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Trichoderma reesei

Trichoderma reesei is a mesophilic, filamentous ascomycete fungus in the genus Trichoderma, best known as the primary industrial producer of cellulases and hemicellulases for degrading lignocellulosic biomass into fermentable sugars. It is the anamorph (asexual form) of the teleomorph Hypocrea jecorina and thrives in soil environments rich in decaying plant material, where it exhibits strong cellulolytic activity. Originally isolated in 1944 during World War II from a site in the Solomon Islands, where it was observed breaking down U.S. Army tent canvas, the species was named in honor of microbiologist Elwyn T. Reese, who advanced its early study for potential applications in cellulose utilization. Since its discovery, T. reesei has been the cornerstone of industrial enzyme production, with strains like QM6a and its derivatives subjected to and to boost secretion capacities—often exceeding 100 g/L of protein in optimized fermentations. Its , sequenced in 2008, reveals a compact 34 Mb assembly with around 9,000 genes, including gene clusters dedicated to carbohydrate-active enzymes (CAZymes) that enable synergistic breakdown of and . The fungus operates heterothallically, with mating-type loci (MAT1-1 and MAT1-2) facilitating under laboratory conditions, which has aided strain improvement efforts. The industrial significance of T. reesei lies in its role across multiple sectors: in biofuels, it powers the enzymatic step for converting agricultural residues into precursors, reducing reliance on fuels. In the , its xylanases and cellulases improve bleaching processes and fiber modification, while in textiles and , they enable biofinishing, denim washing, and juice clarification. Safety assessments confirm T. reesei as non-pathogenic to humans and lacking production under production conditions, supporting its widespread use in commercial formulations. Ongoing focuses on enhancing its robustness against inhibitors and expanding protein expression for broader biotechnological applications.

Taxonomy and history

Taxonomy

Trichoderma reesei is classified within the kingdom Fungi, phylum , subphylum , class , subclass Hypocreomycetidae, order , family Hypocreaceae, and genus . This filamentous ascomycete represents the anamorphic (asexual) stage of its . The species was formally described by Emory G. Simmons in 1977, who proposed it as the anamorph of the teleomorph Hypocrea jecorina, originally described by M.J. and C.E. Broome as Hypocrea jecorina in 1873 (published 1875). Simmons established this connection based on morphological characteristics observed in isolates from tropical regions. Subsequent molecular analyses in 1996 confirmed that T. reesei is a clonal derivative of H. jecorina, supporting the synonymy. Under the "one fungus, one name" principle adopted in fungal nomenclature, Trichoderma reesei is the accepted name, with Hypocrea jecorina recognized as a heterotypic referring to the sexual morph. No other synonyms are currently in widespread use for this .

Discovery and historical development

Trichoderma reesei was first isolated in 1944 from degraded canvas tents used by the U.S. Army on in the during . The fungus was identified as the causative agent of "jungle rot" that was compromising military equipment in the humid tropical environment. This isolation was conducted by Elwyn T. Reese at the Quartermaster Corps Laboratories, leading to the reference strain QM6a, initially classified as a variant of . Following its isolation, early research on T. reesei from the late 1940s through the 1970s was primarily carried out at the U.S. Army Natick Development Center, where Elwyn T. Reese and Mary Mandels investigated its exceptional cellulolytic capabilities for degrading textiles and other cellulosic materials. This work focused on understanding microbial breakdown to address logistical challenges in military supply chains, including the development of assays for activity and optimization of production under various conditions. By the and , studies had revealed T. reesei's superior efficiency in producing extracellular cellulases compared to other fungi, establishing it as a for enzymatic degradation research. The marked a pivotal shift in T. reesei's research trajectory, redirecting efforts from military applications toward biofuel production as part of broader initiatives to convert into fermentable sugars. This transition was driven by the need for alternative energy sources amid global energy shortages, leading to mutagenesis programs at Natick and that enhanced yields by over 20-fold, culminating in hyperproductive strains like RUT-C30. These developments laid the groundwork for industrial-scale production. Key milestones in the late 20th and early 21st centuries further advanced T. reesei's scientific profile. In 1996, molecular analyses confirmed that T. reesei is the anamorph of the ascomycete Hypocrea jecorina, resolving its taxonomic placement through sequence comparisons of regions. The complete genome sequence of the QM6a strain was published in 2008, revealing a compact 34 genome optimized for lignocellulose degradation and providing insights into its regulatory mechanisms.

