MINIREVIEW

Bacterial endophytes: recent developments and applications

Robert P. Ryan1, Kieran Germaine2, Ashley Franks1, David J. Ryan2 & David N. Dowling2
1
BIOMERIT Research Centre, Department of Microbiology, Biosciences Institute, National University of Ireland, Cork, Ireland; and 2Department of
Science & Health, Institute of Technology Carlow, Carlow, Ireland

Correspondence: David N. Dowling, Abstract

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Department Science & Health, Institute of
Technology Carlow, Kilkenny Road, Carlow,
Endophytic bacteria have been found in virtually every plant studied, where they
Ireland. Tel.: 1353 59 9170479; fax: 1353 59 colonize the internal tissues of their host plant and can form a range of different
9170517; e-mail: dowlingd@itcarlow.ie relationships including symbiotic, mutualistic, commensalistic and trophobiotic.
Most endophytes appear to originate from the rhizosphere or phyllosphere; however,
Received 9 August 2007; accepted some may be transmitted through the seed. Endophytic bacteria can promote plant
14 August 2007. growth and yield and can act as biocontrol agents. Endophytes can also be beneficial
First published online December 2007. to their host by producing a range of natural products that could be harnessed for
potential use in medicine, agriculture or industry. In addition, it has been shown
DOI:10.1111/j.1574-6968.2007.00918.x
that they have the potential to remove soil contaminants by enhancing phytoreme-
diation and may play a role in soil fertility through phosphate solubilization and
Editor: Richard Staples
nitrogen fixation. There is increasing interest in developing the potential biotechno-
Keywords
logical applications of endophytes for improving phytoremediation and the sustain-
bioremediation; plant–microbe interaction; able production of nonfood crops for biomass and biofuel production.
biocontrol; plant growth promotion.

Introduction Gnanamanickam, 1997), insects (Azevedo et al., 2000) and

nematodes (Hallmann et al., 1997, 1998). In some cases, they
Beneficial plant–microbe interactions that promote plant can also accelerate seedling emergence, promote plant estab-
health and development have been the subject of consider- lishment under adverse conditions (Chanway, 1997) and
able study. Recent work has also investigated their potential enhance plant growth (Bent & Chanway, 1998). Bacterial
for the enhanced biodegradation of pollutants in soil. Most endophytes have been shown to prevent disease development
of these studies have focused on bacteria from the rhizo- through endophyte-mediated de novo synthesis of novel
sphere of plants (Lindow & Brandl, 2003; Kuiper et al., 2004; compounds and antifungal metabolites. Investigation of the
Berg et al., 2005). Endophytic bacteria can be defined as biodiversity of endophytic strains for novel metabolites may
those bacteria that colonize the internal tissue of the plant identity new drugs for effective treatment of diseases in
showing no external sign of infection or negative effect on humans, plants and animals (Strobel et al., 2004).
their host (Holliday, 1989; Schulz & Boyle, 2006), and of the Along with the production of novel chemicals, many
nearly 300 000 plant species that exist on the earth, each endophytes have shown a natural capacity for xenobiotic
individual plant is host to one or more endophytes (Strobel degradation or may act as vectors to introduce degradative
et al., 2004). Only a few of these plants have ever been traits. The ability of some endophytes to show resistance to
completely studied relative to their endophytic biology. heavy metals/antimicrobials and degrade organic com-
Consequently, the opportunity to find new and beneficial pounds probably stems from their exposure to diverse
endophytic microorganisms among the diversity of plants in compounds in the plant/soil niche. This natural ability to
different ecosystems is considerable. degrade these xenobiotics is being investigated with regard
Bacterial endophytes colonize an ecological niche similar to improving phytoremediation (Siciliano et al., 2001; Barac
to that of phytopathogens, which makes them suitable as et al., 2004; Germaine et al., 2004, 2006; Porteous-Moore
biocontrol agents (Berg et al., 2005). Indeed, numerous et al., 2006; Ryan et al., 2007a). This review aims to provide
reports have shown that endophytic microorganisms can an overview of the potential applications for bacterial
have the capacity to control plant pathogens (Sturz & endophytes particularly in the area of phytoremediation
Matheson, 1996; Duijff et al., 1997; Krishnamurthy & and sustainable agriculture.

