Pseudomonas

P. aeruginosa colonies on an agar plate.

Scientific classification
Kingdom: Bacteria
Phylum: Proteobacteria
Class: Gammaproteobacteria
Order: Pseudomonadales
Family: Pseudomonadaceae
Genus: Pseudomonas
Migula 1894

Characteristics
Members of the genus display the following defining characteristics:[7]

• Rod shaped
• Gram-negative
• One or more polar flagella, providing motility
• Aerobic
• Non–spore forming
• positive catalase test
Other characteristics which tend to be associated with Pseudomonas species (with some
exceptions) include secretion of pyoverdine, a fluorescent yellow-green siderophore[8]
under iron-limiting conditions. Certain Pseudomonas species may also produce additional
types of siderophore, such as pyocyanin by Pseudomonas aeruginosa[9] and
thioquinolobactin by Pseudomonas fluorescens,[10]. Pseudomonas species also typically
give a positive result to the oxidase test, the absence of gas formation from glucose,
glucose is oxidised in oxidation/fermentation test using Hugh and Leifson O/F test, beta
hemolytic (on blood agar), indole negative, methyl red negative, Voges–Proskauer test
negative, and citrate positive.

The members of the genus demonstrate a great deal of metabolic diversity, and
consequently are able to colonise a wide range of niches[11]. Their ease of culture in vitro
and availability of an increasing number of Pseudomonas strain genome sequences has
made the genus an excellent focus for scientific research; the best studied species include
P. aeruginosa in its role as an opportunistic human pathogen, the plant pathogen P.
syringae, the soil bacterium P. putida, and the plant growth promoting P. fluorescens.

Use as biocontrol agents

Since the mid 1980s, certain members of the Pseudomonas genus have been applied to
cereal seeds or applied directly to soils as a way of preventing the growth or
establishment of crop pathogens. This practice is generically referred to as biocontrol.
The biocontrol properties of P. fluorescens strains (CHA0 or Pf-5 for example) are
currently best understood, although it is not clear exactly how the plant growth-
promoting properties of P. fluorescens are achieved. Theories include: that the bacteria
might induce systemic resistance in the host plant, so it can better resist attack by a true
pathogen; the bacteria might out compete other (pathogenic) soil microbes, e.g. by
siderophores giving a competitive advantage at scavenging for iron; the bacteria might
produce compounds antagonistic to other soil microbes, such as phenazine-type
antibiotics or hydrogen cyanide. There is experimental evidence to support all of these
theories, in certain conditions; a good review of the topic is written by Haas and
Defago[18].

Other notable Pseudomonas species with biocontrol properties include P. chlororaphis,

which produces a phenazine-type antibiotic active agent against certain fungal plant
pathogens[19], and the closely related species P. aurantiaca which produces di-2,4-
diacetylfluoroglucylmethan, a compound antibiotically active against Gram-positive
organisms[20].

Rhizobacteria
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Rhizobacteria are root-colonizing bacteria that form a symbiotic relationship with many
legumes. The name comes from the greek rhiza meaning root. Though parasitic varieties
of rhizobacteria exist, the term usually refers to bacteria that form a relationship
beneficial for both parties (mutualism). Such bacteria are often referred to as plant growth
promoting rhizobacteria, or PGPRs.

The mechanism by which the PGPRs promote the plant growth is not clearly understood,
but it is believed to be, by the production of siderophors, assimilation of minerals,
production of phytohormones, asymbiotic Nitrogen fixation, etc. Some of the PGPR
bacteria includes Pseudomonas putida, Azospirillum fluorescens, Azospirilum lipoferum
etc.

Though microbial inoculants are indisputably beneficial for crops, they are not widely
used in industrial agriculture, as large-scale application techniques have yet to become
economically viable. A notable exception is the use of rhizobial inoculants for legumes
such as peas. Inoculation with PGPRs ensure efficient nitrogen fixation, and they have
been employed in North American Agriculture for over 100 years.

Plant-growth promoting rhizobacteria

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Plant growth-promoting rhizobacteria (PGPR) was first defined by Kloepper and

Schroth [1] to describe soil bacteria that colonize the roots of plants following inoculation
onto seed and that enhance plant growth. The following are implicit in the colonization
process: ability to survive inoculation onto seed, to multiply in the spermosphere (region
surrounding the seed) in response to seed exudates, to attach to the root surface, and to
colonize the developing root system [2] . The ineffectiveness of PGPR in the field has
often been attributed to their inability to colonize plant roots [3],[4] . A variety of bacterial
traits and specific genes contribute to this process, but only a few have been identified.
These include motility, chemotaxis to seed and root exudates, production of pili or
fimbriae, production of specific cell surface components, ability to use specific
components of root exudates, protein secretion, and quorum sensing. The generation of
mutants altered in expression of these traits is aiding our understanding of the precise role
each one plays in the colonization process [5],[6]. Progress in the identification of new,
previously uncharacterized genes is being made using nonbiased screening strategies that
rely on gene fusion technologies. These strategies employ reporter transposons [7] and in
vitro expression technology (IVET) [8] to detect genes expressed during colonization.

