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Habitat and geographic distribution of Hypocrea species included in this study 

Habitat and geographic distribution of Hypocrea species included in this study 

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Approximately 950 individual sequences of nonribosomally biosynthesised peptides are produced by the genus Trichoderma/Hypocreathat belong to a perpetually growing class of mostly linear antibiotic oligopeptides, which are rich in the non-proteinogenic α-aminoisobutyric acid (Aib). Thus, they are comprehensively named peptaibiotics. Notably, peptai...

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... An example of the latter is a widespread peptaibols, that is, oligopeptides that contain nonstandard amino acids. They are formed through polyenzymatic synthesis by many terrestrial organisms, in particular, by fungi and play the role of antibiotics that modify microbial membranes and disrupt their function (Raap et al., 2005;Röhrich et al., 2014;Shi et al., 2020). However, in the case of significant changes in the environment, such a modification may lead to membrane stabilization and contribute to the survival of the organism. ...
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... Most data have been published during a period when phylogenetics was not recognized as essential to fungal taxonomy. As a result, many peptaibiotics cannot be linked to a source organism in public collections , and most of the producers reported in literature have never been deposited appropriately (Röhrich et al., 2014;Rush et al., 2021). The efforts devoted to investigate new strains and compounds must be increased and protocols should be updated to state-of-the-art methodologies to increase reliability. ...
Article
Agricultural systems are in need of low-cost, safe antibiotics to protect crops from pests and diseases. Peptaibiotics, a family of linear, membrane-active, amphipathic polypeptides, have been shown to exhibit antibacterial, antifungal, and antiviral activity, and to be inducers of plant resistance against a wide range of phytopathogens. Peptaibiotics belong to the new generation of alternatives to agrochemicals, aligned with the United Nations Sustainable Development Goals and the One Health approach toward ensuring global food security and safety. Despite that, these fungi-derived, non-ribosomal peptides remain surprisingly understudied, especially in agriculture, where only a small number has been tested against a reduced number of phytopathogens. This lack of adoption stems from peptaibiotics' poor water solubility and the difficulty to synthesize and purify them in vitro, which compromises their delivery and inclusion in formulations. In this review, we offer a comprehensive analysis of peptaibiotics' classification, biosynthesis, relevance to plant protection, and mode of action against phytopathogens, along with the techniques enabling researchers to extract, purify, and elucidate their structure, and the databases holding such valuable data. It is also discussed how chemical synthesis and ionic liquids could increase their solubility, how genetic engineering and epigenetics could boost in vitro production, and how omics can reduce screenings' workload through in silico selection of the best candidates. These strategies could turn peptaibiotics into effective, ultra-specific, biodegradable tools for phytopathogen control.
... Trichoderma spp. comprises more than 200 validly described species distributed in soils worldwide and across various habitats and are considered a valuable resource for structurally novel natural products with diverse bioactivities, including biological control of phytopathogens 19 . In the interest of obtaining more effective methods of pathogen control, plant growth-promoting rhizosphere microorganisms have been used as a consortium or in combination with chemical pesticides by our group and other authors [20][21][22][23] . ...
... Trichoderma is a genus that belongs to the family Hypocreaceae and comprises many different fungi strains found in most diverse ecosystems 19 . Trichoderma strains proliferate and have a characteristic morphology, white and cottons at the beginning, then developing into yellowish green to deep green compact tufts. ...
... Direct mechanisms are mycoparasitism and the production of active metabolites and lytic enzymes 3,31 . Mycoparasitism, the ability to parasitize on fungi, is a unique characteristic of Trichoderma since they can parasitize even taxonomically close species 19 . The antifungal activity of Trichoderma against phytopathogenic fungi is attributed to the combined action of secondary metabolites (SMs) and hydrolytic enzymes i.e., cellulases, proteases, chitinases, and xylanases 3,32,33 . ...
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... The important role of Trichoderma strains is partially due to their secondary metabolites, out of which, approximately 80% belong to the peptaibiotics class [57]. One large group of peptaibiotics is the peptaibols which are characterized by the generally acetylated Nterminus, the amino-alcohol group like isoleucinol and phenylalaninol on the C-terminus, and the presence of non-proteinogenic amino acids, such as α-aminoisobutyric acid (Aib), hydroxyproline or D-isovaline [62]. ...
