205_PDFsam_Ehara_Sago Palm

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location. For example, spore-forming bacteria, enumerated as heat-tolerant ones,


showed large fluctuations in their numbers, ranging from 0.01 to 1.2 × 106 g−1 soil
(more than 100 times difference) depending on the sampling sites, but the differ-
ences in total bacterial numbers were only less than five times. Recent studies dem-
onstrate that soil contains as many as 1010–1011 bacteria, 6000–50,000 bacterial
species, and up to 200 m of fungal hyphae per 1 g and the soil microbes play key
roles in ecosystems (van der Heijden et al. 2008).
In addition to soil, plants harbor a diverse and an abundant microbial community
and support their activities as the key primary producers in most terrestrial ecosys-
tems. In particular, the rhizosphere, defined as the soil environment influenced by the
presence and activities of plant roots, has a huge impact on soil microbes. Plant roots
release organic matter called rhizodeposition or root exudate, and the amount of
rhizodeposition represents ca. 11% of carbon fixed by the plant and 27% of carbon
allocated to roots (Dennis et al. 2010). Rhizodeposition supports the microbial activi-
ties in the rhizosphere. A wide range of organisms, i.e., different kinds of bacteria,
fungi, protozoa, and nematodes, show higher populations in the rhizosphere than in
the non-rhizosphere, which is not affected by plant roots. The rhizosphere consists of
endorhizosphere (root tissue area), rhizoplane (root surface), and exorhizosphere
(rhizosphere soil: soil directly surrounding the root). According to a report summa-
rizing the results of 22 different plants, the numbers of aerobic bacteria were on
average 9.9 times (2.6–24.2) higher in rhizosphere soil than in non-­rhizosphere soil
(Lochhead and Rouatt 1955), and microbial densities are markedly higher in the
rhizoplane than in the endorhizosphere. An example showed that the numbers of total
bacteria, Gram-negative bacteria, and fungi were 100–1000 times higher in the rhi-
zoplane than in the endorhizosphere (Fig. 15.1). Rhizosphere soil and the rhizoplane
are microbial hot spots, and many microbes colonize even the endorhizosphere.
In addition to belowground, the aboveground parts of plants also support a vari-
ety of bacteria, yeasts, and fungi. According to Lindow and Brandl (2003), bacteria
are by far the most numerous colonists of leaves, ranging from 106 to 107 cells cm−2
(up to 108 cells g−1). The aerial habitat colonized by these microbes is termed the
phyllosphere, and the inhabitants are called epiphytes. The number of epiphytes is
affected by surrounding environments. For example, Enya et al. (2007) compared
the number of leaf-associated bacteria between field-grown and greenhouse-grown
tomato leaves and found that the former harbored 100 times higher densities and the
population densities increased with age. Since sago palm grows under field condi-
tions, its leaves and stems are colonized by many microbes.
Microbes affect plant productivity through different mechanisms. First, a typical
one is direct effects on plants via plant-associated organisms that form mutualistic
or pathogenic relationships with plants. Another is indirect effects via the action of
free-living microbes that alter rates of nutrient supply and the partitioning of
resources (van der Heijden et al. 2008). Plant productivity is enhanced by different
microbial actions, while there are a number of microbial threats to crop production
(Table 15.1).
Nitrogen-fixing bacteria contribute to plant productivity to the largest extent, as
described below. Arbuscular mycorrhizal (AM) fungi enhance plant productivity,
especially in grasslands (van der Heijden et al. 1998). These two symbiotic micro-

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