International Journal of Agricultural Science and Research (IJASR) ISSN 2250-0057 Vol.

3, Issue 1, Mar 2013, 17-24 TJPRC Pvt. Ltd.

REVIEW ON ROLE OF VAM FUNGI IN CROP PLANT-SOIL SYSTEM

AMEETA SHARMA1 & SAVITA YADAV2
1 2

Sr. Assistant Professor, Department of Biotechnology, The IIS University, Jaipur, Rajasthan, India

Dissertation Scholar, Department of Biotechnology, The IIS University, ICG Campus, Jaipur, Rajasthan, India

ABSTRACT
VA Mycorrhizal fungi occur in nearly all soils on earth and form a symbiotic relationship with roots of most terrestrial plants and have a wide range of roles. This review discusses the diversity of interactions between mycorrhizal fungi and plants, its occurrence, distribution, significance in environmentally stressed soil conditions such as drought, water logging and salinity. The improved nutrient absorption by mycorrhiza plants especially phosphorus under drought conditions have had a direct impact on growth parameters. Although VA mycorrhizal fungi do not show any host specificity, there is increasing evidence that various climatic and edaphic environmental factors influence their occurrence, taxonomic distribution and effectiveness.VAM fungi do have a potential to enhance plant growth but further studies are needed with additional plants, fungi under variable environmental conditions.

KEYWORDS: Mycorrhiza, VAM, Soil Factors, Agricultural Practices, Environmental Stress, Soil Pathogen INTRODUCTION
Most mycorrhizas are balanced mutualistic associations in which the fungus and plant exchange commodities required for their growth and survival. Myco-heterotrophic plants have exploitative mycorrhizas where transfer processes apparently benefit only plants. Exploitative associations are symbiotic in the broad sense, but are not mutualistic. A new definition of mycorrhizas that encompasses all types of these associations while excluding other plant-fungus interactions recognizes the importance of nutrient transfer at an interface resulting from synchronized plant-fungus development. Mycorrhizal fungi also function as endophytes, necrotrophs and antagonists of host or non-host plants, with roles that vary during the lifespan of their associations. The purpose of this review is to consider and examine the diversity of mycorrhizal associations and the interactions between mycorrhizal fungi and plants and factors regulating their associations. Unique characteristics of mycorrhizas are generally identified and used to formulate a comprehensive definition of these associations that excludes other plant-fungus interactions. These plant fungal associations primarily responsible for nutrient transfer are essential to one or both organisms and involve synchronized development. The mycorrhizal morphology is the basis for grouping them into two major groups: ectomycorrhizae and endomycorrhizae. In this relationship, the fungi obtain carbon compounds and other nutritional requirements from the symbiotic plant roots, and in return, supply the plant with most of the immobile mineral elements such as Nitrogen (N), Phosphorus (P), Potassium (K), Calcium (Ca), Copper (Cu) and Zinc (Zn) from the soil solution, thus, becoming a significant component in low-input agricultural systems (Barea & Jeffries, 1995). Vesicular Arbuscular Mycorrhiza (VAM) is the utmost profuse kind of mycorrhiza Defined as a universal plant symbioses. This symbiosis is a highly progressed mutualistic relationship found between fungi and plants, most prevalent plant symbiosis known, and AM is found in 80% of vascular plant families in existence today. The marvelous advances in research on mycorrhizal physiology and ecology over the past 40 years have led to a larger understanding of the multiple roles of