Using molecular markers such as green fluorescent protein or fluorescent antibodies it is

possible to monitor the location of individual rhizobacteria on the root using confocal
laser scanning microscopy [9],[10],[11],. This approach has also been combined with an
rRNA-targeting probe to monitor the metabolic activity of a rhizobacterial strain in the
rhizosphere and showed that bacteria located at the root tip were most active [12]

Mechanisms of Action
PGPR enhance plant growth by direct and indirect means, but the specific mechanisms
involved have not all been well-characterized [13], Direct mechanisms of plant growth
promotion by PGPR can be demonstrated in the absence of plant pathogens or other
rhizosphere microorganisms, while indirect mechanisms involve the ability of PGPR to
reduce the deleterious effects of plant pathogens on crop yield. PGPR have been reported
to directly enhance plant growth by a variety of mechanisms: fixation of atmospheric
nitrogen that is transferred to the plant, production of siderophores that chelate iron and
make it available to the plant root, solubilization of minerals such as phosphorus, and
synthesis of phytohormones. Direct enhancement of mineral uptake due to increases in
specific ion fluxes at the root surface in the presence of PGPR has also been reported .
PGPR strains may use one or more of these mechanisms in the rhizosphere. Molecular
approaches using microbial and plant mutants altered in their ability to synthesize or
respond to specific phytohormones have increased our understanding of the role of
phytohormone synthesis as a direct mechanism of plant growth enhancement by PGPR
[14]
PGPR that synthesize auxins and cytokinins or that interfere with plant ethylene
synthesis have been identified.

References
1. ^ Kloepper, J. W., and Schroth, M. N. 1978. Plant growth-promoting
rhizobacteria on radishes. Pages 879-882 in: Proc. of the 4th Internat. Conf. on
Plant Pathogenic Bacter. Vol. 2, Station de Pathologie Vegetale et
Phytobacteriologie, INRA, Angers, France.
2. ^ Kloepper, J. W. 1993. Plant growth-promoting rhizobacteria as biological
control agents. Pages 255-274 in: Soil Microbial Ecology: Applications in
Agricultural and Environmental Management. F. B. Metting, Jr., ed. Marcel
Dekker Inc., New York, USA.
3. ^ Benizri, E., Baudoin, E., and Guckert, A. 2001. Root colonization by inoculated
plant growth promoting rhizobacteria. Biocontrol Sci. Technol. 11:557-574.
4. ^ Bloemberg, G. V., and Lugtenberg, B. J. J. 2001. Molecular basis of plant
growth promotion and biocontrol by rhizobacteria. Curr. Opin. Plant Biol. 4:343-
350.
5. ^ Lugtenberg, B. J. J., Dekkers, L., and Bloemberg, G. V. 2001. Molecular
determinants of rhizosphere colonization by Pseudomonas. Ann. Rev.
Phytopathol. 38:461-490.
6. ^ Persello-Cartieaux, F., Nussaume, L., and Robaglia, C. 2003. Tales from the
underground: Molecular plant-rhizobacteria interactions. Plant Cell Environ.
26:189-199.
7. ^ Roberts, D. P., Yucel, I., and Larkin, R. P. 1998. Genetic approaches for
analysis and manipulation of rhizosphere colonization by bacterial biocontrol
agents. Pages 415-431 in: Plant-Microbe interactions and Biological Control. G. J.
Boland and L. D. Kuykendall, eds. Books in Soils, Plants, and the Environment,
vol. 63. Marcel Dekker Inc., New York, USA.
8. ^ Rainey, P. B. 1999. Adaptation of Pseudomonas fluorescens to the plant
rhizosphere. Environ. Microbiol. 1:243-257.
9. ^ Bloemberg, G. V., Wijfjes, A. H. M., Lamers, G. E. M., Stuurman, N., and
Lugtenberg, B. J. J. 2000. Simultaneous imaging of Pseudomonas fluorescens
WCS365 populations expressing three different autofluorescent proteins in the
rhizosphere: New perspectives for studying microbial communities. Mol. Plant-
Microbe Interact. 13:1170-1176.
10. ^ Bloemberg, G. V., and Lugtenberg, B. J. J. 2001. Molecular basis of plant
growth promotion and biocontrol by rhizobacteria. Curr. Opin. Plant Biol. 4:343-
350.
11. ^ Sorensen, J., Jensen, L. E., and Nybroe, O. 2001. Soil and rhizosphere as
habitats for Pseudomonas inoculants: New knowledge on distribution, activity and
physiological state derived from micro-scale and single-cell studies. Plant Soil
232:97-108.
12. ^ Lubeck, P. S., Hansen, M., and Sorensen, J. 2000. Simultaneous detection of the
establishment of seed-inoculated Pseudomonas fluorescens strain DR54 and
native soil bacteria on sugar beet root surfaces using fluorescence antibody and in
situ hybridization techniques. FEMS Microbiol. Ecol. 33:11-19.
13. ^ Kloepper, J. W. 1993. Plant growth-promoting rhizobacteria as biological
control agents. Pages 255-274 in: Soil Microbial Ecology: Applications in
Agricultural and Environmental Management. F. B. Metting, Jr., ed. Marcel
Dekker Inc., New York, USA.
14. ^ Glick, B. R. 1995. The enhancement of plant growth by free-living bacteria.
Can. J. Microbiol. 41:109-117.