Morphology and ecology

Morphological characteristics

Trichoderma reesei is a mesophilic filamentous fungus, exhibiting optimal growth at temperatures between 25 and 30°C. It possesses septate hyphae that are , lacking pigmentation in the vegetative stage, which facilitates its rapid colonization of substrates. The hyphae are typically branched and extend apically, forming a network of that supports efficient uptake and . On solid agar media such as potato dextrose agar (PDA), T. reesei forms fast-growing colonies with a cottony, white aerial mycelium that transitions to green as conidiophores develop and produce conidia. The reverse side of colonies often appears yellow due to diffusible pigments. When grown on cellulosic substrates like carboxymethyl cellulose (CMC) agar, the colonies exhibit visible clearing zones or halos around them, resulting from the degradation of the substrate by secreted cellulolytic enzymes. This morphological trait underscores its role as a potent decomposer. Microscopically, T. reesei features branched conidiophores arising from the hyphae, terminating in phialides that are lageniform to ampulliform in shape. These phialides produce chains of green conidia, which are subglobose to ellipsoidal, smooth-walled, and measure 3–5 μm in length. The conidia are typically one-celled and serve as the primary means of dispersal.

Habitat and ecological role

Trichoderma reesei is primarily found in tropical and subtropical soils rich in , as well as on decaying wood and plant debris, where it thrives in environments abundant with lignocellulosic substrates. However, wild isolates of T. reesei remain rare, with only a few documented beyond the original tropical site, highlighting its primary association with industrial and laboratory settings. Originally isolated from a tent in the during , the fungus was identified for its ability to degrade cellulose-based materials. Despite its tropical origins, T. reesei exhibits a , having been reported in diverse global locations including temperate regions like and various forest ecosystems. Ecologically, T. reesei occupies a saprotrophic niche as a key of lignocellulose, contributing to the breakdown of biomass and facilitating nutrient recycling in ecosystems. It produces a suite of extracellular enzymes, including cellulases and hemicellulases, that enable efficient degradation of complex carbohydrates, playing a vital role in carbon turnover within litter and wood decay processes. Like other species, T. reesei possesses potential for mycoparasitism, antagonizing pathogenic fungi through and limited direct , though it is primarily recognized for its saprotrophic role rather than strong biocontrol activity. The has adapted to its habitats through tolerance to acidic conditions, with optimal occurring at levels between 3 and 6, common in organic-rich soils. This resilience, combined with its proficiency in colonizing cellulose-rich substrates such as pre-degraded wood and root exudates in the , allows T. reesei to exploit nutrient-scarce niches opportunistically. These traits enhance its competitive edge in microbial communities, supporting its persistence across varied environmental gradients.

Reproduction and life cycle

Asexual reproduction

Trichoderma reesei primarily reproduces asexually through conidiation, a in which unicellular conidia are produced mitotically on specialized structures called phialides, which arise from branched conidiophores. These conidiophores emerge as erect, variably branched hyphae that form in unpaired patterns, supporting flask-shaped (lageniform or ampulliform) phialides measuring approximately 5–10 μm in length. The phialides enteroblastically generate chains of conidia, which are green, slightly roughened, and ellipsoidal to subglobose in shape, typically 2.5–4.0 × 2.0–3.0 μm in size. This clonal propagation enables efficient dispersal and colonization in diverse environments. The asexual life cycle begins with conidial under favorable conditions, such as nutrient availability and , where a swells and extends one or more germ tubes to form initial hyphae. These hyphae exhibit septate, branching growth, establishing a vegetative that expands radially on solid substrates. Under nutrient-rich conditions, like those provided by malt extract , the differentiates into aerial hyphae after 18–24 hours of growth; these aerial structures then branch into conidiophores, initiating sporulation around 21–27 hours. Mature conidia accumulate, turning the dark green, and serve as propagules for new infections or inocula. Conidiation in T. reesei is tightly regulated by environmental cues, including and nutrient status. Exposure to (380–500 nm) or a brief UV pulse triggers photoconidiation via photoreceptors BLR1 and BLR2, leading to rapid aerial formation and a ring of spores at the colony periphery within 36 hours; continuous promotes uniform sporulation, while light-dark cycles produce concentric rings. availability also modulates the process: vegetative hyphal growth occurs on carbon-rich media, but carbon or deprivation, particularly a high C:N ratio or low , enhances sporulation rates, often synergizing with light signals. Additionally, air exposure generates that initiate differentiation, while mechanical injury can induce localized conidiation under acidic conditions (pH 2.8–4.4). These triggers ensure sporulation aligns with dispersal opportunities, contrasting with the rare, genetically diverse sexual cycle.