18

Ameeta Sharma & Savita Yadav

VAMF in the ecosystem. This knowledge is applicable to human endeavors of ecosystem management, ecosystem restoration, and agriculture. Lack of host specificity is even more distinctive of this symbiosis than other types known. Mycorrhiza is the mutualistic symbiosis (non-pathogenic association) between soil-borne fungi with the roots of higher plants. Early morphological classifications separated mycorrhizas into endomycorrhizal, ectomycorrhizal and ectendomycorrhizal associations based on the relative location of fungi in roots. Mycorrhizas are multifaceted associations comprising diverse morphological, functional and evolutionary categories (Brundrett, 2002). Most mycorrhizas occur in roots, which evolved to house fungi (Brundrett, 2002), but they also occur in the subterranean stems of certain plants and the thallus of bryophytes (Read et al., 2000). Mycorrhizal associations where fungi do not seem to receive any benefits from plants have been called epiparasitic, myco-heterotrophic, or cheating associations. These fungi are considered to be primitive due to their relatively simple spores, their lack of sexual reproduction and because there are relatively few species of these fungi and they associate with a wide diversity of plants. Wet sieving and decanting methods for spore isolation are used, modified from Gerdemann (1955), and Gerdemann & Nicolson (1963). Mycorrhizas are the rule in nature, not the exception. In a mycorrhizal association, the fungus may colonize the roots of a host plant, either intracellularly or extracellularly. Mycorrhizae are present in 92% of plant families (80% of species) (Wang & Qiu, 2006), with endomycorrhizae or Arbuscular Mycorrhizae (AM) being the ancestral and predominant form and indeed the most prevalent symbiotic association found in all the plant kingdom. Conservative estimates suggest that this ancient symbiosis, dating back to the early Devonian age (398 million years ago), affects approximately 90% of the Earth's land plant species (Remy et al., 1994). Mycorrhiza association is a symbiotic non-pathogenic relationship between plant roots and fungal hyphal with a fungal connection between the soil and the root. Arbuscules consist of hyphae that branch dichotomously and profusely within root cortical cells. Arbuscules and other fungal structures do not penetrate host cell membranes, but invaginate them. Most species of plants are capable of associating with fungi of a single family, Endogonaceae, to form V-A mycorrhizae (Gerdemann, 1968). Arbuscular mycorrhizal fungi belong to the order Glomales, sub orders Glomineae and Gigasporineae and in the Zygomycetes habitats mycorrhizas may provide little benefit if the establishment of early successional species with low mycorrhizal dependency normally precedes successful establishment of mycorrhizal species.

VAM AND SOIL FACTORS

The mycorrhizal connections between plants have special significance for the tripartite legume association because of the latters role as a source of N to the associated soil ecosystem. Legumes with less extensive root systems are less efficient nutrient absorbers than grasses. Higher soil infectivity was observed under reduced or no tillage practices (Mozafar et al., 2000) and limited increased mycorrrhizal colonization of barley root and soil infectivity. Relative to conventional management, there is evidence that organic farming practices can enhance the amounts of AMF inoculum (Bending et al., 2004). New molecular methods have been applied to investigate mycorrhizal fungal communities (Bruns & Bidartondo, 2002) and greater attention has been paid to their possible effects at the level of the plant community. The intense exploitation of tropical forests has led to degradation of once stable ecosystems. There have been changes in abiotic and biotic soil properties, which hamper the re-establishment of proper vegetation cover. Under certain conditions, mycorrhiza is known to absorb fixed phosphate and even to stimulate root phytase activities. The higher height increment registered with inoculated plants could be as a result of enhanced inorganic nutrient absorption (Cooper, 1984) and greater rates of photosynthesis. Since nitrogen (N), phosphorus (P), and carbon (C)

Review on Role of VAM Fungi in Crop Plant-Soil System

19

are each required by mycorrhizal fungi, the availability of each nutrient could control mycorrhizal abundance. Plants provide C by transferring carbohydrates via roots; soils supply N and P. One of the more widely tested hypotheses within the field of mycorrhizal ecology is that plants should invest more C in mycorrhizal fungi where N or P are limiting to plant growth, since mycorrhizal fungi contribute to nutrient uptake by plants Alternately, mycorrhizal growth may be N-limited in some ecosystems (Treseder & Allen, 2002) so that N fertilization increases mycorrhizal abundance. Nonetheless, the mycorrhizal component may disappear or, at least, be severely depleted in degraded soils, so it may be necessary to reinforce or replace it by appropriate inoculation. Enzymatic activities are considered as major factors contributing to overall soil microbial activity and soil quality. The presence in the soil of given mycorrhizal species can influence the persistence and activities of native bacteria and vice versa (Andrade et al., 1998).

INTERACTION WITH AGRICULTURAL PRACTICES

Vesicular arbuscular mycorrhizae can improve plant acquisition of soil minerals by soil exploration and can better enable a plant to withstand environmental stresses. In natural ecosystems, both VAM colonization and spore number show seasonal variation and other agricultural soils also show an increase in spore numbers late in the season and at crop senescence. VAM fungi are able to absorb and transfer all of the 15 major, macro and micro nutrients necessary for plant growth (Lester, 2009). In general, agricultural practices have a negative impact on the AM association and agricultural soils are AMF impoverished, particularly in terms of number of species (Menendez et al., 2001). AM fungi respond to host exudates with extensive hyphal growth and branching. The establishment of a mycorrhizal symbiosis can improve the performance of the shrubs. Mycorrhiza represents ecological key factors governing the cycles of major plant nutrients and has significant influence on plant health and productivity (Jeffries et al., 2003). Agricultural practices can have major short or long term impacts on mycorrhizal fungi, as well as on other soil microorganisms. Scullion et al., (1998) studied the effectiveness of VAM spores from organically and conventionally managed soils and found that white clover only benefited from mycorrhizal infection in a low-fertility organically managed soil. Soil microbial biomass is dictated by a variety of soil and environmental parameters including soil texture and structure, pH, air/moisture content, and soil temperature. In natural ecosystems, much of the root system can be colonized by mycorrhizal fungi, which are restricted to the root cortex and do not enter the vascular cylinder. Perennial plants typically also have much more root structural diversity than annual plants. The plant roots provide substances for the fungi and the fungi transfer nutrients and water to the plant roots (Adholeya et al., 2005).

