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CHAPTER 1.

25 Root and Stem Nodule Bacteria of Legumes 821


Habitat
Rhizobia through their ability to x N
2
in symbi-
osis with legumes play a central role in the N
supply of most natural ecosystems. The Ameri-
can tall grass prairie is but one ecosystem in
which plant diversity and productivity is con-
trolled in large measure by N availability (Col-
lins et al., 1998). Rhizobia, although thought to
be solely soil saprophytes, can also be found in
aquatic systems associated with water-growing
leguminous plants. Owing to cultural and agri-
cultural practices, the migration of birds and
animals, and atmospheric deposition of soil par-
ticles, there are relatively few soils in the world
that do not contain some rhizobia. Rhizobia have
been shown to exist in soils for a relatively long
time in the absence of a host plant (Bottomley,
1992; Brunel et al., 1988; Kucey and Hynes, 1989;
Sanginga et al., 1994; Slattery and Coventry,
1993; Weaver et al., 1972).
Rhizobia have been recognized as being
important for the functioning of soil ecosystems
for centuries (Fred et al., 1932). Shortly, after
legume root nodules were shown conclusively to
assimilate atmospheric N
2
(Hellriegel and
Wilfarth, 1888), Nodbe and Hiltner applied for,
and were granted, a patent for the use of these
microorganisms as legume inoculants (Elkan
and Bunn, 1994). This and subsequent farming
and cultural practices have led to the dissemina-
tion of rhizobia on a global basis.
Rhizobia in soils may be introduced by appli-
cation of commercial inoculants or, as in many
cases, be the normal ora present as microsym-
bionts of an indigenous legume. Inoculants
applied to seed, as recommended by their man-
ufacturer, achieve inoculation rates of 10
3
10
6
rhizobia seed
-1
(Somasegaran and Hoben, 1994).
This corresponds to application rates of up to 8
10
10
rhizobia ha
-1
(Brockwell and Bottomley,
1995). At these rates, inoculant strains often
dominate in nodulation in the rst year of a
newly introduced crop (Brockwell et al., 1982;
Gibson et al., 1976; Singleton and Tavares, 1986).
Moreover, inoculant strains contribute to the
rapid buildup of rhizobia in the soil once nod-
ulessenesce and release large numbers of viable
rhizobia into the soil system (McDermott et al.,
1987; Sutton, 1983). Several studies have docu-
mented that inoculant strains dominate in nod-
ules 515 years after initial inoculation (Brunel
et al., 1988; Diatloff, 1977; Lindstrom et al.,
1990). It should be noted, however, that not all
introduced legumes receive inoculation, and in
such situations, seed, soil or aerial contamination
will usually lead to some initial nodule forma-
tion, and over a period of 45 years, a buildup
of soil rhizobial populations (Sadowsky and
Graham, 1998a). Moreover, diverse rhizobial
populations can develop in association with
species that are not initially indigenous to a
particular region (Leung et al., 1994). Although
it is thought that rhizobia in soil have a clonal
Table 2. Differences among genera of root nodule bacteria in the carbon compounds used for growth
1
.
Symbols: +, positive reaction; -, negative reaction; +/-, discriminatory within the genus; (+), mainly positive reaction; (-),
mainly negative reaction.
1
Includes data from Elkan and Brunn, 1992; de Lajudie et al., 1994, 1998; Rome et al., 1996; Jarvis et al., 1997 and Wang
et al., 1999.
Genus of nodule bacteria
Carbon source Rhizobium Sinorhizobium Mesorhizobium Allorhizobium Bradyrhizobium Azorhizobium
Adonitol + + + - -
D-Arabinose + + + - + -
L-Arabinose (+) + + -
D-Cellobiose + + + - -
L-Fucose + +/- +/- - - -
Inositol + + +/- + - -
Gluconate + (+) - + +
Lactose + + (+) + - -
L-Lysine +/- (-) - - -
DL-Malate (+) (+) +/- + (+) +
D-Maltose + + + + - -
D-Mannose + + (+) + + -
Mannitol + (+) + + (+) -
D-Mellibiose + + (-) - -
D-Rafnose + + +/- - - -
Ribose + + + + -
L-Rhamnose + + + + (+) -
Sucrose + + + (-) + -
Trehalose + + + (-) (+) -
D-Xylose + + + - + -