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Singh Exudates Root 2006

Singh Exudates Root 2006

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Published by: Gregorio Arone on Oct 28, 2012
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Geeta Singh, Krishna G. Mukerji
The plant-soil interrelationship is a subject of interest in ecology and agri-cultural sciences. While it is common knowledge that some plants only grow in special soils or that the yield depends on the condition of the soil,the fact that the plant may strongly influence the soil in which it grows hasnotveryoftenbeentakenintoaccount.Manyoftheseeffectsmaybecausedby specific ionic uptake or the exudation of specific organic compoundsor other rhizosphere effects (Marschner 1984; Peters and Long 1988). Theecological importance of such rhizosphere effects has been emphasizedtime and again (Mukerji 2002).The rhizosphere, generally defined as that volume of soil adjacent toand influenced by the plant root, is regarded as a “hot spot” for micro-bial colonization and activity (Metting 1993). In contrast to bulk soil,where available organic carbon sources are only at low concentrations,rhizospheres are supplied with higher concentrations of nutrient sourcesgenerated during plant photosynthesis. (Duineveld et al. 1998, 2001).Microorganisms in the rhizosphere of plants dominate the cycling of nutrients in soil-plant systems. Rhizosphere microorganisms increase theabilityofplantstoacquirenutrientsfromsoilbyeitherincreasingtheextentof the root system (e.g. through fungal hyphae) or solubilizing macronu-trients such as phosphorus or sulfur (Smith and Read 1997). In fact, plantgrowthinnutrient-poorenvironmentsislinkedtotheformationofmutual-isticassociationswithsoilmicroorganismssuchasarbuscularmycorrhizalfungi (Smith and Read 1997). Soil microorganisms also influence plant-pathogen interactions. For example, bacteria inoculated onto cucumberroot (
Cucumis sativus
) penetrate the root and induce systemic resistanceagainst a range of cucumber diseases (Wei et al. 1996).The abundance and activities of soil microorganisms are influenced by variousenvironmental(e.g.soiltype,nutrientsstatus,pH,texture,organic
Geeta Singh: Division of Microbiology, Indian Agricultural Research Institute, New Delhi-12, IndiaKrishna G. Mukerji: Department of Botany, University of Delhi, Delhi-110007, IndiaSoil Biology, Volume 7Microbial Activity in the RhizosphereK.G. Mukerji, C. Manoharachary, J. Singh (Eds.)
Springer-Verlag Berlin Heidelberg 2006
40 G. Singh, K.G. Mukerji
matter content, rootzonelocation,moistureandinteractionsofthesevari-ables etc. (Gorlenko et al. 1997; Kreitz and Anderson 1997; Marschner etal. 1989) as well as plant factors (e.g. species, age). Survival of differentintroduced strains of fluorescent pseudomonads varies in soils of differenttextures (Araujo et al. 1994). However, microbial growth in soil is carbonlimitedandtherefore,thepresenceoforganicmatterhasthegreatestinflu-enceonmicrobialpopulations(Wardle1992;Mukerji2002).
Pseudomonas,Flavobacterium, Alcaligenes
species have been shownto be particularly stimulated in the rhizosphere due to release of exudatesand lysates (Curl and Truelove 1986). The variety of organic compoundsreleased by plants has been postulated to be a key factor influencing thediversity of microorganisms in the rhizospheres of different plant species(Bolton et al. 1992; Bansal and Mukerji 1994). However, there is no directevidence to support this hypothesis.Rhizosphere is chemically, physically and biologically complex. Plantrootreleasenearly20%oftheirphotosyntheticassimilatesaswatersolublesugars, amino acids, hormones, vitamins; root exudates in the form of organic acids, amino compounds and sugar phosphate ester (Uren 2001).The released low molecular weight exudates, lysates, gases, e.g. ethyleneandCO
, mucilageandorganiccompoundshavebeenpostulatedto bekey factorsinfluencingthediversityandgrowthratesofmicroorganismsintherhizosphere of different plant species. The effect of root exudates dependson the distance that they can diffuse away from the rhizoplane (Gupta andMukerji 2002).The identification and quantification of root exudates is of significanceasthesecompoundsaltertheflowofnutrientsintherhizosphereandstim-ulate natural transformations in soils (Marschner et al. 1989; Nielsen andElsas 2001). The diffusion rate into soil is a function of the size and chargeof the compound and soil properties such as soil texture, organic mattercontent, pH, moisture and temperature. Although roots release a variety of organic substances, water-soluble compounds are most readily decom-posable substances which serves as a source of nutrients for microbialpopulations colonizing root surfaces. The microorganism in turn can af-fect growth and physiology of plants by various microbial metabolites orby modifying soil root environment.Rootexudates may create a niche that influences which microorganismsare to colonize the rhizosphere, thereby altering the composition and di-versity of micro organism colonizing the rhizosphere in a plant specificmanner (Grayston et al. 1998).Duineveld et al. (1998) observed that the effect of root exudates ondominant bacterial species can be marginal as most micro organisms areoligotrophic and thus respond slowly to root exudation. However Falchiniet al. (2003) showed that Copiotrophs are favoured by the addition of 
3 Root Exudates as Determinant of Rhizospheric Microbial Biodiversity 41
low molecular substrates passively released by roots. However in reality roots exude a spectra of substrates following a certain dynamic pattern(Kuzyakov and Cheng 2001), thus masking the effect of the individualcompound.
Knowledgeofthecompositionandquantityoforganicsubstancesreleasedfrom roots of different plant species gives an insight into chemical andbiologicalprocessesoccurringintherhizosphere.Rootexudatescomprisesprimary metabolites such as sugars, amino acids, sugar alcohols, organicacids and secondary metabolites (Gupta and Mukerji 2002).The major portion of microbial biodiversity is created by secondary metabolites. Plant roots also accumulate and exude numerous terpenoids,coumarins and flavonoids.
The qualitative and quantitative composition of root exudates are affectedby various environmental factors, including pH, soil type, oxygen status,light intensity, soil temperature, nutrient availability and the presence of microorganisms. These factors may have a greater impact upon root exu-dation than differences due to plant species.Soil texture has been shown to affect root exudates as it alters the me-chanical impedanceto the root(Boeuf-Tremblayet al. 1995).Similarlyclacontent influences the binding of different amino acids and peptides. Mi-croorganisms are known to stimulate amounts and composition of rootexudates (Meharg and Killham 1995). Plant species and the age of a plantisknown to influence the composition of rootexudates (Jaeger et al. 1999).Soilmicroorganismshavebeenshowntorespondtoplantexudationandplantspeciescanhavedifferentrootexudationpatterns(Brimecombeetal.2001).Peasand oatsexudedifferent amounts ofamino acids. Several pineshavedifferentexudationpatterns.Youngmaplesareknowntoexudegreaterand more diverse carbohydrates compared to mature trees, while maturetrees exude greater and more diverse amino acids. Chantigny et al. (1996)found a correlation between the water-soluble organic C and microbialbiomass C found under various annual and perennial species, suggestingdifferences in C deposition from the plant species. Martin (1971) showedthat clover had a greater exudation rate than wheat, which was higher than

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