EFFECT ON DROUGHT AND SALINITY

Vesicular arbuscular mycorrhizal fungi can affect the water balance of both adequately watered and droughted host plants. The effects of rootstock, irrigation regime and water salinity on the vertical distribution of vesicular arbuscular mycorrhiza (VAM) of citrus roots were investigated in the field. Rootstocks differed in the uniformity of the vertical distribution of VAM, notably Rough lemon exhibited a uniform vertical distribution and Sour orange a non-uniform distribution with decreased VAM infection at depth. Decreasing the interval between irrigations reduced VAM infection of Sour orange, especially in the upper layers of soil. Increased salinity of irrigation-water reduced VAM in deep layers of soil. Effect on drought and salinity colonization by VAM fungi improves the drought resistance of plants ( Puppi and Bras, 1990).These differential effects might be due to the origin of the VAM fungi since mycorrhizal to edaphic conditions fungi adapt. Mycorrhizal improvement of drought tolerance occurs via drought avoidance. It can be a function of the often observed improved acquisition of phosphorus, nitrogen and other growth promoting nutrients by AMF plants (Aug,

20

Ameeta Sharma & Savita Yadav

2001). Mycorrhizae often improve plant growth and survival by facilitating uptake of nutrients and increasing drought tolerance. In addition to increased nutrition, mycorrhizal plants also show increased resistance to root pathogens and tolerance to drought stress, and their hormonal balance is altered. In greenhouse experiments, mycorrhizas| have been shown to increase the drought resistance of cultivated crops, such as wheat, onion and pepper. As the basic setting in which processes in soils take place, soil structure influences many biotic, physical and chemical aspects of soils, reviewed previously (Six et al., 2004). VAM fungi are abundant in grassland, savannah and tropical forests and associate with many grasses, herbs, tropical trees and shrubs (Read and Perez-Moreno, 2003). Furthermore, mutualistic symbiosis with mycorrhizal and endophytic fungi can confer salt tolerance to plants and decrease yield losses in cultivated crops grown in saline soils (Rodriguez et al., 2004). Under drought conditions the uptake of highly mobile nutrients such as NO3- can also be enhanced by mycorrhizal associations (Subramanian and Charest, 1999). Drought stress is a major agricultural constraint in the semi-arid tropics. It is known to have a considerable negative impact on nodule function. Drought inhibits photosynthesis and disturbs the delicate mechanism of oxygen control in nodules. The latter is essential for active nitrogen fixation. As mycorrhizae may enhance the ability of the plant to scope with water stress situtations associated to nutrient deficiency and drought, mycorrhizal inoculation with suitable fungi has been proposed as a promising tool for improving restoration success in semi-arid degraded areas. VAM colonization has been found in plants growing in frequently waterlogged soils. VA-mycorrhizal fungi most commonly observed in saline soils are Glomus spp. which suggests that this may be adapted to grow in saline conditions, but ecological specificity has not been demonstrated.

INTERACTION WITH SOIL PATHOGENS

The placement of the residue in soil can lead to the dislodgment of a pathogen from its preferred niche, thinning the pathogens capacity to survive (Bailey and Lazarovits, 2003). Rotation is most successful in limiting the impact of biotrophic pathogens that require living host tissues, or those pathogens with low saprophytic continued existence capability. Even in natural systems, plants with fibrous root architecture show only marginal reduction in P achievement when their mycorrhizas are impaired by fungicides. Many problems are associated with controlling pathogens with longterm persistent survival structures due to difficulties in reducing pathogen inoculum and lack of good sources of plant resistance. Data supports the Newsham, et al., (1995) hypothesis that plant identity can determine the degree to which AM fungi can protect plant roots from pathogens. The VAM fungal partner played a larger role in protecting the root from a pathogen in the fine-rooted plant compared to the coarse-rooted plant. However, in addition, our data also support the hypothesis that the identity of the AM fungi influences the ability of the mycorrhiza to reduce pathogen infection as previously demonstrated by Maherali and Klironomos (2007). Timing of inoculations may be a key determinant of AM fungal-mediated pathogen protection particularly if priority effects determine the outcome of the interaction (Kennedy and Bruns, 2005). VAM fungi can also affect plant growth indirectly by improving the soil structure, providing antagonist effects against pathogens and altered water relationships. Rotation is most successful in limiting the impact of biotrophic pathogens that require living host tissues, or those pathogens with low saprophytic survival capability.

