Biotechnol Lett (2010) 32:15591570

DOI 10.1007/s10529-010-0347-0

REVIEW

Beneficial bacteria of agricultural importance

Olubukola Oluranti Babalola

Received: 10 May 2010 / Accepted: 28 June 2010 / Published online: 16 July 2010
Springer Science+Business Media B.V. 2010

Abstract The rhizosphere is the soilplant root

interphase and in practice consists of the soil
adhering to the root besides the loose soil surrounding
it. Plant growth-promoting rhizobacteria (PGPR) are
potential agents for the biological control of plant
pathogens. A biocontrol strain should be able to
protect the host plant from pathogens and fulfill the
requirement for strong colonization. Numerous compounds that are toxic to pathogens, such as HCN,
phenazines, pyrrolnitrin, and pyoluteorin as well as,
other enzymes, antibiotics, metabolites and phytohormones are the means by which PGPR act, just as
quorum sensing and chemotaxis which are vital for
rhizosphere competence and colonization. The presence of root exudates has a pronounced effect on the
rhizosphere where they serve as an energy source,
promoting growth and influencing the root system for
the rhizobacteria. In certain instances they have
products that inhibit the growth of soil-borne pathogens to the advantage of the plant root. A major
source of concern is reproducibility in the field due to
the complex interaction between the plant (plant
species), microbe and the environment (soil fertility
and moisture, day length, light intensity, length of

O. O. Babalola (&)
Department of Biological Sciences, Faculty
of Agriculture, Science and Technology, North-West
University, Private Bag X2046, Mmabatho 2735,
South Africa
e-mail: olubukola.babalola@nwu.ac.za

growing season, and temperature). This review listed

most of the documented PGPR genera and discussed
their exploitation.
Keywords Biocontrol
Exudate

PGPR
Phytohormones

Introduction
Plant growth-promoting rhizobacteria (PGPR) are
free-living bacteria of beneficial agricultural importance. The PGPR present encourage beneficial effects
on plant health and growth, suppress disease-causing
microbes and accelerate nutrient availability and
assimilation. Thus, in the quest to improve soil
fertility and crop yield and to reduce the negative
impacts of chemical fertilizers on the environment,
there is a need to exploit PGPR for continued
beneficial agricultural purposes. PGPR exist in the
rhizosphere and this is defined as the region around
the root. Seed inoculation with PGPR could be by
drench application, seed bacterization, or via dual
treatment. PGPR compensate for the reduction in
plant growth caused by weed infestation (Babalola
et al. 2007b), drought stress (Zahir et al. 2008), heavy
metals (Kumar et al. 2009), salt stress (Egamberdieva
2008; Kaymak et al. 2009) and some other unfavorable environmental conditions. One of the mechanisms by which bacteria are adsorbed onto soil

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particles is by simple ion exchange and a soil is said

to be naturally fertile when the soil organisms are
releasing inorganic nutrients from the organic
reserves at a rate sufficient to sustain rapid plant
growth. These bacteria belong to the genera
Acetobacter, Acinetobacter, Alcaligenes, Arthrobacter, Azoarcus, Azospirillum, Azotobacter, Bacillus,
Beijerinckia, Burkholderia, Derxia, Enterobacter,
Gluconacetobacter, Herbaspirillum, Klebsiella, Ochrobactrum, Pantoae, Pseudomonas, Rhodococcus, Serratia, Stenotrophomonas and Zoogloea and have been
subject of extensive research throughout the years.
Some PGPR may have more than one mechanism
for accomplishing plant growth (Ahmad et al. 2008).
Included in such mechanisms are antagonism to
pathogenic fungi, siderophore production, nitrogen
fixation, phosphate solubilization, the production of
organic acids and indole acetic acid (IAA), NH3,
HCN, the release of enzymes (soil dehydrogenase,
phosphatase, nitrogenase, etc.), and the induction of
systemic disease resistance (ISR). Similarly, the plant
beneficial effects of bacteria include attributes such
as the production of phytohormones, nitrate reduction, and nitrogen fixation. For that reason, potential
PGPR are sometimes screened in vitro for such
multiple plant growth promoting traits. PGPR as
these make use of the plant growth promoting traits to
improve the health of forest tree species (Barriuso
et al. 2008).
The use of beneficial soil microorganisms as
agricultural inputs for improved crop production
requires the selection of rhizosphere-competent
microorganisms with plant growth-promoting attributes (Hynes et al. 2008). The beneficial bacteria
termed PGPR are disease-suppressive microorganisms that improve plant health. Findings and documentation abound, and all point towards the need to
commercially exploit PGPR as biofertilizers for their
agricultural benefits. From a general perspective,
however, the problems of variability in colonization
efficiency, field performance and rhizosphere competence are controversial issues. This review provides
(1) an overview of these PGPR mechanisms and their
applications for beneficial agricultural purposes, (2) it
highlights the current knowledge on PGPR and (3)
examines the interactions of bacteria used as biocontrol agents of bacterial and fungal plant pathogens;
consideration is given to biofertilizer, quorum sensing, PGPR bioactive factors, plant growth promoters,