FEMS Microbiol Lett 278 (2008) 1–9

c 2007 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
2 R.P. Ryan et al.

Isolation and biodiversity of bacterial Studies that make use of both culture-based and culture-
endophytes independent techniques can be particularly useful. The
presence and taxonomy of endophytic bacteria of the entire
The endophytic niche offers protection from the environ- aerial parts of Crocus (Crocus albiflorus) was investigated by
ment for those bacteria that can colonize and establish Reiter & Sessitsch (2006). Their results suggest that Crocus
in planta. These bacteria generally colonize the intercellular supports a diverse bacterial microbial communities resem-
spaces, and they have been isolated from all plant compart- bling the microbial communities that have been described
ments including seeds (Posada & Vega, 2005). Endophytic for other plants, but also containing species that have not
bacteria have been isolated from both monocotyledonous been described in association with plants before. A combi-
and dicotyledonous plants, ranging from woody tree species, nation of plating of plant macerates, isolation and 16S rRNA
such as oak and pear, to herbaceous crop plants such as gene sequence identification of isolates and whole-commu-
sugar beet and maize. Classical studies on the diversity of nity fingerprinting was used. The results clearly indicated

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bacterial endophytes have focused on characterization of that a wide range of bacteria from diverse phylogenetic
isolates obtained from internal tissues following disinfection affiliation, mainly Gammaproteobacteria and Firmicutes, live
of plant surfaces with sodium hypochlorite or similar agents in association with plants of Crocus albiflorus. The commu-
(Miche & Balandreau, 2001). A review by Lodewyckx et al. nity composition of the culturable component of the micro-
(2002) highlights the methods used to isolate and character- bial communities was different from that of the 16S rRNA
ize endophytic bacteria from different plant species. A very gene clone library. Only three bacterial divisions were found
comprehensive list of bacterial endophytes isolated from in the culture collection, which represented 17 phylotypes,
a broad range of plants is provided by Rosenblueth & whereas six divisions were identified in the library analysis
Martinez-Romero (2006) and Berg & Hallmann (2006), comprising 38 phylotypes. The predominant group in the
which updates the groundwork laid by Hallmann et al. culture collection was the low G1C Gram-positive group,
(1997) and Lodewyckx et al. (2002). whereas in the clone library, the Gammaproteobacteria pre-
A recent study by Porteous-Moore et al. (2006) describes dominated. This study confirms that the culturable endo-
the diversity of endophytes found in poplar trees, growing at phytes are a subset of total endophyte biodiversity.
a phytoremediation field site contaminated with toluene,
with the aim of identifying potential candidates for enhancing
Plant colonization and inoculation with
phytoremediation of toluene, ethylbenzene, and xylene
(BTEX) compounds. Endophytic bacteria were isolated from
endophytes
two varieties of Poplar. The isolated endophytic bacteria were Autofluorescent protein (AFP) methods are now a key tool
characterized by comparative sequence analysis of partial 16S for studying processes such as microbe–plant interactions
RNA genes, BOX-PCR profiling of genomic DNA and and biofilm formation (for a recent review, see Larrainzar
physiological characterization, with respect to substrate utili- et al., 2005). These techniques have been utilized to detect
zation, antibiotics and heavy-metal sensitivities. This study and enumerate microorganisms in situ on plant surfaces
and those of Germaine et al. (2004, 2006) demonstrated that and in planta (Gage et al., 1996; Tombolini et al., 1997;
within the diverse bacterial communities found in Poplar Tombolini & Jansson, 1998). One of these AFP strategies
trees, several endophytic strains were present that had the uses a marker system, which encodes the green fluorescent
potential to enhance phytoremediation of volatile organics protein (GFP). This technique has shown promise in
and herbicides. monitoring pseudomonads in root tissues (Tombolini
Molecular approaches for the isolation and characteriza- et al., 1997; Tombolini & Jansson, 1998). GFP is a useful
tion of bacterial endophytes and plant-associated bacteria AFP biomarker because it does not require any substrate or
and communities have been reviewed recently by Franks cofactor in order to fluoresce. Workers have developed GFP
et al. (2006). Microbial communities inhabiting stems, roots cassettes for chromosomal integration and expression of gfp
and tubers of various varieties of plants were analysed by 16S in a variety of bacteria (Tombolini et al., 1997; Tombolini
rRNA gene-based techniques such as terminal restriction & Jansson, 1998; Xi et al., 1999). Bacterial cells with
fragment length polymorphism analysis, denaturing gradi- chromosomal integration of gfp can be identified by
ent gel electrophoresis as well as 16S rRNA gene cloning and epifluorescence microscopy or confocal laser scanning
sequencing. Five taxa exhibiting the most promising levels microscopy (Villacieros et al., 2003; Germaine et al., 2004).
of colonization and an ability to persist were identified as Germaine et al. (2004) investigated a number of GFP-
Cellulomonas, Clavibacter, Curtobacterium, Pseudomonas labelled Poplar endophytes for their colonization ability and
and Microbacterium by 16S rRNA gene sequence, fatty acid also explored different methods of inoculation; a simple
and carbon source utilization analyses (Elvira-Recuenco & ‘stick dipping’ method was found to be very efficient,
van Vuurde, 2000; Zinniel et al., 2002). leading to extensive colonization of the specific tissues of