Sexual reproduction

Trichoderma reesei, the anamorph of the ascomycete Hypocrea jecorina, exhibits a heterothallic sexual reproductive system that requires compatible strains of opposite mating types, MAT1-1 and MAT1-2, for successful crossing. This sexual cycle was first demonstrated in 2009, when researchers induced fruiting body formation by pairing the MAT1-2 strain QM6a with a MAT1-1 strain (CBS999.97) derived from H. jecorina, overturning the long-held assumption that T. reesei was strictly asexual. In nature, the teleomorph H. jecorina undergoes sexual reproduction relatively infrequently compared to asexual conidiation, but evidence from wild isolates shows an equal distribution of both mating types (approximately 50% each across global populations), indicating occasional recombination events. Laboratory induction, however, is readily achieved under controlled conditions, such as on malt extract agar at 20–22°C with a 12-hour light-dark cycle, highlighting the potential for directed genetic manipulation. The sexual process begins with the confrontation of compatible hyphae from opposite mating types, leading to plasmogamy and the formation of ascogenous hyphae. These hyphae fuse, and stromata—compact masses of hyphae—develop on the substrate surface, typically decaying wood in natural settings where H. jecorina stromata have been observed. Within 7–10 days, perithecia (flask-shaped structures) embed in the stroma, where karyogamy occurs followed by meiosis in linear asci. Each ascus undergoes meiosis to produce four haploid nuclei, which then undergo two mitotic divisions, yielding 16 uninucleate ascospores that mature and are forcibly discharged upon perithecial maturation. Light is essential for stroma and perithecium development, with blue light signaling pathways regulating pheromone production and hyphal communication to initiate these stages. The primary outcome of sexual reproduction in T. reesei is increased through meiotic recombination, which shuffles alleles and can generate progeny with novel trait combinations, as evidenced by 50% inheritance rates of selectable markers in lab crosses. This contrasts with the clonal propagation via conidia, promoting adaptability in natural populations despite the predominance of modes. In the , this facilitates targeted for enhanced traits, though natural occurrences remain rare due to environmental constraints and the fungus's opportunistic ecology.

Genetics and strain engineering

Genome and genetic features

The genome of Trichoderma reesei strain QM6a, the reference wild-type isolate, was initially sequenced in 2008 using a whole-genome shotgun approach, yielding a draft assembly of approximately 34 Mb comprising 89 scaffolds and encoding 9,129 protein-coding genes, with an overall GC content of 52% and coding regions around 56.5%. A complete telomere-to-telomere assembly was published in 2017, spanning 34.9 Mb across 7 chromosomes with no gaps and 10,877 predicted protein-coding genes, reflecting an efficient genomic architecture adapted for lignocellulose degradation. A hallmark of the T. reesei is the organization of carbohydrate-active (CAZyme) genes into discrete clusters, totaling 316 CAZyme loci across 25 clusters ranging from 14 kb to 275 kb. Notably, these include seven major genes, such as the cellobiohydrolases encoded by cbh1 (cellobiohydrolase I) and cbh2 (cellobiohydrolase II), which account for over 60% of secreted cellulolytic activity under inducing conditions. Approximately 63% of these CAZymes are hydrolases (GHs) targeted at , underscoring the fungus's role as a premier producer. Gene regulation is tightly controlled, with carbon (CCR) mediated by the Cys₂His₂ Cre1, which binds GC-rich motifs in promoters to repress expression in the presence of readily metabolizable sugars like glucose. Comparative genomics between the wild-type QM6a and hyperproducing industrial strains, such as NG14 and RUT-C30, reveals genomic evolution driven by , including single-nucleotide variants, insertions/deletions, and larger structural changes affecting up to 125 loci in advanced mutants. These alterations disproportionately impact pathways for protein and vacuolar targeting, with about 17% of the affected genes in NG14 related to secretion machinery, facilitating higher extracellular yields without core gene modifications. Such adaptations highlight how industrial selection has amplified T. reesei's native secretory capacity, evolving from a saprotroph to an optimized biotechnological platform. Recent multi-omics studies as of 2025 continue to uncover regulatory networks enhancing production.