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metabolites, release of enzymes, and production of

antibiotics among others.

PGPR/biofertilizer
PGPR are bacteria that produce hormones, vitamins
and growth factors that improve plant growth and
increase plant yield. The endorhizosphere and exorhizosphere are the regions around the root. The
rhizosphere is richer in microflora and available
nutrients such as photosynthates from the roots than
the bulk soil. Manipulating microorganisms in the
rhizosphere is by diverse, competitive, complex,
highly regulated cell-to-cell communication and by
using signaling compounds to monitor their surroundings and alter their activities. Plants are also
involved in these signaling mechanisms. Microbial
activity in the rhizosphere may affect the acquisition
of mineral nutrients by roots either directly via effects
on mobilization and/or immobilization or indirectly
via effects on root morphology and/or physiology. A
large proportion of manganese oxidizing microorganisms in the rhizosphere may, therefore, either
decrease the risk of manganese toxicity in plants
growing in poorly aerated or waterlogged soils, or
increase the risk of manganese deficient in wellaerated calcareous soils.
The term Plant Growth-Promoting Rhizobacteria
(PGPR) was coined over three decades ago; they are
non-pathogenic, strongly root colonizing bacteria on
the surface of plants roots which increase plants
yield by one or more mechanisms. The PGPR
enhance the availability and uptake of plant nutrients,
the production of substances promoting plant growth
and the suppression of deleterious bacteria (Table 1).
Unlike the adverse effect of the continuous use of
chemical fertilizers, PGPR when applied to the soil
improve the soil structure, leaving no toxic effects.
Microbial inoculants or biofertilizers are an important
component of organic farming (Table 1); accounting
for about 65% of the nitrogen supply to crops
worldwide. PGPR are known to fix atmospheric
molecular nitrogen through symbiotic and asymbiotic
or associative nitrogen fixing processes (Anjum et al.
2007). With more and more emphasis being placed
on organic farming, PGPR are finding increasing
applications today as biofertilizers (Table 1).

Increased total microbial activity, shoot and root length,

total dry weight
Increased root and shoot growth, nodulation in legume

Arabidopsis

Broad beans
Wild plants

Bacillus subtilis GB03, B. amyloliquefaciens

IN937a, B. pumilus SE-34, B. pumilus T4,
B. pasteurii C9, Paenibacillus polymyxa E681,
P. fluorescens 89B-61, and S. marcescens
90-166

P. aeruginosa and Serratia liquefaciens

Pseudomonas spp.

Increased root elongation in cadmium supplemented

soil in pot

Improved fresh and dry weight, root length, shoot

length, number of leaves per plant, and water use
efficiency under drought stress
Increased total soluble solids, total sugar and reduced
sugar

Indian mustard and rape

Pea

Strawberry
Stored potatoes
Radish

Rice

Pseudomonas brassicacearum, P. marginalis,

P. oryzihabitans, P. putida, Alcaligenes
xylosoxidans, B. pumilus and Rhodococcus
spp.
P. fluorescens biotype G (ACC-5), P. fluorescens
(ACC-14), and P. putida biotype A (Q-7)

Pseudomonas BA-8, Bacillus OSU-142, and

Bacillus M-3

B. cepacia strain OSU-7

Agrobacterium rubi strain A 16, Burkholderia
gladii strain BA 7, P. putida strain BA 8,
B. subtilis strain BA 142, and B. megaterium
strain M 3

A. amazonense

Increased grain dry matter accumulation (711.6%), the

number of panicles (318.6%) and nitrogen
accumulation at grain maturation (3.518.5%)

