Mycoparasitism

A mycoparasite is an organism with the ability to be a parasite to fungi. A variety of fungicolous fungi have been found in nature, either as parasites, commensals or saprobes.[1] Biotrophic mycoparasites get nutrients from living host cells. Necrotrophic mycoparasites rely on decayed matter (saprophytic growth).[1][2]

Types of mycoparasitism

Biotrophic mycoparasites and nectrophic mycoparasites are two main groups of mycoparasite.[3]

Biotrophic and necrotrophic mycoparasites

Biotrophic mycoparasites get nutrients from living host cells and their growth of these parasites is greatly influenced by the metabolism of the host.[4] Biotrophic mycoparasites tend to have a high host specificity, and often form specialized infection structures or a host parasite transition.[5] Necrotrophic mycoparasites could only rely on saprophytic growth.[2] On the other hand, the antagonistic action is strongly aggressive in necrotrophic relationships which is dominated by necrotrophic mycoparasites. Necrotrophic parasites tend to have a low host specificity, and are relatively unspecialized in their mechanism of parasitism.[5]

Balanced and destructive mycoparasites

Balanced mycoparsites have little or no destructive effect on the host, whereas destructive mycoparasites have the opposite effect.[6] Biotrophic mycoparasites are generally considered as balanced mycoparasites; Necrotrophic mycoparasites usually have low host specificity and could use toxic and related enzyme to kill host, therefore necrotrophic mycoparasites are usually considered as destructive mycoparasites. However, in some combinations, the parasite may live during its early development as a biotrophic parasite, then kill their host and change into destructive mycoparasites in their late stage of parasitization.[4][6]

Myco-heterotrophy

Monotropa uniflora (or other angiosperm species), which are parasitic on fungus and do not photosynthesize, are sometimes classified as mycoparasites, however they should be referred to as myco-heterotrophs.

Mechanism

There are normally four continuous steps of mycoparasitism: target location; recognition; contact and penetration; nutrient acquisition.[7]

Target location

Many research indicate that the growth direction of mycelium, spore germination, and bud tube elongation of mycoparasites have tropic reaction or tropism before attaching the host fungi.[8] This tropic reaction is resulted from chemical stimulants secreted from mycohost and the direction of concentration gradient determines the growth direction of the parasite.[9] As the mycoparasitic interaction is host-specific and not merely a contact response, it is likely that signals from the host fungus are recognized by mycoparasites such as Trichoderma and provoke transcription of mycoparasitism-related genes.[10][11]

Recognition

When mycoparasites get attachment of mycohost, they will recognize each other. It is indicated that the recognition between mycoparasites and their host fungi is related to the agglutinin on the cell surface of the mycohost. The carbohydrate residues on the cell wall of mycoparasites could specifically bind to the lectins on the surface of the host fungi to achieve mutual recognition.[12]

Contact and penetration

Once mycoparasites and mycohost recognize each other, both of them would have some changes in external form and internal structure to some extents.[13][14] The manifestations of the mycoparasitic fungi are usually as follows: (1) the hypha grows rapidly on the host fungi and coils around the hypha of the host fungi; (2) the hypha penetrates and elongates the hypha of the host fungi (3) the infection filament is produced and penetrates the cells of host fungi to build parasitic relationship.[15] In terms of necrotrophic/destructive mycoparasites, on the other hand, the hyphae of host fungi, due to the role of mycoparasites, will stop the growth, deform, shrivel and even dissolve.[2]

Application

Recently, it is popular to use integrated pest management (IPM) as an approach to reduce the amount of pesticides used, through preventive cultural practices, the use of disease resistant plant cultivars, and the use of mechanical and biological control of pathogen populations. Biocontrol of plant pathogens by microbial antagonists is one promising component in future disease control strategies that is compatible with both organic agriculture and IPM.[16] As some mycoparasitic fungi have the ability to parasite and kill other fungi, these fungi could be used to antagonize plant pathogenic fungi directly as biocontrol agent. Mycoparasitism of plant pathogenic fungi by Trichoderma isolates has been well researched and is widely considered to be a major contributing factor to the biocontrol of a range of commercially important diseases.[7] In, among others, the United States, India, Israel, New Zealand, and Sweden have applied Trichoderma species commercially such as Rhizoctonia solani, Botrytis cinerea, Sclerotium rolfsii, Sclerotinia sclerotiorum, Pythium spp., and Fusarium spp. as a promising alternative to chemical pesticides.[17][18] Moreover, with the better understanding of mycoparasitism, more bioactive compounds including biopesticides and biofertilizers would be introduced as beneficial products.[19]

List of fungal bioagents with their trade and manufacturers name[5]
Commercial products Bioagents used Name of the manufacturer
AQ10 biofungicide Ampelomyces quisqualis

isolate M-10

 Ecogen, Inc. Israel
Anti-Fungus Trichoderma spp. Grondortsmettingen De Cuester, Belgium
Biofungus Trichoderma spp. Grondortsmettingen De

Cuester n. V.Belgium

Bas-derma Trichoderma viride Basarass Biocontrol Res.