Strain improvement techniques

Strain improvement in Trichoderma reesei has primarily relied on classical techniques to generate hyper-producing , followed by advanced methods to achieve precise modifications. Early efforts in the involved to induce random mutations, leading to the development of key strains such as NG-14 and Rut-C30 from the parental strain QM6a. NG-14 was obtained through sequential steps, including UV exposure and chemical treatment with N-methyl-N'-nitro-N-nitrosoguanidine, resulting in enhanced production capabilities. Subsequent UV of NG-14 produced the hypersecretory Rut-C30, which features multiple genomic alterations affecting 43 genes compared to the wild-type, including single-nucleotide , insertions/deletions, and large deletions, enabling significantly improved protein secretion. These classical approaches, while effective for increasing overall enzymatic output, often introduce unintended mutations that require extensive screening to identify beneficial . Genetic engineering techniques have enabled more targeted strain optimization in T. reesei, with protoplast transformation serving as a foundational method for introducing foreign DNA. Protoplast-mediated transformation, developed in the 1980s, involves enzymatic removal of the cell wall to generate protoplasts, which are then fused with linear DNA for stable integration into the genome, facilitating heterologous gene expression. This technique has been widely adopted for overexpressing native or foreign genes, such as cellulases, by integrating constructs under strong promoters like cbh1, with transformation efficiencies reaching up to 1,000 transformants per microgram of DNA in optimized protocols. More recently, CRISPR-Cas9 has revolutionized precise genome editing in T. reesei, allowing efficient gene knockouts and modifications. For instance, CRISPR-Cas9 systems using multiple single-guide RNAs have been employed to disrupt the cre1 gene, which encodes a carbon catabolite repressor, thereby alleviating glucose repression and enhancing cellulase expression in strains like Rut-C30. These tools achieve high editing efficiencies, often exceeding 80% for single-gene knockouts, and support multiplex editing for simultaneous modifications. Contemporary strategies focus on refining regulatory elements and reproductive capabilities to further boost strain performance. Promoter engineering targets inducible systems, such as the native cbh1 promoter, which is strongly activated by inducers like sophorose, to drive controlled . Modifications, including deletion of repressor binding sites or fusion with synthetic elements, have strengthened this promoter's activity under diverse carbon sources, enabling up to twofold increases in target transcription without altering the native structure. Additionally, mating-type manipulation has unlocked sexual crossing for , as T. reesei was historically considered but possesses cryptic sexual cycles. By genetically switching the mating type locus (e.g., from MAT1-2 to MAT1-1 via ), researchers have enabled controlled between industrial strains, generating progeny with transgressive traits like improved growth rates or enzyme profiles. This approach, combined with fusion, facilitates the introgression of beneficial alleles while maintaining industrial compatibility. As of 2025, advances include identification of novel repressors like Rme1 and strategies for ultrahigh protein secretion exceeding 10 g/L.