Biocontrol agent of Fusarium dry rot

Improved the percentage of seed germination under
saline conditions

Improved plant growth and nutrition under salt stress

Mung bean
Wheat

Bradyrhizobium and PGPR

Acinetobacter, Alcaligenes faecalis, Bacillus

cereus, Enterobacter hormaechei, Pantoae,
P. aeruginosa

Better phytoremediation potential of broad bean plants

grown in oily sand

Increased foliar fresh weight

Decreased the number of diseased plants caused by

Phytophthora. Increases in shoot length (81%), shoot
dry weight (68%), root length (152%), and root dry
weight (290%)

Betelvine

Serratia marcescens NBRI1213

Effect compared with control

Plant species

PGPR

Table 1 Some PGPR and their beneficial effects on plants

Rodrigues et al. (2008)

Recep et al. (2009)

Kaymak et al. (2009)

Pirlak and Kose (2009)

Zahir et al. (2008)

Belimov et al. (2005)

Egamberdieva (2008)

Shaharoona et al. (2006)

Ahn et al. (2007)

Radwan et al. (2005)

Ryu et al. (2005)

Lavania et al. (2006)

References

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Quorum sensing (QS)

In QS behavior small diffusible extracellular signaling molecules meditate cellcell communication. The
signaling molecules for Gram-negative bacteria are
named autoinducers, usually acylated homoserine
lactones (AHLs). The Gram-positive bacteria use
peptide-signaling molecules for QS. At a particular
population density, bacteria exert a concerted action;
the signals activate transcription factors that induce
specific genes in the bacteria. Examples of rhizobacteria that utilize AHL signals to regulate the expression of traits important in plantmicrobe interaction
are Pantoea stewartii, Pseudomonas aureofaciens,
and Pseudomonas syringae. Studies indicate that host
plant can interfere with the QS of bacteria. The QS
molecules in Azospirillum lipoferum are strain-specific and are associated with rhizosphere competence
and adaptation to plant roots (Boyer et al. 2008).
Plantmicrobe interactions such as biofertilization,
rhizoremediation, biocontrol and phytostimulation
may be QS-dependent. Some studies have highlighted the fact that bacteria form microcolonies on
plant roots and these are assumed to be suitable
setting for QS as a result of the high bacterial cell
density. In addition, the mucigel, secreted from the
roots, can cover the bacteria and this could retard
diffusion of N-acyl homoserine lactones required for
QS. The degradation of AHLs is a mechanism for
biocontrol. Two N-acyl homoserine lactone systems,
RhlI/R and LasI/R, in Pseudomonas aeruginosa
PUPa3 from rice rhizosphere are involved in the
regulation of plant growth-promoting traits (Steindler
et al. 2009).

Chemotaxis and mobility

Findings by some researchers using spinach roots
indicated that gravitational water flow in irrigation
plays an important role in the spread of the colonization of roots by PGPR Pseudomonas strains
(Urashima et al. 2004). When the colonization pattern
and the effect of bacterization on the early root
development of Pseudomonas fluorescens 5.014 on
tomato seeds were monitored, the inoculated strains
were distributed over the entire root and the distal
segment was more densely colonized compared with

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the proximal and middle segments. The density of the

indigenous seed flora was uneven but was stabilized
by the application of PGPR. Azospirillum brasilense
confirms both chemotaxis and chemokinesis reaction
to the sequential gradients of diverse chemo-effectors, thereby increasing the chance of rootbacterial
interactions. Genera Azospirillum needs bacterial
motility for establishment on plant root as demonstrated using non-flagellated mutants and a
non-chemotactic mutant. The understanding that
chemotaxis is essential for colonization has aided
the selection of bacterial strains that are good
colonizers of cereal root. Examples of chemotactic
agents in the rhizosphere are amino acids and sugar.
The root exudates of monocots improved growth
yield because the root tips of monocots, e.g. perennial
ryegrass, support ten times more bacteria than those
of dicots, e.g. potato, radish and tomato (Lugtenberg
et al. 2001).
Bacteria can move only in water films and,
although many are smaller than 1 lm diameter, they
appear to be readily motile only in films appreciably
thicker than this, although they may well be capable
of growth at lower moisture contents. For all motile
bacteria, the rate of movement is dependent on the
moisture content of the soil and is thus dependent on
soil structure. Moisture content affects the activity of
the soil population in a number of ways: through the
thickness of the water films in the soil and consequently its aeration; through the reduction in the free
energy of the water as the soil becomes drier and the
films become thinner, so affecting water availability
and the cell osmotic relationships; and finally through
the effects that can be demonstrated when soil
moisture fluctuates from the dry state to that of
becoming adequate. Movement becomes restricted as
the soil becomes drier, partly because the water film
pathway between two points in the soil becomes more
tortuous, and partly due to organisms becoming
trapped in moisture filled pores occluded by narrow
necks. Studies have indicated the non-uniform distribution and the lognormal distribution of root-colonizing bacteria in the rhizosphere. A number of areas
on the roots are virtually free from bacteria while
other areas can be densely inhabited. Root hairs
provide a hospitable biological surface for chemotaxis, migration and colonization due to microorganisms (Pothier et al. 2007).

