TASK 2 JIB101

ASSIGNMENT 2 DISCUSS THE BENEFIT OF MYCORRHIZA TO SOIL.

Mycorrhiza y Definition of Mycorrhiza: symbiotic/ mutualistic associations that form between the roots of most plant species and fungi. y y y Greek: myco= fungus; rhiza= root. Described in 1885 by the German forest pathologist A.B. Frank. Vast majority of land plants form symbiotic associations with fungi: ~ 95% of all plant species. y Characterized by bi-directional movement of nutrients where C fungus and inorganic nutrients y plant: providing a critical linkage between plant root and soil.

In infertile soils, nutrients taken up by the mycorrhizal fungi can lead to improved plant growth (agronomic) and reproduction (ecology). As a result, mycorrhizal plants are often more competitive and better able to tolerate environmental stresses than are non mycorrhizal plants.

y y

Mycorrhizal fungi usually proliferate both in the root and in the soil. The soil borne or extrametrical hyphae take up nutrients from the soil solution and transport them to the root. By this mechanism, mycorrhizae increase the effective absorptive surface area of the plant.

The best known benefit of mycorrhizal inoculation is the plant growth response; not always the most important benefit.

Real benefits may be improved soil structure, protection from pathogens, and in a restored plant community, increased seedling survivorship, higher species diversity, and greater resistance to invasion by weeds.

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What does fungus get from plant? y sugars (monosaccharides) that are transported from the plant to the fungus and converted into trehalose, mannitol & glycogen, which are generally not permeable back across fungal cell membrane y y B vitamins; stimulate spore germination. Estimated that trees invest 10% of photosynthesis to their mycobionts.

What does plant get from fungus? y y y y P, Ca, K, Cu, Mo, Mg, Zn nitrogen water: source-sink and water potential protection against pathogens

How do mycorrhizal relationships form? y Mycorrhizal associations start when soil hyphae respond to the presence of a root by growing towards it, establishing contact and growing along its surface; root exudates promotes spore germination and growth. y y One or more hyphae produce swellings called appressoria between epidermal cells. Root penetration occurs when hyphae from the appressoria penetrate epidermal or cortical cells to enter the root. These hyphae cross the hypodermis and start branching in the outer cortex; form hartig net / fungal mantel. y y extension of hyphae from mantel into soil Unique root growth and morphology i. ii. iii. iv. fungus secretes auxins, gibberellins, & cytokines enhances root growth maintains juvenile roots; no root hairs mycorrhizal root tip is more often branched than non mycorrhizal root tip, e.g., dichotomous branching in Pinaceae y However, it is important to distinguish between specificity, innate ability to colonize, infectiveness, amount of colonization, and effectiveness, plant response to colonization. AM fungi differ widely in the level of colonization they produce in a root system and in their impact on nutrient uptake and plant growth.

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(Vesicular) Arbuscular Mycorrhiza A. Endomycorrhiza y The most common: arbuscular mycorrhiza/ vesicular-arbuscular mycorrhiza(AM/ VAM); found on grasses, most crop plants, many trees, shrubs, flowers, and ~80% world's plant species. y The diagnostic feature of AM is the development of a highly branched arbusculewithin root cortical cells. y The fungus initially grows between cortical cells, but soon penetrates the host cell wall and grows within the cell. In this association neither the fungal cell wall nor the host cell membrane are breached. As the fungus grows, the host cell membrane invaginates and envelops the fungus, creating a new compartment where material of high molecular complexity is deposited. y This apoplastic space prevents direct contact between the plant and fungus cytoplasm and allows for efficient transfer of nutrients between the symbionts. y VAM was originally applied to symbiotic associations formed by all fungi in the Zygomycete order Glomales, but because a major suborder lacks the ability to form vesicles in roots, AM is now the preferred acronym. The order Glomalesis further divided into families and genera according to the method of spore formation. y The arbuscules are relatively short lived, less than 15 days, and are often difficult to see in field-collected samples. y Other structures produced by some AM fungi:1. Vesicles: thin-walled, lipid-filled structures that usually form in intercellular spaces. Their primary function: storage; can also serve as reproductive propagulesfor the fungus. 2. Auxiliary cells: formed in the soil and can be coiled or knobby. The function: unknown. Reproductive spores can be formed either in the root or more commonly in the soil. 3. Spores: produced by fungi forming AM associations are asexual, forming by the differentiation of vegetative hyphae. For some fungi (e.g., Glomusintraradices), vesicles in the root undergo secondary thickening, and a septum (cross wall) is

