [Shrestha et al. 2012]. ...
... Another reason lies with the ecology of the species. Cordyceps militaris has a cosmopolitan distribution, occurring in much of Europe, and with over 30 species of lepidopterans and hymenopterans reported as hosts [Kryukov et al. 2011;Shrestha et al. 2012]. Research on the insect infection process of C. militaris is still rudimentary, despite the potential for this species to become a useful insect pathogen for use in pest control. ...
Cordyceps militaris is a widespread entomopathogenic fungus found in Europe, Asia, and North America, with a large number of insect hosts, predominantly lepidopteran larvae (caterpillars). This species is well known for its production of the nucleoside analogue cordycepin (3’-deoxyadenosine). Co-produced with its protector molecule pentostatin, cordycepin is a polyadenylation inhibitor, via its active modified form as cordycepin triphosphate. Pentostatin protects cordycepin from degradation to 3’-deoxyinosine by inhibition of the enzyme adenosine deaminase. It has been shown to have anti-inflammatory effects and hence has been the subject of much pharmacological research. Until recently, very little was known about the role of cordycepin in the ecology of the fungus, or why its production is favoured by natural selection. There are also gaps in the understanding of the process of infection of the host by the fungus, which is directly followed by sexual development and the formation of stromata bearing sexual fruiting bodies and spores (ascospores). Understanding these areas could have implications for biological control of insect pests. Indeed, the related species Beauveria bassiana and Metarhizium anisopliae have been used as bioinsecticides, precluding the use of harmful chemical insecticides. Culture degeneration is a phenomenon defined previously as a reduction in the production of cordycepin by C. militaris. Experiments comparing a degenerated strain of an isolate of C. militaris with its parental control strain were performed, involving the use of gene expression analysis and metabolomics. Reduced cordycepin production in the degenerated strain was shown to be accompanied by declines in sexual development-related gene expression, and reduced production of other metabolites involved in the citrate cycle and purine metabolism. This suggested a link between cordycepin production, primary metabolism, and sexual development. We hypothesised that the production of cordycepin by C. militaris aids the infection of the insect by suppression of the host immune system, and that pentostatin, by providing molecular protection, enhances this effect. In a caterpillar infection assay system involving the injection of spores into the model species Galleria mellonella (greater wax moth) caterpillars, the lower cordycepin-producing degenerated strain was shown to produce a significantly-decreased pathogenic response, marked by reduced fungal growth in the host. When spores of the degenerated strain were supplemented by cordycepin and pentostatin, fungal emergence rates and levels significantly increased, restoring the infection performance of the degenerated strain to that of the parental control. Assays of insect gene expression were also performed, and cordycepin was demonstrated to suppress the upregulation of immune response genes in both Drosophila melanogaster Schneider 2 cells and G. mellonella haemolymph cells. Pentostatin enhanced the effects of low cordycepin concentrations in both models. These findings support the hypothesis that cordycepin has an important role in aiding insect infection by the fungus via immune suppression, and that the effect of cordycepin on host cell responses is maintained by pentostatin. Biosynthesis genes (Cns genes) for cordycepin and pentostatin are located in the same gene cluster. We hypothesised that cordycepin-pentostatin co-production was a rare trait, and its evolution had been resultant partly due to horizontal gene transfer between different species. This was due to the lack of cordycepin in other Cordyceps species, and genetic evidence of its production only found previously in two other, distantly-related species. Bioinformatics work involving tBLASTn searches through the sequenced genomes of over two and a half thousand fungal species uncovered evidence of homologous Cns gene clusters in five new species. This together with consideration of protein structures suggests that the development of cordycepin-pentostatin co-production has occurred by convergent evolution involving duplication and subfunctionalisation of genes involved in the purine synthesis pathway, and/or through horizontal gene transfer.
... Hosts. Commonly on Lepidoptera larvae and pupae, infrequently on Hymenoptera (Kobayasi 1941;Kryukov et al. 2011). ...
Species of Cordyceps sensu lato (Hypocreales, Sordariomycetes) have always attracted much scientific attention for their abundant species diversity, important medicinal values and biological control applications. The insect superfamilies Elateroidea and Tenebrionoidea are two large groups of Coleoptera and their larvae are generally called wireworms. Most wireworms inhabit humid soil or fallen wood and are often infected with Cordyceps s.l. However, the species diversity of Cordyceps s.l. on Elateroidea and Tenebrionoidea is poorly known. In the present work, we summarise taxonomic information of 63 Cordyceps s.l. species that have been reported as pathogens of wireworms. We review their hosts and geographic distributions and provide taxonomic notes for species. Of those, 60 fungal species are accepted as natural pathogens of wireworms and three species ( Cordyceps militaris , Ophiocordyceps ferruginosa and O. variabilis ) are excluded. Two new species, O. borealis from Russia (Primorsky Krai) and O. spicatus from China (Guizhou), are described and compared with their closest allies. Polycephalomyces formosus is also described because it is reported as a pathogen of wireworms for the first time. Phylogeny was reconstructed from a combined dataset, comprising SSU, LSU and TEF1-α gene sequences. The results, presented in this study, support the establishment of the new species and confirm the identification of P. formosus .