Biotechnol Lett (2010) 32:15591570

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PGPR bioactive factors

Vitamins

PGPR bioactive factors are substances that impact on

growth. Examples include: root exudates, vitamins,
and amino acids.

Vitamin producing bacteria are widespread in the

rhizosphere because many bacteria excrete this compound. It should however be mentioned that the
vitamins are at time not in sufficient quantity. A large
proportion of indigenous bacteria requires or is stimulated by water-soluble B vitamins and amino acids.
They will not grow in simple laboratory media unless
supplemented with the appropriate substances. In the
absence of vitamins (thiamine, biotin, pantothenic
acid, folic acid, riboflavin, vitamin B12, nicotinic acid,
pyridoxine, and myoinositol), inoculation with a PGPR
P. putida strain G11-32 promoted the biomass of
soybean callus in a tissue culture bioassay. Similarly, a
thiamine auxotroph of P. fluorescens WCS365
appeared to be a poor competitor in the tomato
rhizosphere. The ability of PGPR to synthesize biotin
and/or thiamine appears to be an important colonization trait (Lugtenberg et al. 2001). Thiamine is the most
frequently required vitamin, but biotin and vitamin B12
also are essential for a large number of bacteria.

Root exudates
Root exudates are chemical compounds photosynthates, organic acid, sugar (Castro-Sowinski et al. 2007;
Kamilova et al. 2006), polyamine putrescine (Kuiper
et al. 2001) excreted from root tissues; the process is
called rhizodeposition. Indirect interactions between
plants and microbes occur in the rhizosphere due to
root exudates. Energy-rich organic acid materials
such as citric, lactic, succinic, amides, malic, oxalic,
pyruvic, aliphatic and aromatic acids, and fructose
(simple sugar), carbohydrates (glucose, xylose) loss
by exudation, secretion or autolysis of older root
cells; leaked moisture and root-growth-lubricant
mucigel secreted from the roots all provide nutrients
for microorganism growth (Alexandre et al. 2000). In
addition to such solubles and diffusibles, there is the
entire range of insolubles occurring in the root (e.g.,
cellulose, lignin, protein) and lost from it by cell
exfoliation and root pruning. Organic acids are 83%
of the total exudates and sugars roots increased
during plant growth (Kamilova et al. 2006). Some of
the matter lost by the roots has a low molecular
weight and is metabolized by the microbes. Rhizosphere bacteria are ecophysiologically adapted to
these chemical metabolites. The abundant supply of
organic carbon contained in or emanating from plant
root makes the rhizosphere an oasis for growth and
maintenance of bacteria in the bulk soil. Some rootassociated populations may be more sensitive to a
change in exudates (as may occur during plant
growth) than others (Herschkovitz et al. 2005).
Apparently, chemical signals between plant roots
and microbes influence community structure and
functions. PGPR are exogenous bacteria introduced
into agricultural ecosystems that act positively upon
plant development (Castro-Sowinski et al. 2007)
showed that P. putida strain PCL1444 cultivated in
the presence of grass seedlings cv. Barmultra in soil
led to a tenfold increase of cells. Based on the
underlying principle that root exudate is the best
nutrient source available in soil.

Amino acids
The ability to synthesize amino acids is an important
colonization trait. Notable among the amino acids are
glutamine and asparagine with glutamic acid, alanine,
serine, valine, isoleucine, aspartic and leucine which
are also present. The loss of amino acids by exudation
has been reported (Alexandre et al. 2000); the amount
of loss may vary with plants species, age and
developmental stage. The role of amino acid is to
support growth of auxotrophs.
Growth promoters
Growth promoters are organic molecules required in
trace quantities for growth. The need for growth
factors has considerable ecological importance
because a species needing them will only grow in
habitats where the organic molecules are present.