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laid down across the hyphal attachment leading to spore formation, but more often spores develop in the soil from hyphal swellings. y The spores of AM fungi are very distinctive. Diameter: 10 mm for Glomus tenueto more than 1m for some Scutellospora spp. Spores vary in colour (hyaline black) and in surface texture (smooth-highly ornamented). Glomus forms spores on the ends of hyphae, Acaulospora forms spores laterally from the neck of a swollen hyphal terminus, and Entrophospora forms spores within the neck of the hyphal terminus. y The AM type of symbiosis is very common as the fungi involved can colonize a vast taxonomic range of both herbaceous and woody plants, indicating a general lack of host specificity among this type.

B. Ericaceous Mycorrhiza y The term ericaceous is applied to mycorrhizal associations found on plants in the order Ericales. y y No arbuscules are formed. On fine root system; extensive hyphal development around root; slight development on root surface; within cells extensive hyphal coils. y Three major forms of ericaceous mycorrhiza have been described: i. Ericoid Cells of the inner cortex become packed with fungal hyphae. A loose welt of hyphaegrows over the root surface, but a true mantle is not formed. and Plants -Calluna(heather), and

Rhododendron(azaleas

rhododendrons)

Vaccinium(blueberries) that have very fine root systems and typically grow in acid, peaty soils. Fungi -

ascomycetes(Hymenoscyphus). ii. Arbutoid Intracellular penetration can occur, a mantle forms, and a Hartig net is present. Plants: Arbutus (eg. Pacific madrone), Arctostaphylos (eg. bearberry), and Pyrolaceae spp. Fungi -

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basidiomycetes. Epidermal cells are colonized by hyphae that form coils; coils disintegrate but cells dont; may be colonized again. iii. Monotropoid The fungi colonize a chlorophyllous (lacking chlorophyll) plants in Monotropaceae (e.g., Indian pipe), producing the Hartig net and mantle. The same fungi also form ECM associations with trees and thereby form a link through which carbon and other nutrients can flow from the autotrophic host plant to the heterotrophic, parasitic plant.

C. Orchidaceous Mycorrhiza y y y y orchid seeds and young plants associated with Rhizoctonia seeds and seedlings apparently lack some enzymatic machinery; can't metabolize sulfur Many orchids are achlorophyllous; get carbon from fungus. Rhizoctoniais normally a potent pathogen of plant; often considered to be captured by the orchid; orchids produce many secondary products that may inhibit fungus. y y Dont have any major effect on root morphology; form loops in root cells. Orchid mycorrhizasare unique in that they are required for seed germination. Some kinds of orchids never photosynthesize, but instead parasitize the mycorrhizal fungi. y Orchids typically have very small seeds with little nutrient reserve. The plant becomes colonized shortly after germination, and the mycorrhizal fungus supplies carbon and vitamins to the developing embryo. y For achlorophyllous species, the plant depends on the fungal partner to supply carbon throughout its life. The fungus grows into the plant cell, invaginated the cell membrane and forming hyphal coils within the cell. These coils are active for only a few days, after which they lose turgor and degenerate and the nutrient contents are absorbed by the developing orchid. y Fungi participating in the symbiosis are basidiomycetes similar to those involved in decaying wood (e.g., Coriolus, Fomes, Marasmius) and pathogenesis (e.g., Armillariaand