Review on Role of VAM Fungi in Crop Plant-Soil System

21

CHALLENGES AND OPPORTUNITIES

Fungi are ubiquitous; some having beneficial effects on plants, while others may be detrimental (Ipsilantis and Sylvia, 2007). This review highlights the existing potential of mycorrhiza in the complex plant-soil system. VAM hyphae may explain antagonistic effects observed between VAM colonization and nodulation. Mycorrhizal fungi improve crop yields, especially in infertile soil (Hayman, 1982).The occurrence of two types of mycorrhizas in the same root system raises important questions about the relative benefits they provide to plants. Plants ultimately control the extent of mycorrhizal formation by regulating root growth and turnover. AM fungi can also reduce growth of several rudderal plant species, perhaps suggesting that rudderal species are more sensitive to microbial colonization. Mycorrhizal fungi microbial associations with mycorrhizal fungi and bacteria that live on and near the roots) offer several benefits to the host plant, including faster growth, improved nutrition, greater drought resistance and protection from pathogens (Cooper and Grandison, 1986). Vesicular-arbuscular mycorrhiza colonization also protects roots from soil pathogens and thereby increase root growth and nutrients acquisition of the host plants. Vesicular arbuscular mycorrhiza inoculation has a high potential in agroforestry as a bio-fertilizer. The mycorrhiza fungi live symbiotically with the roots of different architecture such as those involving cereal-legume mixtures. Inoculation with vesicular-arbuscular mycorrhiza increased plant tissue phosphorus, nitrogen and potassium content. The better growth response of mycorrhizal plants was attributed to improvement in nutrient uptake, especially phosphorous, nitrogen and potassium. They may enhance the uptake of water and nutrients (macro- and micro-elements) from the soil making them available to plants and hence meeting some of the mineral nutritional requirements. This could also help to reduce some of the fertility constraints, moisture requirements, and increases the drought resistance of the crops. Generally, mycorrhiza will improve soil structure, enhance plant health and vigor and minimize stress caused by pathogenic fungi, weeds and pollution from heavy metals. The interaction of these factors with vesicular-arbuscular mycorrhizae (VAM) is poorly understood except in a few cases. It is now very clear that VA mycorrhizal associations are ecologically significant factors that require more attention than previously accorded.

REFERENCES
1. Adholeya, A., Tiwari, P. & Singh, R. (2005). Large scale inoculum production of arbuscular mycorrhizal fungi on root organs and inoculation strategies. In: Soil Biology, In Vitro Culture of Mycorrhizae (Eds. S. Declerck, D.G. Strullu and A. Fortin). Springer-Verlag Berlin Heidelberg: 4 (1) 315-338. 2. Andrade, G., Linderman, R.G. & Bethlenfalvay, G.J. (1998). Bacterial associations with the mycorrhizosphere and hyphosphere of the arbuscular mycorrhizal fungus Glomus mosseae. Plant Soil, 202(1) 79-87. 3. Auge, R. M. (2001). Water relations, drought and vesicular-arbuscular mycorrhizal

symbiosis. Mycorrhiza :11(4) 342. 4. Bailey, K.L. & Lazarovits, G. (2003). Suppressing soil-borne diseases with residue management and organic amendments. Soil Till Res: 72 (3) 169-80. 5. Barea, J.M. & Jeffries, P. (1995). Arbuscular mycorrhizas in sustainable soil plant systems. In Mycorrhiza: Structure, Function, Molecular Biology and Biotechnology.( Eds Varma A & Hock B ) Springer-Verlag, Berlin: 53 (374) 521560. 6. Bending, G.D., Turner M.K., Rayns F.R., Marx, M.C. & M. Wood. (2004). Microbial and biochemical indicators of soil quality and their potential for differentiating areas under contrasting agricultural management regimes. Soil. Biol. Biochem. 36 (11) 1785-1792. 7. Brundrett, M. C. (2002). Coevolution of roots and mycorrhizas of land plants. New Phytologist 154 (2) 275304.

22

Ameeta Sharma & Savita Yadav

8.

Bruns, T. D. & Bidartondo, M. I. (2002). Molecular windows into the below-ground interactions of ectomycorrhizal fungi. Mycologist: 16 (2) 47-50.

9.