Industrial and biotechnological applications

Enzyme production and mechanisms

Trichoderma reesei is renowned for its high-yield production of cellulolytic enzymes, primarily consisting of cellulases such as endoglucanases (e.g., EGI/Cel7B and EGII/Cel5A), exoglucanases (e.g., CBH1/Cel7A and CBH2/Cel6A), and β-glucosidases (e.g., BGL1/Cel3A), alongside hemicellulases like xylanases. These enzymes belong to various glycoside hydrolase (GH) families, with CBH1 classified in GH7 and CBH2 in GH6, enabling efficient biomass degradation. Industrial strains of T. reesei can secrete enzyme cocktails at concentrations exceeding 100 g/L under optimized conditions, making it a cornerstone for biotechnological applications. The production of these enzymes is tightly regulated at the transcriptional level, with induction primarily triggered by or its hydrolysis products like sophorose and , mediated by transcription factors such as Xyr1 and Ace2. Conversely, glucose exerts through the Cre1 regulator, which binds to promoter regions of cellulase genes to inhibit their expression in the presence of readily available sugars. This dual mechanism ensures that enzyme synthesis is activated only when complex are the carbon source, optimizing in the . Enzyme secretion in T. reesei follows the classical eukaryotic pathway, where N-terminal signal peptides direct nascent polypeptides to the () for translocation and initial folding, assisted by chaperones like BiP1 and (). From the , proteins traffic through the Golgi apparatus for further processing and sorting, culminating in via vesicles; the unfolded protein response regulator Hac1 plays a critical role in maintaining during high-level secretion. This pathway supports the massive extracellular accumulation of enzymes without compromising cellular integrity. The synergistic action of T. reesei cocktails is essential for lignocellulose , where endoglucanases randomly cleave internal β-1,4-glycosidic bonds to create nicks in chains, exoglucanases like CBH1 (GH7) and CBH2 (GH6) processively release from chain ends, and β-glucosidases hydrolyze to glucose, preventing product inhibition. Hemicellulases complement this by degrading hemicellulosic components, exposing fibrils for access, resulting in comprehensive that no single can achieve alone. This coordinated mechanism, refined through evolutionary adaptation, underpins the fungus's prowess as a and industrial producer.

Commercial uses and advancements

Trichoderma reesei plays a pivotal role in the biofuels industry, particularly in the production of , where its enzymes efficiently hydrolyze into fermentable sugars. This capability has made it a cornerstone for second-generation processes, enabling the conversion of agricultural residues and forestry waste into sustainable fuels. In the textile sector, enzymes from T. reesei are employed for biostoning and biopolishing, replacing harsh chemical treatments like stone washing with eco-friendly alternatives that soften fabrics and remove fuzz without damaging fibers. Additionally, its detergent-compatible enzymes enhance laundry performance by breaking down starch-based stains and improving fabric care in household and industrial cleaning products. In food processing, T. reesei-derived beta-glucanases are utilized to improve handling and quality in by degrading beta-glucans in , resulting in better texture and volume. Beyond native enzymes, T. reesei serves as a versatile host for recombinant protein , including fusion proteins at yields up to 19 g/L, and has been engineered to express ovalbumin for animal-free , as demonstrated in a 2021 study achieving sustainable via fungal . While primarily known for , its eukaryotic secretion machinery also supports the expression of therapeutic proteins, positioning it as a platform for vaccines and other biologics in filamentous fungi systems. Recent advancements have focused on co-cultures involving T. reesei to boost enzyme yields, with 2021 studies showing enhanced lignocellulolytic activities through synergistic interactions with fungi like Aspergillus niger, improving overall saccharification efficiency compared to monocultures. The global industrial enzymes market, heavily influenced by T. reesei contributions, reached approximately $8.5 billion in 2025. In 2025, efforts to engineer T. reesei as a "super-host" have advanced heterologous protein expression by addressing secretion limitations through promoter engineering, protease deletions, and process optimization, with perspectives for improved yields in non-native proteins.