Production of phytohormones
Phytohormones produced by bacteria contributed to
the host root respiration rate, metabolism and root

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abundance and hence improved the mineral and water

uptake in inoculated plants. Gibberellins (GAs) may
influence seed germination, stem elongation, flowering, and fruit setting of higher plants. Four GAs (GA1,
GA3, GA4, and GA20) from seven species of bacteria,
Acetobacter diazotrophicus, A. lipoferum, A. brasilense, Bacillus licheniformis, Bacillus pumilus, Herbaspirillum seropedicea and Rhizobium phaseoli, have
been identified (MacMillan 2002). Azotobacter chroococcum, an inoculant that fixes atmospheric nitrogen,
produces on average 0.05 lg gibberellic acid (GA3)
equivalents per milliliter of culture in 14 days; and
when tomato seedling roots are treated with a culture,
growth is drastically changed and found similar to
treatment with authentic GA3. Bacterial inoculants,
when applied to plants, probably contain sufficient
growth regulators to influence future plant development, but as they multiply around seedling roots,
possibly more hormone is produced and absorbed by
the plants at the critical differentiation stage.
PGPR synthesize auxins, and these influence root
growth, cell elongation, tissue differentiation, plant
growth promotion and responses to light and gravity.
Auxins are the most important plant hormone
produced by Azospirilum, Bacillus megaterium and
Pseudomonas spp. HPLC analysis confirmed the
presence of IAA and indole acetamide (IAM) as the
major auxins in the culture filtrates of some rhizobacteria (Lopez-Bucio et al. 2007).

Biotechnol Lett (2010) 32:15591570

stress hormone when under conditions such as those

generated by salinity, drought, waterlogging, heavy
metals and pathogenicity. Under stress conditions, the
endogenous production of ethylene is accelerated
substantially which adversely affects root growth and
consequently the growth of the plant as a whole
(Saleem et al. 2007). The ability of 1-aminocyclopropane-1-carboxylate (ACC)-utilizing PGPR to
ameliorate plant growth when inhibited by ethylene
through a decrease in ACC content and the enzymatic
removal of ethylene has been demonstrated. Another
way bacteria stimulate plant growth is with ACC
deaminase that hydrolyzes the ethylene precursor
ACC and thus presents a promising opportunity to
increase crop yields. The bacteria occurring on the
root surface degrade ACC to ammonium and
a-ketobutyrate for use as carbon and nitrogen sources.
The results of (Shaharoona et al. 2007) demonstrated
that ACC-deaminase activity is an efficient parameter
for the selection of promising PGPR under axenic
conditions. Ethylene plays a key role in inducing
multifarious physiological changes in plants at the
molecular level (Saleem et al. 2007).

Metabolites
Hydrogen cyanide (HCN), 2,4-diacetylphloroglucinol
(DAPG) and siderophores are antifungal compounds.
Their production assists in the control of soil-borne
diseases.

Regulating ethylene production in roots

Ethylene is a plant growth regulator. For many plants,
a burst of ethylene, a gaseous natural byproduct of
plant metabolism, is required to break seed dormancy, increase the number of roots, shoot and root
growth differentiation, adventitious root formation,
leaf and fruit abscission, the induction of flowering
and increased femaleness in dioecious plants, flower
and leaf senescence, and fruit ripening (Babalola
et al. 2007b) but following seed germination, a
sustained high level of ethylene would inhibit root
elongation, and lead to abnormal root growth, which
would seriously impede plant growth and development. Although ethylene is involved in the regulation
of numerous physiological processes in plants, it is
also produced by PGPR (Babalola 2010). Apart from
being a plant growth regulator, ethylene acts as a

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Diacetylphloroglucinol (DAPG) production

DAPG is a polyketide antibiotic that suppresses a wide
variety of soil-borne fungal pathogens, including
Gaeumannomyces graminis var. tritici and Fusarium
oxysporum f. sp. radicis-lycopersici (Duffy et al. 2004).
An example of DAPG producer is P. fluorescens Q2-87
(Duffy et al. 2004). Strains of P. fluorescens that
generate DAPG are efficient PGPR controlling root
and seedling diseases.

Production of siderophores
Low molecular weight siderophores synthesized by
PGPR can solubilize and sequester iron from the soil

Biotechnol Lett (2010) 32:15591570

and then provide it to the plant cells. Ochrobactrum

anthropi TRS-2 (Chakraborty et al. 2009) isolated
from the rhizosphere of tea produces siderophore and
IAA in vitro. Pseudomonas spp. produces fluorescent
siderophores. These bacteria have biological control
and growth promoting capability.

