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论文范文
1. Trichoderma reesei: Environmental and Lignocellulosic Biomass Degrader Trichoderma species are ubiquitous and cosmopolitan. They are very efficient colonizers of a variety of habitats and can be found from the tundra to the tropics [1], especially in lignocellulosic material and plant rhizospheres, and this effectiveness is translated by the ability of competently degrading the available substrate and of secreting different enzymes and metabolites used in the process [2–5]. The capability of growing in such a variety of carbon sources is also due to the high and fast capacity of responding to diverse environmental signals, being able to adapt according to that current background and regulate its growth, conidiation, and the production of enzymes and secondary metabolites. These signals may vary from different nutrients found in the milieu to the absence and presence of light, and adjusting to them is crucial for the survival of the microorganism [2, 4, 6–8]. As a result of this versatility, Trichoderma species are very useful in many aspects that range from plant biocontrol [9] to various sorts of industries [10–13], especially for the cellulolytic enzymes produced by them. Among all species from this genus and which are industrially used, Trichoderma reesei is the most studied one regarding lignocellulosic biomass degradation, since it is the main producer of cellulolytic and xylanolytic enzymes [14–17]. The ability of growing in a wide range of carbon sources allows great variability in the production of cellulases, since the gene expression and secretion of enzymes are directly dependent on the different chemical signals produced from the diverse substrates. Considering that the plant biomass, one of the most important and complex substrates used by Trichoderma, is composed of mono-, di-, and polysaccharides, the different sugars may have different levels of induction or repression of cellulase genes. Some of the cellulase inducers are cellulose, β-glucan, xylan, lactose, cellobiose, and sophorose, while glucose is the main repressor carbon source [18]. When T. reesei degrades the lignocellulosic biomass, cellobiose may be converted into sophorose by a transglycosylation activity of a β-glucosidase [19, 20]. The comparison of the genomes of Trichoderma species, including T. reesei, suggests they have a mycoparasitic common ancestral, probably from fungi that degrade lignocellulosic material. Considering this, T. reesei may have maintained the mycoparasitic characteristic, which allows it to have advantages over other species when competing for substrate, through the conversion of cellobiose into sophorose by transglycosylation to be metabolized [21, 22]. Differently from other fungi, in T. reesei, sophorose acts as a very potent cellulase inducer in very low concentrations, being able to induce the expression of some xylanases as well [23, 24]. 
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