c 2007 Federation of European Microbiological Societies FEMS Microbiol Lett 278 (2008) 1–9
Published by Blackwell Publishing Ltd. All rights reserved
Bacterial endophytes 3

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Fig. 1. Schematic diagram of the different plant–bacterial endophyte interactions that have been studied and their applications

Poplar plants (see Fig. 1) at levels of 102 to 104 CFU g 1

tissue depending on the strain, whereas a seed ‘imbibing’
method was efficient for inoculation of pea plants
(Germaine et al., 2006; Germaine, 2007) (Fig. 2).
Endophyte colonization has also been visualized with the
use of the b-glucuronidase (GUS) reporter system. A GUS-
marked strain of Herbaspirillum seropedicae Z67 was inocu-
lated onto rice seedlings. GUS staining was most intense on
coleoptiles, lateral roots, and also at some of the junctions of
the main and lateral roots (James et al., 2002). This study by
James et al. (2002) showed that endophytes entered the roots
through cracks at the point of lateral root emergence.
Herbaspirillum seropedicae subsequently colonized the root
intercellular spaces, aerenchyma and cortical cells, with a few
penetrating the stele to enter the vascular tissue. The xylem
vessels in leaves and stems were also colonized.
Successful endophyte colonization also involves a compa-
tible host plant. Recently, work by Miche et al. (2006)
investigated the obligate nitrogen-fixing endophyte Azoarcus
sp. strain BH72, which expresses nitrogenase (nif) genes
inside rice roots. They used a proteomic approach to dissect
responses of rice roots toward bacterial colonization and
jasmonic acid treatment (which induces plant defence pro-
teins). Data suggest that induced plant defence responses may
contribute to restricting endophytic colonization in grasses.
Fig. 2. Application of AFPs for studying bacterial endophyte–plant
Few studies have been published describing the molecular
interactions. (a) Xylem tracheid pits of inoculated poplar trees showing
basis of the interactions between endophytic bacteria and colonization by Pseudomonas putida VM1453 cells (  1000) (Germaine
plants. Adapting strategies that have been used to study et al., 2004). Scale bar = 10 mM. (b) Pseudomonas putida VM1450 micro-
bacterial gene expression in the rhizosphere and phyllo- colony within the root cortex of inoculated pea plants exposed to 54 mg
sphere such as in vivo expression technology (IVET) and 2,4-D (  1000) (Germaine et al., 2006).