Cooper, K.M. (1984). Physiology of VA mycorrhizal associations. In: VA Mycorrhiza (Eds. C. L. Powell & D. J. Bagyaraj), CRC Press, Inc., Boca Raton, Florida:57 (5) 155-186

10. Cooper, K.M. & Grandison, G.S. (1986). Interaction of VAM fungi and root knor nematode on cultivars of tomato and white clover susceptible to Meliodogyne hapla. Annal. Biol., 108, 555-65. 11. Gerdemann, J. W. (1955). Relation of a large soil-borne spore to phytomycetous mycorrhizal infections. Mycologia 47 (5) 619-632. 12. Gerdemann, J.W. & Nicholson, T. H. (1963). Spores of mycorrhizal Endogone species extracted from soil by wet sieving and decanting. Transactions of the British Mycological Society: 46 (2) 235-244. 13. Gerdemann, J.W. (1968). Vesicular-arbuscular mycorrhizae and plant growth. Annual Review of Phytopathology: 6 (4) 397-418. 14. Hayman, D.S. (1982). Practical Aspects of Vesicular-Arbuscular Mycorrhiza. In: Advances in Agricultural Microbiology, (Ed. SubbraRao, N.S.). Oxford and IBM Publishing Company, New Delhi: 9 (2) 325-373. 15. Ipsilantis, I. & Sylvia, D.M. (2007). Abundance of fungi and bacteria in a nutrient-impacted Florida wetland Applied Soil Ecology :35 (2) 272-280. 16. Jeffries, P., Gianinazzi, S., Perotto, S., Turnau, K. & Barea, J. M. (2003). The contribution of arbuscular mycorrhizal fungi in sustainable maintenance of plant health and soil fertility. Biol. Frtil. Soils: 37 (3) 1-16. 17. Lester, D. (2009). Buying and applying mycorrhizal fungi. Max. Yield. USA: 483 (4) 126-131. 18. Maherali, H. & Klironomos, J.N. (2007). Influence of Phylogeny on fungal community assembly and ecosystem functioning. Science: 316 (5832) 17461748. 19. Menendez, A.B., J.M. Scervino & A.M. Godeas. (2001). Arbuscular mycorrhizal population. Associated with natural and cultivated vegetation on a site of Buenos Aires Province, Argentina. Biol. Ferti. Soils: 33 (12) 373381. 20. Mozafar, A. T., Anken, R. R. & Frossard, E. (2000). Tillage intensity, mycorrhizal and nonmycorrhizal fungi and nutrient concentrations in maize, wheat and canola. Agron. J: 92 (6) 1117-1124. 21. Newsham, K.K., Fitter, A.H. & Watkinson, A.R. (1995). Multi-functionality and biodiversity in arbuscular mycorrhizas. Trends in Ecology & Evolution: 10 (10) 407411. 22. Puppi, G. & Bras, A. (1990). Nutrient and water relations of mycorrhizal white clover. Agric Ecosyst. Environ. 29: 317-322. 23. Read, D. J., Duckett, J. G., Francis, R., Ligrone, R. & Russell, A. (2000). Symbiotic fungal associations in lower land plants. Philosophical Transactions of the Royal Society of London, B 355: (1398) 815831. 24. Remy, W., Taylor, T.N., Hass, H. & Kerp, H. (1994). Four hundred million year old vesicular arbuscular mycorrhizae. Proc Natl Acad Sci USA: 91 (2) 1184111843. 25. Rodriguez R.J., Redman R.S. & Henson J. (2004). The role of fungal symbioses in the adaptation of plants to high stress environments. Mitigation and Adaptation Strategies for Global Change: 9 (5) 261272. 26. Scullion, J., Eason, W.R, & Scott, E.P. (1998). The affectivity of arbuscular mycorrhizal fungi from high input conventional and organic grassland and grass-arable rotations. Plant Soil: 204 (2) 243-54. 27. Six, J., Bossuyt, H., Degryze, S. & Denef, K. (2004). A history of research on the link between (micro) aggregates, soil biota, and soil organic matter dynamics. Soil and Tillage Research: 79 (1) 731.

Review on Role of VAM Fungi in Crop Plant-Soil System

23

28. Subramanian, K.S. & Charest, C. (1999). Acquisition of N by external hyphae of an arbuscular mycorrhizal fungus and its impact on physiological responses in maize under drought-stressed and well watered conditions. Mycorrhiza 9 (2) 69-75. 29. Treseder, K.K. & Allen, M.F. (2002). Direct N and P limitation of arbuscular mycorrhizal fungi: a model and field test. New Phytologist : 155(3) 507515. 30. Wang, B. & Qui, Y.L. (2006). Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza: 16(5) 299-363.