... While present each year, it was the most prevalent entomopathogen in 2015 and 2016. Species from other families should be considered potential (opportunistic) pathogens of Lepidoptera as most of them belong to large genera including saprotrophs and pathogens of various hosts, with some representatives occasionally found also on insects [23][24][25]. There are exceptions from this general scheme, however. ...
... The lab-based estimates can underestimate the prevalence of entomopathogens in nature as the insects in the laboratory culture should be less exposed to various potential sources of infection than in the field, e.g., soil or infected insects [23,27]. Alternatively, laboratory mass rearings of insects might be prone to disease outbreaks, leading to higher prevalence values in the lab compared to the field. ...
The knowledge about the diversity and ecological role of entomopathogenic fungi is primarily based on agroecosystems whereas information derived from natural insect populations is much more limited. To contribute to filling this gap, we recorded the prevalence of fungal infections in laboratory rearing experiments with five species of Lepidoptera, and in a field rearing experiment including one of these moths. The diversity of detected fungi was found to be high; we isolated 25 species of fungi from insects that had died in the course of these experiments. Six species belonged to the family Cordycipitaceae known to include unambiguous insect pathogens. The trophic niche of the representatives of other taxa is less clear and requires further studies. Regarding the most abundant species, Cordyceps farinosa, in which this question could be addressed, there was no indication of specialization on particular insect hosts, whereas several of the less common species may have been recorded from lepidopteran hosts for the first time. Across the subsets of the data, the prevalence of fungal infections generally remained below 5%. Our results are thus consistent with the idea that entomopathogenic fungi are always present in insect populations but rarely reach epizootic levels. The detected species richness shows that much is to be gained from mapping the diversity of fungal species associated with folivorous insects in natural populations.
... The ascomycete Cordyceps militaris is characterized by a restricted host and habitat range and has a highly reduced number of genes involved in secondary metabolism and the synthesis of proteases compared to generalists such as Metarhizium robertsii and Beauveria bassiana [50,53]. This fungus mainly infects forest lepidopterans (Lepidoptera, Macroheterocera) in the larval and pupal stages which are located in the soil, forest flour and fallen wood [54][55][56]. Previously, natural infections of insects with C. militaris were studied insufficiently. It is known that larvae and pupae could be infected by topical application with ascospores or conidia in a laboratory [22,57]. ...
... However this method is difficult to reproduce, and the outcomes strongly depend on the physiological state of the host [22]. In contrast, injection of lepidopteran larvae and pupae with blastospores or conidia has led to more stable development of the mycosis [55,58,59]. Importantly, spontaneous bacterial infections have been constantly documented after infection of C. militaris with insects in laboratory conditions [22,58,59]. ...
... This is consistent with the more specific conditions for cultivation required by C. militaris in vivo or in vitro [70]. Occurrence of bacterioses after topical infection or injection of C. militaris in insects has been documented previously [22,55,58,71]. We suggest that C. militaris has a limited ability to suppress host commensal bacteria, as the fungus is associated with narrow environmental requirements, including a specific temperature range [22]. ...
Various insect bacterial associates are involved in pathogeneses caused by entomopathogenic fungi. The outcome of infection (fungal growth or decomposition) may depend on environmental factors such as temperature. The aim of this study was to analyze the bacterial communities and immune response of Galleria mellonella larvae injected with Cordyceps militaris and incubated at 15 • C and 25 • C. We examined changes in the bacterial CFUs, bacterial communities (Illumina MiSeq 16S rRNA gene sequencing) and expression of immune, apoptosis, ROS and stress-related genes (qPCR) in larval tissues in response to fungal infection at the mentioned temperatures. Increased survival of larvae after C. militaris injection was observed at 25 • C, although more frequent episodes of spontaneous bacteriosis were observed at this temperature compared to 15 • C. We revealed an increase in the abundance of enterococci and enterobacteria in the midgut and hemolymph in response to infection at 25 • C, which was not observed at 15 • C. Antifungal peptide genes showed the highest expression at 25 • C, while antibacterial peptides and inhibitor of apoptosis genes were strongly expressed at 15 • C. Cultivable bacteria significantly suppressed the growth of C. militaris. We suggest that fungi such as C. militaris may need low temperatures to avoid competition with host bacterial associates.
... Cordyceps militaris (Cm) is a species that forms teleomorphs and has a restricted host range. The fungus causes mycoses mainly among moth (Lepidoptera, Macroheterocera) larvae and pupae that inhabit forest coenoses [7][8][9][10]. C. militaris is characterized by a prolonged life cycle, https://doi.org/10.1016/j.micpath.2020.103995 Received 31 December 2019; Received in revised form 18 January 2020; Accepted 21 January 2020 at least 30 days from infection to perithecia formation [11], as well as by relatively slow mycelial growth on artificial media. ...