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Rhizoctonia). In mature orchids, mycorrhizae also have roles in nutrient uptake and translocation. D. Ectomycorrhiza y y y Hyphae do not penetrate cells of cortex in root tips Next most common kind after VAM. The host plants are pines, firs, spruce, oaks, and several otherkinds of plants, mostly forest trees (Gymnosperm or Angiosperm). The fungi often form mushrooms or truffles. e.g. Amanita, Russula, Lactarius, Laccaria, Cortinarius, Inocybe, Boletusspp. 2000 spp. of plants; 5000 spp. Fungi y y y y y y Hartig net: growth of hyphaein between and around cortex cells Form fungal mantle; only form in actively growing roots Most ECM fungi unable to degrade cellulose Obligate vs. facultative; enhanced by high light, decreased by high N & P levels Ectomycorrhiza(ECM) = dynamic process Succession: individual mycorrhiza active for 1-3 yrs; root may out grow fungus and be colonized by a alternative ectomycorrhizal fungus or a more aggressive ectomycorrhizal fungus may replace the previous. y y y Ectendomycorrhiza: characteristics of ectosbut exhibit intracellular penetration MRAs(Mycelium radicisatrovirens); dark septatemycorrhiza; ascomycetes Most plants with ECM have roots with a modified lateral root branching pattern. This pattern, which is called heterorhizy, consists of short mycorrhizal lateral roots (called short roots) supported by a network of long roots. y The long and short roots in heterorhizic root systems are fundamentally similar in structure, but short roots normally grow much more slowly than long roots. Hyphae contact, recognize and adhere to root epidermal cells near the apex of young, actively growing, high-order, lateral root.

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Importance of mycorrhiza
Global perspective y Ericaceousplants, that dominate the acidic, high-organic heathlandsoils of subarcticand subalpineregions, are colonized by a group of ascomycetous fungi. y Moving down the environmental gradient coniferous trees as the dominant vegetation. These trees are colonized by a wide range of mostly basidiomycetous fungi. These hyphae function in the absorption and translocation of inorganic nutrients and water, but also release nutrients from litter layers by production of enzymes involved in mineralization of organic matter. y At the warmer and drier end of the environmental gradient, grasslands often form the dominant vegetation. Here, nutrient use is high and phosphorus is frequently a limiting element for growth. y Grasses and a wide variety of other plants are colonized by fungi belonging to the order Glomales. These fungi form arbuscules and extensive extrametrical hyphae and can significantly increasephosphorus-inflowrates of the plants they colonize.

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Benefits to plants
y Increased plant nutrient supply: extending the volume of soil accessible to plants; acquiring nutrient forms that would not normally be available to plants. y y Greater yield, nutrient accumulation, and/or reproductive success. Some ECM and ericoid fungi have the capacity to breakdown phenolic compounds in soils which can interfere with nutrient uptake. y Root colonization by ECM and VAM fungi can provide protection from parasitic fungi and nematodes. y y Suppression of competing non-host plants, bymycorrhizal fungi has been observed Significant amounts of C transfer through ECM connecting different plant species has been measured reduce competition between plants and contribute to the stability and diversity of ecosystems. y Networks of hyphae supported by dominant trees may help seedlings become established or contribute to the growth of shaded under storey plants. y y Nutrient transfer from dead to living plants can occur. Non-nutritional benefits to plants due to changes in water relations, phytohormone levels, carbon assimilation, etc. have been reported, but are difficult to interpret. y C fluxes-up regulation of the rate of net photosynthesis of host

Uptake and Transfer of soil nutrients

y When a nutrient is deficient in soil solution, the critical root parameter controlling its uptake is surface area.