Production of volatile compounds

Low molecular weight metabolites, such as HCN, a
biocide, with antifungal activity is released in a blend
of volatile organic compounds such as 2,3-butanediol
and acetoin that promote the growth of Arabidopsis
thaliana. In a comparison of the importance of HCN
and DAPG to P. fluorescens CHA0, HCN not
repressed by fusaric acid, played a more important
role in disease suppression than DAPG (Duffy et al.
2004). The production of HCN was a more common
trait of Pseudomonas (88.89%) (Ahmad et al. 2008).
Among other substances reported are pyoluteorin
(Baehler et al. 2006) and auxofuran (Riedlinger et al.
2006).

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Production of exo-enzymes that suppress

deleterious microbes
PGPR may synthesize some enzymes that modulate
plant growth and development. The enzymes are
usually for defence and they could be cell walldegrading hydrolytic enzymes. The production of
enzymes includes chitinase, cellulase, b-1,3-glucanase, protease, lipase can lyse some fungal cells
(Muleta et al. 2007) and suppress deleterious rhizobacteria. Examples of the best characterized defence
enzymes are peroxidase, polyphenol oxidase and
phenylalanine ammonia-lyase (Latha et al. 2009).
Organic and inorganic phosphate solubilization
Phosphorus is an essential nutrient for plants. There
are a number of microorganisms which can solubilize
the economical sources of phosphorus, for instance
rock phosphate. Among such phosphate-solubilizing
bacteria are Pseudomonas striata, Enterobacter,
Erwinia, B. megaterium and O. anthropi TRS-2
(Chakraborty et al. 2009). They solubilize the bound
phosphorus, mineralize organic phosphorus and
release soluble inorganic phosphate into soil through
decomposition of phosphate-rich organic compounds.

Release of enzymes
Production of antibiotics
Fixation of N2 that is transferred to the plant
The atmosphere contains 78% nitrogen. Leguminous
plants have a symbiotic (mutually advantageous)
relationship with the bacteria that provide fixed
nitrogen. In the rhizosphere, nitrogen-fixing (diazotrophic) bacteria are also present. Examples of
nitrogen-fixing bacteria are Azotobacter vinelandii,
Beijerinckia derxii and Zoogloea strain Ky1
(Meunchang et al. 2006). Culturable diazotrophs
from two traditional Indian rice cultivars, Sataria and
Kartiki, are H. seropedicae, Azospirillum amazonense, Burkholderia cepacia/vietnamiensis, and Pseudomonas spp. (Jha et al. 2009). Other bacterial genera
are Acetobacter, Arthrobacter, Alcaligenes, Bacillus,
Enterobacter, Klebsiella and Pseudomonas. These
are broadly divided into three categories, viz.,
symbiotic microorganism e.g. legumeRhizobium
symbiosis, asymbiotic or free living e.g. Azotobacter
and associative symbiosis, e.g. Azospirillum.

Some Bacillus spp. produce antibiotics and promote

plant growth (Choudhary and Johri 2009). The
production of phenazine antibiotics contributes to
ecological fitness by competing with the resident
microflora. The modes of action of these antibiotics
are not well known but they do suppress root
diseases. O. anthropi TRS-2 (Chakraborty et al.
2009) and Bacillus sp. MEP2 18 and ARP (2) 3
exhibit antagonistic activity against some phytopathogenic fungi (Principe et al. 2007). Also elaborated
are the antifungal properties of pyoluteorin, pyrrolnitrin and iturin.
Nematophagous bacteria
Nematophagous bacteria exhibit diverse modes of
action including promotion of plant health. They act
synergistically through the direct suppression of
nematodes, promoting plant growth, and facilitating

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the rhizosphere colonization and activity of microbial