References

  1. [1]
    Genome sequencing and analysis of the biomass-degrading fungus ...
    May 4, 2008 · Trichoderma reesei is the main industrial source of cellulases and hemicellulases used to depolymerize biomass to simple sugars that are ...
  2. [2]
    Trichoderma reesei - NCBI - NIH
    Molecular evidence that the asexual industrial fungus Trichoderma reesei is a clonal derivative of the ascomycete Hypocrea jecorina.Missing: history | Show results with:history
  3. [3]
    Tracking the roots of cellulase hyperproduction by the fungus ...
    Trichoderma reesei (teleomorph Hypocrea jecorina) is the main industrial ... reesei strain QM6a that was isolated from US Army tent canvas in 1944 in ...Results · Mutation Patterns In The Two... · Discussion
  4. [4]
    Fungal Cellulases | Chemical Reviews - ACS Publications
    Jan 28, 2015 · viride was eventually reclassified as T. reesei in honor of Elwyn Reese. (6) Postwar work at Natick by Reese and Mandels led to the modern ...<|control11|><|separator|>
  5. [5]
    Cellulases and beyond: the first 70 years of the enzyme producer ...
    Jun 10, 2016 · More than 70 years ago, the filamentous ascomycete Trichoderma reesei was isolated on the Solomon Islands due to its ability to degrade and thrive on cellulose ...Missing: taxonomy | Show results with:taxonomy
  6. [6]
    Trichoderma reesei Isolated From Austrian Soil With High Potential ...
    reesei is a heterothallic fungus with a common mating type structure of three mat-genes in the MAT1-1 locus and one mat-gene in the MAT1-2 locus (Seidl et al., ...
  7. [7]
    Enhancement of Cellulase Production in Trichoderma reesei via ...
    The filamentous fungus Trichoderma reesei is one of the most studied cellulolytic organisms and the major producer of cellulases for industrial applications.
  8. [8]
    The phenomenon of degeneration of industrial Trichoderma reesei ...
    Oct 1, 2021 · Strains of the genera Trichoderma reesei are used for large-scale production of cellulases, which are needed in food and feed, textile, paper ...
  9. [9]
    On the safety of Trichoderma reesei - ScienceDirect
    T. reesei is non-pathogenic for man and it has been shown not to produce fungal toxins or antibiotics under conditions used for enzyme production.
  10. [10]
    Industrial Relevance of Trichoderma reesei as an Enzyme Producer
    reesei's role in bio-ethanol production, it is used to produce industrial enzymes with a broad range of applications supporting the bio-based economy. To date ...
  11. [11]
    Hypocrea jecorina Berk. & Broome 1873 [1875] - Biota of NZ
    Jan 17, 2025 · Taxonomic and nomenclatural information for the scientific name: Hypocrea jecorina ... Synonym of Trichoderma reesei E.G. Simmons.
  12. [12]
    Trichoderma reesei Isolated From Austrian Soil With High Potential ...
    The teleomorph of T. reesei was initially known as Hypocrea jecorina, a name which was frequently used in literature of this species as well, until the one ...
  13. [13]
    Trichoderma reesei - Leibniz Institute DSMZ
    ATCC 13631,CCM F-560,CMI 045548,IMI 045548,CBS 383.78. Isolated from: cotton duck shelter. Country: Solomon Islands Bougainville Island. Date of sampling: 1944.
  14. [14]
    Tracking the roots of cellulase hyperproduction by the ... - PNAS
    reesei strain QM6a that was isolated from US Army tent canvas in 1944 in the Solomon Islands (5). Despite these years of research and development in the area, ...
  15. [15]
    Remembering Mary (1917 to 2008) - PubMed Central - NIH
    Sep 1, 2009 · The fungus was identified as Trichoderma 'viride' strain QM 6a, which was later renamed Trichoderma reesei QM 6a, in honour of its discoverer.
  16. [16]
    Use of Cellulases from Trichoderma reesei in the Twenty-First ...
    The two first energy crises in 1973 and 1979 raised in western countries the question of fossil fuel costs and dependence on volatile middle-east politics.Chapter 17 - Use Of... · Overview Of The Global... · Current Applications
  17. [17]
    Fermentation of trichoderma reesei and apparatus therefor
    The method comprises fermenting the Trichoderma reesei at a temperature of between about 26° C. and 31° C.<|control11|><|separator|>
  18. [18]
    Trichoderma reesei Simmons - 56766 - ATCC
    On PDA medium at 25°C after 21 days, mycelium green, powdery to velutinous. Reverse bright yellow. Hyphae hyaline, guttulate. Conidia green, ellipsoidal ...
  19. [19]
    Morphology and enzyme production of Trichoderma reesei Rut C-30 ...
    The filamentous fungus Trichoderma reesei is one of the main microorganisms for industrial production of biomass degrading enzymes and in academic research ( ...Missing: Reuben Reese