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y The absorbing surface area of the root is greatly increase by the distribution and function of the fungi extrametrical hyphae. y To be effective in nutrient uptake, the hyphae must be distributed beyond the nutrient depletion zone around the root. For a poorly-mobile ion such as P, Zn, Cu, a narrow depletion zone develops close to the root. Hyphae can readily bridge this zone and grow into soil with an adequate supply of phosphorus. y Another factor contributing to the effective absorption of nutrients by mycorrhizae is their narrow diameter relative to roots. Furthermore, narrow hyphae can grow into small soil pores inaccessible to roots or even root hairs. y Some EM fungi produce large quantities of oxalic acid, and this may partially explain enhanced nutrient uptake by EM roots. y Another mechanism by which mycorrhizal fungi release inorganic phosphorus is through mineralization of organic matter. This occurs by phosphatase-mediated hydrolysis of organic phosphate (C-O-P) ester bonds. Significant phosphatase activity has been documented for mycorrhizal fungi grown in pure cultures and for excised and intact EM short roots. y Ericoid and EM have a special role in the mineralization of nitrogen. Most plant litter entering the soil has a high C:N ratio and is rich in lignin and tannins. Only a few mycorrhizal fungi can mobilize nutrients from these primary sources. However, a wide range of ericoid and EM fungi can obtain nitrogen and other nutrients from secondary sources of organic matter such as dead microbial biomass. A wide range of hydrolytic and oxidative enzymes capable of depolymerizing organic nitrogen have been demonstrated. These types of mycorrhizae may have an important role in nitrogen cycling in the acidic and highly organic soils where they predominate.

Sustainable Plant - Soil Ecosystems Roles in ecosystem:

y Soil hyphae are likely to have an important role in nutrient cycling by helping to prevent losses from the system, especially at times when roots are inactive. y Epigeous and hypogeous sporocarps of ECM and VAM fungi are important food sources for placental and marsupial mammals.

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y Mycorrhizal roots and fungus hyphae/ fruit bodies are important as food sources and habitats for invertebrates. y Mycorrhizas influence soil microbial populations and exudates in the mycorrhiza sphere and hyphosphere. y Contribution to soil structure; simple entanglement microaggregation (glycoprotein secretions such as glomalin by AM) soil aggregate formation of stable soil

aggregates. Typically ~1 to 20 m of AM hyphae g soil. y Mycorrhizal fungi contribute to carbon storage (nutrient reserve) in soil by altering the quality and quantity of soil organic matter.

Importance to human
y y y ECM fungi are economically and nutritionally important as human food resources. As medicines and natural dyes. Aesthetic values and an important part of the as culture, folklore and appreciation of nature by many people. y y Fungal diversity is a bio-indicator of environmental quality. Fungi which have adapted to local soil conditions are required for agriculture, horticulture and forestry.

Conclusion
Mycorrhiza is mutually beneficial relationships between such fungi and plant roots. Mycorrhizal fungi, can deliver phosphate ions and other minerals to plants, which the plants themselves cannot acquire on their own. In exchange, the plants supply the fungi with the organic nutrients. There are several different types of mychorrhiza. Ectomycorrhiza, which fungal cells form sheaths of hyphae over the surface of a root and also grow into the extracellular spaces of the root cortex. Endomychorriza, which is the fungal cells extend their hyphae through the root cell wall and into tubes formed by invagination or pushing inward of the root cell membrane.

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LIST OF REFERENCE BOOKS
1. Eldra P. Solomon, Linda R. Berg and Diana W. Martin (2002). BIOLOGY ( 6th Edition) BROOKS/COLE. 2. Neil A. Campbell and Jane B. Reece (2005) INTERNATIONAL EDITION, BIOLOGY (7th Edition). PEARSON BENJAMIN CUMMINGS 3. Gerald Karp (2007). CELL AND MOLECULAR BIOLOGY (CONCEPT AND EXPERIMENT) (5th Edition). WILEY INTERNATIONAL STUDENT VERSION. 4. Kenneth R. Miller and Joseph S. Levine (2002). PRENTICE HALL BIOLOGY. Pearson Education, Inc., Upper Saddle River, New Jersey.

LIST OF REFERENCE WEBSITES.

1. http://www.mycorrhiza.net/structure.htm http://cropsoil.psu.edu/sylvia/mycorrhiza.htm http://cropsoil.psu.edu/sylvia/mycorrhiza.htm http://Verdant Horticulture - Horticulture 101 - Mycorrhiza The Secret Root System.mht

2. 3. 4.

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