antagonists in the rhizosphere.
Actinomycetes
Actinomycetes (Babalola et al. 2009), when associated to root systems, are considered rhizobacteria and
some behave as true PGPR, showing good potential
as inducer of ISR where the interaction has been
mentioned to interfere with the autoregulation of
nodulation in alfalfa (Solans et al. 2009). Streptomyces successfully suppresses the growth of plantpathogenic fungi (Riedlinger et al. 2006). This is not
to say that all Streptomyces spp. are PGPR. The plant
growth-promoter as well as plant growth-retardant
constitutes the genus Actinomyces.
PGPR application methods
Treatments with PGPR include drench application
(Babalola et al. 2007b) and seed bacterization
(Babalola et al. 2007a; Kumar et al. 2009). Also
documented are seedling treatment (Babalola et al.
2007a), bioformulation, biopreparation and dual
treatment (Lavania et al. 2006). Coating seeds with
commercial inoculants reduces inconsistency in the
field, that notwithstanding, field-tested commercial
formulations, mostly based on dry powder (charcoal,
lignite, farmyard manure, etc.) have inherent problems of appropriate shelf-life and cell viability.
Biological control
The potential of PGPR for biological control can
result from one or more mechanisms, including
antagonism, competition, and the production of
antibiotics or siderophores or by inducing disease
resistance and/or direct growth by improving nutrient
uptake for plants by the alteration of plant hormone
levels (Anith et al. 2004). Combining multiple PGPR
types can suppress disease development in many crop
plants and protect them against a broad range of soilborne plant pathogens. The strategy to study root
growth is essential as it helps in increasing root
branching, root mass, root length, and/or the amount
of root hairs and therefore the creation of a greater
root surface area for nutrient absorption. To effectively reduce disease symptoms, the bacteria must be
able to colonize the roots aggressively and have the

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potential to dominate the ecological niche. PGPR

B. pumilus SE 34, P. putida 89B61, BioYield and
Equity Biological control has emerged as one of the
important methods in the management of soil-borne
plant pathogens (Anith et al. 2004). The dependence
on chemicals for plant disease control could be
reduced to non-economic and environmentally
friendly level.
Laboratory, screenhouse and field studies
Striga hermonthica and Orobanche are two of the
most devastating parasitic weeds. They require specific host factors to break seed dormancy, thereby
ensuring the immediate availability of a host root.
The most common germination stimulants are
sesquiterpene lactones, collectively referred to as
strigolactones, including strigol. It was earlier hypothesized that Striga spp. has the same host range as
rhizospheric bacteria of the genus Azospirillum. In a
recent work, Babalola and colleagues (2007b)
describe the potential significance of PGPR for
agro-industry in promoting the growth of leguminous
crops with respect to their biomass and grain yield and
use of the PGPR to promote plant growth in Striga
infested soils and increase plant yield component.
Under optimal laboratory conditions, bacterial
cultures in a concentration equal to that in the
rhizosphere produce phytohormones to a level comparable to concentrations around field-grown maize
roots. In various studies, Pseudomonas, Klebsiella
oxytoca and Enterobacter sakazakki increased the
percentage germination of S. hermonthica over
the non-inoculated control (Babalola et al. 2007a).
The study demonstrated that there is some potential
in certain rhizosphere bacteria to induce the suicidal
germination of S. hermonthica.
Bacterial isolates were grown on Tryptone Soy
Agar (TSA) and Kings medium B (KB). All cultures
were grown for 48 h at 28C on appropriate agar
plates as stated above. Individual plates were harvested by adding 10 ml of 0.01 M phosphate buffer
(PBS, pH 7.0) solution and scrapping with a sterile
cotton swab. The harvested bacteria were pooled
together according to the bacterial type, washed
twice, and adjusted to the desired concentration by
optical density. The bacteria were serially diluted in
buffer and plate counts were made on the inoculum
prior to inoculation. The average colony forming

Biotechnol Lett (2010) 32:15591570

units (c.f.u.) from triplicate plates was determined by

counting after 48 h incubation, this was used to
express the quantity of inoculum added. A 250 ml,
24 h culture of each bacterial cell, free from culture
filtrate was drenched on the non-sterilized potted soil
in the screenhouse. Pots that received only water
served as the control. The results suggest that PGPR
were able to promote maize growth (Fig. 1) even in
S. hermonthica infested soil (Fig. 2). In the case of
Fig. 2, the soil was passed through a 10-mesh sieve

Fig. 1 A typical large pot screenhouse experiment with or

without added PGPR to maize planted in Striga-free soil.
Plantings were done in screenhouse potted soil. The soil was
not sterilized. A mixture of bacterial cells from 24-h-old
culture resuspended and washed clean of culture media were
drenched into the soil. Pots which did not receive bacterial
cells but only water, served as control