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Integrated disease management and plant protection have been discussed with much fervor in the past decade due to the rising environmental concerns of using industrially produced pesticides. Members of the genus Trichoderma are a subject of considerable research today due to their several properties as biocontrol agents. In our study, the peptaibol production of Trichoderma longibrachiatum SZMC 1775, T. longibrachiatum f. bissettii SZMC 12546, T. reesei SZMC 22616, T. reesei SZMC 22614, T. saturnisporum SZMC 22606 and T. effusum SZMC 22611 were investigated to elucidate structure-activity relationships (SARs) between the properties of peptaibols and their 3D structures. The effects of peptaibol mixtures obtained from every Trichoderma strain were examined against nine commonly known bacteria. The lowest minimum inhibitory concentrations (MIC, mg ml−1) were exerted by T. longibrachiatum f. bissettii SZMC 12546 against Gram-positive bacteria, which was also able to inhibit the plant pathogenic Gram-negative Rhizobium radiobacter. Accelerated molecular dynamics (aMD) simulations were performed in aqueous solvent to explore the folding dynamics of 12 selected peptaibol sequences. The most characteristic difference between the peptaibols from group A and B relies in the ‘Gly-Leu-Aib-Pro’ and ‘Gly-Aib-Aib-Pro’ motifs (‘Aib’ stands for α-aminoisobutyric acid), which imparted a significant effect on the folding dynamics in water and might be correlated with their expressed bioactivity. In our aMD simulation experiments, Group A peptaibols showed more restricted folding dynamics with well-folded helical conformations as the most stable representative structures. This structural stability and dynamics may contribute to their bioactivity against the selected bacterial species.
... Several therapeutic agents and major compounds of commercially important medicines comprise natural products used in therapies for autoimmune disorders, cancer, microbial infections, neurological disorders, and cardiovascular diseases (Newman and Cragg, 2016). Some agriculture-related chemicals are also natural by-products (Rimando and Duke, 2006;Asolkar et al., 2013). Mainly, fungi are a significant source of bioactive secondary metabolites (Peláez, 2005) that have immensely improved animal and human health. ...
... Many secondary metabolites are identified due to their association with a biological process or activity sensed outside laboratory conditions, e.g., pigments, mycotoxins (fungal toxins), phytotoxins. However, secondary metabolites of some fungi are characterized and correlated with phylogenetic classifications (Röhrich et al., 2014;Stadler et al., 2014). ...
... Several therapeutic agents and major compounds of commercially important medicines comprise natural products used in therapies for autoimmune disorders, cancer, microbial infections, neurological disorders, and cardiovascular diseases (Newman and Cragg, 2016). Some agriculture-related chemicals are also natural by-products (Rimando and Duke, 2006;Asolkar et al., 2013). Mainly, fungi are a significant source of bioactive secondary metabolites (Peláez, 2005) that have immensely improved animal and human health. ...
... Many secondary metabolites are identified due to their association with a biological process or activity sensed outside laboratory conditions, e.g., pigments, mycotoxins (fungal toxins), phytotoxins. However, secondary metabolites of some fungi are characterized and correlated with phylogenetic classifications (Röhrich et al., 2014;Stadler et al., 2014). ...
... Several therapeutic agents and major compounds of commercially important medicines comprise natural products used in therapies for autoimmune disorders, cancer, microbial infections, neurological disorders, and cardiovascular diseases (Newman and Cragg, 2016). Some agriculture-related chemicals are also natural by-products (Rimando and Duke, 2006;Asolkar et al., 2013). Mainly, fungi are a significant source of bioactive secondary metabolites (Peláez, 2005) that have immensely improved animal and human health. ...
... Many secondary metabolites are identified due to their association with a biological process or activity sensed outside laboratory conditions, e.g., pigments, mycotoxins (fungal toxins), phytotoxins. However, secondary metabolites of some fungi are characterized and correlated with phylogenetic classifications (Röhrich et al., 2014;Stadler et al., 2014). ...
... Another striking property of Trichoderma that makes it a useful model of taxonomic studies is the evident lack of hidden diversity or "dark Trichoderma species" (Migheli et al. 2009;Friedl and Druzhinina 2012;Hagn et al. 2007;Meincke et al. 2010;López-Quintero et al. 2013;Röhrich et al. 2014;Jaklitsch 2009Jaklitsch , 2011 2015), meaning that most or all species can be successfully cultivated in vitro. Therefore, Trichoderma spp. ...