  20. [20]
    The micromorphology of Trichoderma reesei analyzed in cultivations ...
    Aug 9, 2016 · Because of the strong influence on the enzyme production, the morphology of the filamentous fungi is a key parameter for process optimization.Missing: conidia phialides
  21. [21]
  22. [22]
    The clear zone of Aspergillus niger (right) and Trichoderma reesei...
    A. niger and T. reesei grew in CMC solid medium form a clear zone around colony that indicated the cellulolytic activity.
  23. [23]
    Cellulase Enzyme Production from Filamentous Fungi Trichoderma ...
    Conidiophores were erect and formed in variable branches. Phialides were lageniform in shape and more or less ampulliform (Figure 2). All these features showed ...
  24. [24]
    Trichoderma diversity from karst area in Yunnan, Shilin, and four ...
    Aug 8, 2025 · For microscopic morphology, including hyphae, conidiophores, phialides, conidia, and other structures, images were taken using an Olympus BX51 ...
  25. [25]
    Characterization and Strain Improvement of a Hypercellulytic Variant ...
    Aug 28, 2016 · Trichoderma reesei is a cellulolytic filamentous fungus originally isolated in the Solomon Islands during World War II. T. reesei QM9414 and RUT ...<|control11|><|separator|>
  26. [26]
    [PDF] TRICHODERMA: THE GENOMICS OF OPPORTUNISTIC SUCCESS
    Oct 9, 2011 · Trichoderma: the genomics of opportunistic success. Permalink https://escholarship.org/uc/item/4hc5r81s. Author. Druzhinina, Irina S.
  27. [27]
    Ecological functions of Trichoderma spp. and their secondary ...
    Feb 22, 2016 · Trichoderma species have been used to suppress plant diseases and the growth of pathogens in contact with plant tissues or in terms of pathogen ...
  28. [28]
    [PDF] Cellulolytic enzyme production and response to pH and temperature ...
    The best performance was seen with fermentations conducted at pH 4.5 or 6.0 and temperature of 23˚C.
  29. [29]
    [PDF] Optimization of Solid State Fermentation Conditions for the ...
    Effect of pH on enzyme activity​​ The enzyme exhibited 82% activity at pH 4, 100% activity at pH 5 and 83% activity at pH 6, after 24 hrs of incubation. Solid ...
  30. [30]
    The Longibrachiatum Clade of Trichoderma: a revision with new ...
    Feb 5, 2012 · ... conidia also formed from phialides formed along hyphae submerged in the agar. Conidia slowly turning pale green, held in wet heads.
  31. [31]
    Expression of Biomass-Degrading Enzymes Is a Major Event during ...
    To investigate the transcriptional changes taking place during the transition from growing hyphae to conidia in Trichoderma reesei, microarray experiments were ...
  32. [32]
    Conidiation in the filamentous fungus Trichoderma - ResearchGate
    Aug 6, 2025 · Five stages of conidiation can be described: vertical hyphae appear and then branch forming conidiophores, phialides develop on the ...
  33. [33]
    Trichoderma in the light of day – Physiology and development - PMC
    ... filamentous fungus Hypocreajecorina (anamorph Trichoderma reesei) BMC Microbiol. ... Sexual development in the industrial workhorse Trichoderma reesei.
  34. [34]
    Sexual development in the industrial workhorse Trichoderma reesei | PNAS
    ### Summary of Sexual Reproduction in Trichoderma reesei
  35. [35]
    Trichoderma reesei meiosis generates segmentally aneuploid ...
    Feb 25, 2015 · Hypocrea jecorina is the sexual form of the industrial workhorse fungus Trichoderma reesei that secretes cellulases and hemicellulases to ...Missing: stroma | Show results with:stroma
  36. [36]
    Whole-genome metabolic model of Trichoderma reesei built by ...
    Nov 21, 2016 · The amount of deoxyribonucleotides in DNA was calculated based on the published GC content of the genome, 52.0% [56]. The amount of RNA in ...
  37. [37]
    Trichoderma reesei complete genome sequence, repeat-induced ...
    Trichoderma reesei undergoes a heterothallic reproductive cycle and generates fruiting bodies (stromata) containing linear asci with 16 ascospores [7]. These 16 ...Missing: shape | Show results with:shape
  38. [38]
    Crel, the carbon catabolite repressor protein from Trichoderma reesei
    Nov 27, 1995 · The presence of the crel translation product was demonstrated with polyclonal antibodies against Cre1, which identified a protein of 43 (±2) kDa ...<|control11|><|separator|>
  39. [39]
    Comparative Genomics Analysis of Trichoderma reesei Strains