Fig. 2 A typical large pot screenhouse experiment of presence

or absence of added PGPR in maize planted in S. hermonthicainfested soil

1567

and thoroughly mixed with S. hermonthica seeds

within the first 010 cm depth at 120 seeds per kg
soil 14 days before planting. Growth promotion may
be as a result of the production of plant growth
producing substances, the suppression of deleterious
bacteria, improved soil structure due to PGPR, or the
alleviation of Striga induced stress on the maize host.
The yield effect of PGPR was plant cultivardependent (Babalola et al. 2007b) (Table 2). The
application of bacterial inoculants as biofertilizers
has improved the growth and yield of cereal crops
(Babalola et al. 2007b). All the bacteria affected crop
growth similarly, frequently increasing seed germination rates, elongating stems, enlarging leaves and
causing early flowering and fruiting effects indicative
of a common cause, related not to mineralization but
to growth regulators. The bacterial inoculants were
more effective when used in conjunction with
mineral and organic fertilizers (Kumar et al. 2009)
and worked best in soils supporting increased plant
and microbial growth. Tests by various workers for
the production of growth regulators by the bacteria
used as inoculants have shown that all of them
produce traces of IAA (Chakraborty et al. 2009) and
gibberellin-like substances, sufficient to cause
changes in plant morphology, for only trace amounts;
if applied at the right stage of plant development, will
change growth completely.
To study whether PGPR also has a beneficial
effect on plants under Striga infestation, the growth
patterns of maize were investigated. The PGPR
E. sakazakii and Pseudomonas spp. promoted a
significant increase in the associated trap crop,
cowpea (Vigna unguiculata), pod weight, fresh
biomass, pod number and pod wall thickness (Babalola et al. 2007b). The PGPR reduce the debilitating
effects of the root parasitic weed, S. hermonthica, on
host plant maize (Babalola et al. 2007a). The use of
bacteria for the biological control of S. hermonthica
is relatively new (Babalola et al. 2002). In the field,
results from the use of PGPR have been less
consistent and their ineffectiveness has often been
attributed to their inability to colonize plant roots.
This inconsistency in PGPR field results might be due
to many factors such as complex interactions among
hosts, rhizobacteria, pathogens, and climate. The
soils environmental conditions, e.g., texture, water
content, soil temperature and pH value could also
contribute to the inconsistency reported.

123

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Biotechnol Lett (2010) 32:15591570

Table 2 Effect of E. sakazakii 8MR5, Pseudomonas 44MS8, and Pseudomonas 10MS applied at planting on yield components of
cowpea sown in S. hermonthica-infested soil (from Babalola et al. 2007b)
Cowpea
variety
IT95K-286-4

Bacterial
isolatea

Pod
number

Pod
weight (g)

Fresh
biomass (g)

Dry
biomass (g)

13

55

43

464

71

45

13

41

31

361

50

46

12

47

22

400

60

47

12

50

30

363

58

43

SE

0.6

0.2

4.1

6.0

34.5

4.2

0.8

8MR5

13

23

24

420

72

47

44MS8

15

20

27

414

69

48

10M3

14

20

17

441

69

48

Controlb

13

16

11

536

89

49

SE

0.5

0.6

2.2

2.9

26.3

3.7

0.2

8MR5
44MS8

6
5

18
17

44
45

47
45

496
442

67
65

47
48

10M3

18

47

51

424

54

47

Controlb

17

19

21

268

56

47

SE

0.5

0.3

4.5

6.6

39.2

4.4

0.3

8MR5 E. Sakazakii (1.68 9 10 c.f.u.), 44MS8 Pseudomonas (1.33 9 10 c.f.u.) and 10M3 (1.10 9 10 c.f.u.)

Check without bacterial seed treatment

Conclusions and future perspectives

This review has shown that PGPR are free-living
bacteria of beneficial importance to agriculture and
abound in the rhizosphere, the region around the root.
Some PGPR have more than one mechanism of
accomplishing increased plant growth, such as the
production of enzymes, bioactive factors, antibiotics,
metabolites as well as growth promoters. The mechanisms of action presented are not an all-encompassing list. Although variability in field performance is
common, PGPR are environmentally friendly, unlike
the overuse of chemical fertilizers. Certain bacteria
containing ACC deaminase can show both pathogenic and growth-promoting properties in their
interaction with plants. Inconsistency in experimental
results lies as a result of fluctuations in environmental
conditions but role of biological factors cannot be
ruled out. Chemical fertilizers increase yield in
agriculture but are expensive and harm the environment. They deplete non-renewable energy via side
effects, such as leaching out, and polluting water
basins, destroying micro-organisms and friendly
insects, making the crop more susceptible to the
attack of diseases, reducing soil fertility, thereby
causing irreparable damage to the overall system.

123

Days to
flower

8MR5

Control

Rabuor (local)

Pod length
(cm)

44MS8
10M3

IT94D-437-1

Pod thickness
(cm)

The use of PGPR could be a better alternative to

chemical fertilizers. They are economical, not harmful to the environment and could easily be found.

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