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182j 9 Rice–Rhizobia Association: Evolution of an Alternate Niche<br />

Table 9.3 Colonization potential of R. leguminosarum bv. trifolii<br />

strain E11 and isolate BSS 202 from S. spontaneum on four rice<br />

varieties preferred <strong>by</strong> low-income Filipino farmers.<br />

Inoculum strain Sample location Colonization potential (viable plate count/three rice<br />

roots)<br />

Rlt E11 External rooting medium 1.03 10 7<br />

BSS 202 External rooting medium 9.17 10 7<br />

Rlt E11 Root surface 6.34 10 7<br />

BSS 202 Root surface 1.08 10 7<br />

Rlt E11 Root interior 1.50 10 8<br />

BSS 202 Root interior 7.50 10 7<br />

9.7<br />

Mechanisms of <strong>Plant</strong> Growth Promotion <strong>by</strong> Endophytic Rhizobia<br />

The ability of some endophytic rhizobial strains to promote the growth of rice and<br />

wheat prompted follow-up studies to identify possible mechanisms operative in this<br />

beneficial plant–microbe interaction. These studies have focused primarily on rice,<br />

addressing the following possible mechanism of rhizobial growth promotion: (1)<br />

induction of an expansive root architecture having enhanced efficiency in plant<br />

mineral nutrient uptake; (2) production of extracellular growth-regulating phytohormones;<br />

(3) solubilization of precipitated inorganic and organic phosphate complexes,<br />

there<strong>by</strong> increasing the bioavailability of this important plant nutrient; (4)<br />

endophytic nitrogen fixation; (5) production of Fe-chelating siderophores; and (6)<br />

induction of systemic disease resistance.<br />

9.7.1<br />

Stimulation of Root Growth and Nutrient Uptake Efficiency<br />

Sinandomeng PSBRC 74 PSBRC 58 PSBRC 18<br />

1.50 10 7 7.16 10 7 8.17 10 7<br />

7.00 10 6 8.00 10 7 1.27 10 7<br />

1.05 10 8 6.50 10 7 6.50 10 7<br />

6.00 10 7 5.60 10 7 5.66 10 7<br />

2.19 10 9 1.32 10 8 1.03 10 9<br />

6.30 10 6 7.34 10 7 1.23 10 8<br />

Source: From Yanni et al. [7].<br />

Reprinted with permission from CSIRO Publishing (http://www.publish.csiro.au/journals/<br />

fpb).<br />

Responsive rice varieties commonly develop expanded root architectures when<br />

inoculated with candidate biofertilizer strains of rhizobia. This suggests that these<br />

rhizobial endophytes alter root development in ways that could make them better<br />

miners , more capable of exploiting a larger reservoir of plant nutrients from<br />

existing resources in the soil. This possibility was suggested in early studies showing<br />

significantly increased production of root biomass in plants that had been inoculated<br />

[1,7,42,57] and <strong>by</strong> studies using greenhouse-potted soil indicating significant increases<br />

in N, P, K and Fe uptake <strong>by</strong> rice plants inoculated with selected rhizobia,<br />

including rice endophyte strains [58]. More recent studies have confirmed this result<br />

using plants grown gnotobiotically with rhizobia in nutrient-poor medium (50%<br />

Hoaglands), followed <strong>by</strong> measurements of root architecture and mineral nutrient

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