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Modern taxonomy has developed towards the establishment of global authoritative lists of species that assume the standardized principles of species recognition, at least in a given taxonomic group. However, in fungi, species delimitation is frequently subjective because it depends on the choice of a species concept and the criteria selected by a taxonomist. Contrary to it, identification of fungal species is expected to be accurate and precise because it should predict the properties that are required for applications or that are relevant in pathology. The industrial and plant-beneficial fungi from the genus Trichoderma (Hypocreales) offer a suitable model to address this collision between species delimitation and species identification. A few decades ago, Trichoderma diversity was limited to a few dozen species. The introduction of molecular evolutionary methods resulted in the exponential expansion of Trichoderma taxonomy, with up to 50 new species recognized per year. Here, we have reviewed the genus-wide taxonomy of Trichoderma and compiled a complete inventory of all Trichoderma species and DNA barcoding material deposited in public databases (the inventory is available at the website of the International Subcommission on Taxonomy of Trichoderma www.trichoderma.info ). Among the 375 species with valid names as of July 2020, 361 (96%) have been cultivated in vitro and DNA barcoded. Thus, we have developed a protocol for molecular identification of Trichoderma that requires analysis of the three DNA barcodes (ITS, tef1 , and rpb2 ), and it is supported by online tools that are available on www.trichokey.info . We then used all the whole-genome sequenced (WGS) Trichoderma strains that are available in public databases to provide versatile practical examples of molecular identification, reveal shortcomings, and discuss possible ambiguities. Based on the Trichoderma example, this study shows why the identification of a fungal species is an intricate and laborious task that requires a background in mycology, molecular biological skills, training in molecular evolutionary analysis, and knowledge of taxonomic literature. We provide an in-depth discussion of species concepts that are applied in Trichoderma taxonomy, and conclude that these fungi are particularly suitable for the implementation of a polyphasic approach that was first introduced in Trichoderma taxonomy by John Bissett (1948–2020), whose work inspired the current study. We also propose a regulatory and unifying role of international commissions on the taxonomy of particular fungal groups. An important outcome of this work is the demonstration of an urgent need for cooperation between Trichoderma researchers to get prepared to the efficient use of the upcoming wave of Trichoderma genomic data.
... Within the biosphere, Aib is known to be produced exclusively by saprotrophic filamentous fungi that are both "ubiquitous and cosmopolitan, even occurring in marine, Arctic and Antarctic regions" (Röhrich et al., 2012(Röhrich et al., : 1221. More than 30 genera containing a vast number of species are now known to produce Aib (Brückner et al., 2009;Brückner et al., 2019), and this number is ever-increasing (Röhrich et al., 2012(Röhrich et al., , 2014. Among the most common and speciose genera known to produce Aib today is the genus Trichoderma Pers. ...
... Among the most common and speciose genera known to produce Aib today is the genus Trichoderma Pers. (Hypocreaceae Fr.) (Brückner et al., 2009;Röhrich et al., 2014). Therefore, techniques for distinguishing between biotically and abiotically produced amino acids have been developed (Elsila et al., 2011). ...
... In this study I suggest that a western North American fungal spike may be recorded in the Aib signature of the K/Pg boundary clay, such as at Starkville South or Sussex, Wyoming (Fig. 2). Brückner et al. (2009) andElsila et al. (2011) have implored researchers to question whether claims of extraterrestrial Aib in ancient sediments are instead the result of fungal contamination or the presence of durable Aib-rich peptaibols, which are produced by common soil fungi such as Trichoderma (Brückner et al., 2009;Röhrich et al., 2014;Brückner et al., 2019). Although Brückner et al. (2009) point out that it may be difficult or impossible to differentiate between ancient and modern Aib in sediments, the results of recent phylogenetic and genomic studies conducted by Kubicek et al. (2019) support the contention that Aib within the K/Pg boundary clay could have been produced by Trichoderma or similar soil fungi. ...
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The presence of the amino acid α-aminoisobutyric acid (Aib) within Cretaceous/Paleogene (K/Pg) boundary clay in the Raton and Powder River basins in Colorado and Wyoming, respectively, has been described as compelling evidence that extraterrestrial Aib survived the high-energy Chicxulub impact. Based on contemporary experiments and simulations, however, it is highly unlikely that extraterrestrial Aib survived the impact, which had peak impact pressures and temperatures in excess of 600 GPa and 10,000 K, respectively. In other words, the amino acid signature of the carbonaceous chondritic asteroid that impacted Chicxulub was undoubtedly destroyed upon impact during formation of the vapor plume or so-called “fireball.” The only organisms known to produce Aib are the suite (more than 30 genera) of cosmopolitan saprotrophic filamentous fungi that include Trichoderma Pers., which has recently been hypothesized to have thrived during the K/Pg mass-extinction event. Therefore it is proposed that the Aib horizon in the K/Pg boundary clay in the Raton and Powder River basins correlates with the K/Pg boundary fungal spike, which thus far has only been observed in New Zealand (Southern Hemisphere). This proposition is based upon superimposing the Aib horizon on the well-known iridium and fern-spore spikes, as its stratigraphic position precisely matches that predicted by the fungal spike. If correct, this hypothesis alters the conventional perspective on the tempo and mode of terrestrial ecosystem recovery in western North America, as the heavily sampled K/Pg boundary section in the Raton Basin was instrumental in shaping the traditional narrative of the rapid recolonization of a denuded landscape by ferns via wind-blown spores in the immediate wake of regional deforestation caused by the K/Pg impact event. Perhaps more importantly, it could present an alternative to traditional palynological approaches for locating the fungal spike in other terrestrial K/Pg boundary sections and could provide additional support for the generalization that global mass-extinction events are frequently accompanied by fungal spikes.