Lab., India

Binab T Trichoderma harzianum

(ATCC 20476) and

Trichoderma polysporum

(ATCC 20475)

 Bio-Innovation AB, UK
Bioderma Trichoderma viride/T. harzianum Biotech International Ltd., India
Biofox C Fusarium oxysporum (Non- pathogenic) S. I. A. P. A., Italy
Prestop, Prirnastop Gliocladium catenulatum Kemira Agro. Oy, Finland
Root Pro, Root Prota to Soilgard Trichoderma harzianum/Gliocladium virens strain

GL-21

 Efal Agr, Israel Thermo Trilogy, USA
Root shield, Plant shield,

T-22 Planter box

 Trichoderma harzianum Rifai strain KRL-AG

(T-22)

 Bioworks Inc., USA
Supresivit Trichoderma harzianum Borregaard and Reitzel, Czech Republic
T-22 G, T-22 HB Trichoderma harzianum

strain KRL-AG2

 THT Inc., USA
Trichodex, Trichopel Trichoderma harzianum Makhteshim Chemical Works Ltd., USA
Trichopel, Trichoject, Trichodowels, Trichoseal Trichoderma harzianum

and Trichoderma viride

 Agrimm Technologies Ltd., New Zealand
Trichopel Trichoderma harzianumand Trichoderma viride Agrimm Technologies Ltd., New Zealand
Trichoderma 2000 Trichoderma sp. Myocontrol Ltd., Israel
Tri-control Trichoderma spp. Jeypee Biotechs, India
Trieco Trichoderma viride Ecosense Labs Pvt. Ltd.,

Mumbai, India

TY Trichoderma sp. Mycocontrol, Israel

References
  1. Hawksworth, D.L.; Kirk, P.M.; Sutton, B.C.; Pegler, D.N. (1995). Ainsworth & Bisby's Dictionary of the Fungi. Wallingford, UK: CAB International.
  2. Barnett, H.L. (1963). "The nature of mycoparasitism by fungi". Annu. Rev. Microbiol. 17: 1–14. doi:10.1146/annurev.mi.17.100163.000245.
  3. Boosalis, M G (1964). "Hyperparasitism". Annual Review of Phytopathology. 2 (1): 363–376. doi:10.1146/annurev.py.02.090164.002051. ISSN 0066-4286.
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  5. Ashraf, Shabbir; Zuhaib, Mohammad (2013), "Fungal Biodiversity: A Potential Tool in Plant Disease Management", Management of Microbial Resources in the Environment, Springer Netherlands, pp. 69–90, doi:10.1007/978-94-007-5931-2_4, ISBN 9789400759305
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  10. Druzhinina, Irina S.; Seidl-Seiboth, Verena; Herrera-Estrella, Alfredo; Horwitz, Benjamin A.; Kenerley, Charles M.; Monte, Enrique; Mukherjee, Prasun K.; Zeilinger, Susanne; Grigoriev, Igor V. (2011-09-16). "Trichoderma: the genomics of opportunistic success" (PDF). Nature Reviews Microbiology. 9 (10): 749–759. doi:10.1038/nrmicro2637. ISSN 1740-1526. PMID 21921934.
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  12. Inbar, Jacob; Menendez, Ana; Chet, Ilan (1996). "Hyphal interaction between Trichoderma harzianum and Sclerotinia sclerotiorum and its role in biological control". Soil Biology and Biochemistry. 28 (6): 757–763. doi:10.1016/0038-0717(96)00010-7. ISSN 0038-0717.
  13. Zeilinger, Susanne; Brunner, Kurt; Peterbauer, Clemens K.; Mach, Robert L.; Kubicek, Christian P.; Lorito, Matteo (2003-07-01). "The Nag1 N-acetylglucosaminidase of Trichoderma atroviride is essential for chitinase induction by chitin and of major relevance to biocontrol". Current Genetics. 43 (4): 289–295. doi:10.1007/s00294-003-0399-y. ISSN 0172-8083. PMID 12748812.
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