    Dec 4, 2013 · The genome of the wild type strain QM6a was sequenced by the conventional method in 2008. The comparison of genome sequences between the hyper- ...<|control11|><|separator|>
  40. [40]
    Enhancement of cellulase production in Trichoderma reesei RUT ...
    May 10, 2019 · The well-known hyper-cellulolytic mutant RUT-C30 (ATCC 56765) was also isolated from NG14 using UV mutagenesis (Fig. 1a). To characterize the ...
  41. [41]
    An overview on current molecular tools for heterologous gene ...
    Oct 26, 2021 · In this review, we summarize and discuss several aspects involved in heterologous gene expression in Trichoderma, including transformation methods, genome ...
  42. [42]
    Transformation of Trichoderma reesei with a constitutively ...
    The transformation frequency was high (897–1,421 transformants μg−1 DNA) and up to 70% of the stable transformants exhibited extracellular chitinase activity.
  43. [43]
    An efficient CRISPR/Cas9 genome editing system based on a ...
    Feb 11, 2024 · Truncation of the transcriptional repressor protein Cre1 in Trichoderma reesei Rut-C30 turns it into an activator. Fungal Biol Biotechnol ...
  44. [44]
    Efficient genome editing in filamentous fungus Trichoderma reesei ...
    Our results illustrate that the CRISPR/Cas9 system is a powerful genome-manipulating tool for T. reesei and most likely for other filamentous fungal species, ...Missing: cre1 knockout
  45. [45]
    Engineering the cbh1 Promoter of Trichoderma reesei for Enhanced ...
    Jan 14, 2020 · The strong and inducible cbh1 promoter is most widely used to express heterologous proteins, useful in food and feed industries, in Trichoderma reesei.<|control11|><|separator|>
  46. [46]
    Sexual development in the industrial workhorse Trichoderma reesei
    Aug 18, 2009 · Using gene sequence analysis, Kuhls et al. (7) found that T. reesei is indistinguishable from the the pantropical ascomycete Hypocrea jecorina.
  47. [47]
    Transgressive phenotypes from outbreeding between the ...
    Aug 20, 2024 · Trichoderma reesei, the main filamentous fungus used for industrial cellulase production, was long considered to be asexual.<|control11|><|separator|>
  48. [48]
  49. [49]
  50. [50]
  51. [51]
    Systems biological approaches towards understanding cellulase ...
    The Sordariomycete Trichoderma reesei (teleomorph Hypocrea jecorina) is most widely used for the industrial production of cellulolytic and hemicellulolytic ...5. The T. Reesei... · 6. The T. Reesei Secretome · 7. The T. Reesei Metabolome
  52. [52]
  53. [53]
    Cellulases: From Bioactivity to a Variety of Industrial Applications
    Jul 5, 2021 · Cellulases have been utilized in various commercial sectors including agriculture, brewing, laundry, pulp and paper and textile industry.
  54. [54]
    Microbial cellulase production and its potential application for textile ...
    Apr 10, 2023 · Trichoderma viride and Trichoderma reesei, for example, support cellulase formation in suitable conditions, such as solid and submerged ...Strain Improvement Of... · Biostone Washing · Biopolishing And Finishing<|control11|><|separator|>
  55. [55]
    EPA Issues Final Rule to Reduce Regulatory Burden for ...
    Mar 5, 2020 · Trichoderma reesei is used by the animal feed, baking, beverages, textile processing, detergent, pulp and paper, industrial chemicals and ...
  56. [56]
    Ovalbumin production using Trichoderma reesei culture and low ...
    Dec 16, 2021 · Ovalbumin (OVA) produced using the fungus Trichoderma reesei (Tr-OVA) could become a sustainable replacement for chicken egg white protein powder.Missing: recombinant insulin vaccines
  57. [57]
    Protein Expression Trichoderma Reesei | Bio Tech Resources
    Trichoderma reesei (T. reesei), recently re-classified as Hypocrea jecorina, a mesophilic filamentous fungus, is an attractive expression host for the ...
  58. [58]
    An overview of Trichoderma reesei co-cultures for the production of ...
    Apr 7, 2021 · Several reports have demonstrated that T. reesei co-cultures present higher enzyme activities compared with its monocultures, especially in the ...Missing: enhanced | Show results with:enhanced
  59. [59]
    Industrial Enzymes Market Size to Cross USD 8.46 Billion in 2025
    Jul 31, 2025 · The industrial enzymes market size is projected to grow from USD 8.46 billion in 2025 to USD 16.09 billion by 2034, at a CAGR of 7.4% during ...Missing: Trichoderma reesei
  60. [60]
    Unlocking the Potential of Trichoderma reesei as a Super-Host for ...
    Unlocking the Potential of Trichoderma reesei as a Super-Host for Heterologous Protein Production: Challenges, Advances, and Perspectives. Biotechnol J. 2025 ...