... This lack of information is related to the locality of their distribution, which is caused by the continental climate and is not optimal for the development of these fungi. That have been sporadic studies of these fungi in Siberia [Ogarkov, Ogarkova, 2000;Gorbunova et al., 2011;Kryukov et al., 2011Kryukov et al., , 2018. ...
... Грибы Cordyceps millitaris выступают естественными регуляторами численности чешуекрылых, вы-зывая периодические эпизоотии в их популяциях [Kamata, 2000;Kryukov et al., 2011]. Метаболиты этого гриба активно изучаются в фармакологии [Holliday, Cleaver, 2008;Paterson, 2008;Tuli et al., 2013], однако их роль в развитии микозов насекомых изучена очень слабо. ...
This study examines accumulation dynamics of
Cordyceps millitaris toxins in wax moth Galleria mellonella
larvae after injection of fungal blastospores. At 3 and 4 days
post infection average concentration of cordycepin was 244
± 41 ng per 100 mg of dry mass of larvae. Accumulation of
cordycepin was delayed one day in larvae, that were
envenomated, by parasitoid Habrobracon hebetor (Say).
Cordycepin concentration was correlated with hyphal body
count in hemolymph, but only on tendency level (P > 0.07).
The role of Cordyceps millitaris toxins in the development of
mycoses was discussed.
... Cordyceps militaris is a teleomorphic insect pathogenic ascomycete characterized by long development in/on hosts. In temperate zones, the fungal life cycle lasts up to 1 y (Kryukov et al., 2011;Gedminas et al., 2015). The fungus is broadly distributed in the world, but its habitat and host ranges are restricted. ...
... The fungus is broadly distributed in the world, but its habitat and host ranges are restricted. This pathogen causes substantial mycoses in forest lepidopterans (Lepidoptera, Macroheterocera) and sawflies (Hymenoptera, Cimbicidae), inducing epizootics in these populations (Kamata, 1998(Kamata, , 2000Kryukov et al., 2011;Gedminas et al., 2015). Mycosis develops in larvae, prepupae and pupae hibernating in the forest floor and fallen wood (Kamata, 1998;Gedminas et al., 2015) indicating a certain level of adaptation to the hibernation period of the host life cycle. ...
... Mycosis develops in larvae, prepupae and pupae hibernating in the forest floor and fallen wood (Kamata, 1998;Gedminas et al., 2015) indicating a certain level of adaptation to the hibernation period of the host life cycle. Interestingly, C. militaris often prevails over other fungal species (Beauveria, Metarhizium) killing lepidopterans during hibernation (Kamata, 1998;Kryukov et al., 2011). Thus, it is presumed that C. militaris has some competitive advantages over Metarhizium and Beauveria due to better adaptation to cold conditions and colonization of insects in the state of diapause. ...
Temperature preferences, virulence properties and genetic relationships inferred from tef1-alpha gene sequencing were determined for Cordyceps militaris isolates from the Palearctic region. The variability of the mycelial growth rate of different isolates was extremely low at 5–30 °C. However, correlations among genetic variation, geographic origin and temperature responses were established. C. militaris cultures were not
heat-active but were more psychrotolerant than Beauveria and Metarhizium isolates. The thermo-physiological state of the wax moth Galleria mellonella had a crucial role for the development of C. militaris. Larval diapause induced by decreasing temperature (15 °C) was most favourable for development of mycosis due to a drop in innate immunity. While fought off at 25°C in the larvae, the infection could still be activated in pupae under cold conditions (15 °C). Differences in ecology and pathogenic strategies of C. militaris and the
generalist fungi Beauveria and Metarhizium are discussed.
... The physiological and biochemical mechanisms of the synergistic effect of different pathogens are practically unknown. Earlier we have demonstrated the pathogenic action of the fungus Cor-dyceps militaris on various lepidopteran species, manifested by lower survival rates and a severe development delay in the larvae (Kryukov et al., 2011). In addition, infection of the larvae of the tent caterpillar moth Malacosoma parallela Staud. ...
The synergistic effect in mortality of the greater wax moth Galleria mellonella larvae was recorded after combined treatment with the entomopathogenic fungi Cordyceps militaris (L.: Fr.) Link and Beauveria bassiana (Bals.-Criv.) Vuill. Treatment with C. militaris resulted in development arrest and some changes in immune response (a sharp decrease in the total hemocyte counts and encapsulation rate and an increase in phenoloxidase activity in the hemolymph) which were accompanied by higher susceptibility to B. bassiana. The larvae killed by combined treatment with two pathogens were colonized only by B. bassiana. The mechanisms of synergism under combined treatment of the greater wax moth with C. militaris and B. bassiana are discussed.