FEMS Microbiol Lett 278 (2008) 1–9

c2007 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
4 R.P. Ryan et al.

recombination in vivo expression technology (Leveau & include osmotic adjustment, stomatal regulation, modifica-
Lindow, 2001; Preston et al., 2001; Zhang et al., 2006) may tion of root morphology, enhanced uptake of minerals and
provide an insight into genes that are required by bacteria to alteration of nitrogen accumulation and metabolism (Com-
enter, compete, colonize the plant, suppress pathogens and pant et al., 2005a, b). The recent areas where these plant
generally survive within the plant. growth-promoting bacterial endophytes are being used are
in the developing areas of forest regeneration and phytor-
Genomics of endophyte bacteria emediation of contaminated soils.
To date, few endophytic bacterial genome sequences have
been published; however, genome sequencing of a number Biocontrol and endophytes
of endophytes including Enterobacter sp.638, Stenotropho-
Endophytic bacteria are able to lessen or prevent the
monas maltophilia R551-3, Pseudomonas putida W619,
deleterious effects of certain pathogenic organisms. The

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Serratia proteamaculans 568 and Methylobacterium populi
beneficial effects of bacterial endophytes on their host plant
BJ001 is underway at the United States Department of
appear to occur through similar mechanisms as described
Energy Joint Genome Institute (www.jgi.doe.gov). Recently,
for rhizosphere-associated bacteria. These mechanisms have
the complete genome sequence of the nitrogen-fixing en-
been reviewed in great detail by Kloepper et al. (1999) or,
dophyte, Azoarcus sp. strain BH72 (Hurek & Reinhold-
more recently, by Gray & Smith (2005) and Compant et al.
Hurek, 2003; Krause et al., 2006) has been compared with
(2005a). Diseases of fungal, bacterial, viral origin and in
that of the related soil bacterium strain Azoarcus sp. strain
some instances even damage caused by insects and nema-
EbN1 and other plant-associated bacteria. The BH72 gen-
todes can be reduced following prior inoculation with
ome lacks genes encoding type III and type IV secretion
endophytes (Kerry, 2000; Sturz et al., 2000; Ping & Boland,
systems, toxins, nodulation factors, common enzymes that
2004; Berg & Hallmann, 2006).
hydrolyses plant cell walls and the N-acyl homoserine
It is believed that certain endophyte bacteria trigger
lactone-based quorum-sensing system, which is found
a phenomenon known as induced systemic resistance (ISR),
in many plant-associated bacteria and plant pathogens
which is phenotypically similar to systemic-acquired resis-
(Rainey, 1999; Preston et al., 2001; Büttner & Bonas, 2006).
tance (SAR). SAR develops when plants successfully activate
However, Krause et al. (2006) identified other factors
their defence mechanism in response to primary infection by
encoded by the BH72 genome that may be involved in the
a pathogen, notably when the latter induces a hypersensitive
host interaction. These include type IV pili, surface poly-
reaction through which it becomes limited in a local necrotic
saccharides, type I and II protein secretion systems, flagella
lesion of brown desiccated tissue (van Loon et al., 1998). ISR
and chemotaxis proteins and a large number of ferric-
is effective against different types of pathogens but differs
siderophore uptake systems. The BH72 genome provides
from SAR in that the inducing bacterium does not cause
valuable insights into the biology of bacterial endophytes,
visible symptoms on the host plant (van Loon et al., 1998).
and as more endophyte genome sequences become available,
Bacterial endophytes and their role in ISR have been
this will provide a rational basis to design experiments
reviewed recently by Kloepper & Ryu (2006).
to investigate the mechanisms involved in successful
endophyte colonization.
Natural products from endophytic
Plant growth-promoting endophytes bacteria
Research has been conducted on the plant growth-promot- Many endophytes are members of common soil bacterial
ing abilities of various rhizobacteria. They differ from genera, such as Pseudomonas, Burkholderia and Bacillus
biocontrol strains in that they do not necessarily inhibit (Lodewyckx et al., 2002). These genera are well known for
pathogens but increase plant growth through the improved their diverse range of secondary metabolic products includ-
cycling of nutrients and minerals such as nitrogen, phos- ing antibiotics, anticancer compounds, volatile organic
phate and other nutrients. compounds, antifungal, antiviral, insecticidal and immuno-
Endophytes also promote plant growth by a number of suppressant agents. While a wide range of biologically active
similar mechanisms. These include phosphate solubilization compounds have been isolated from endophytic organisms,
activity (Verma et al., 2001; Wakelin et al., 2004), indole they still remain a relatively untapped source of novel
acetic acid production (Lee et al., 2004) and the production natural products.
of a siderophore (Costa & Loper, 1994). Endophytic organ- While most research has focused on fungal-based pro-
isms can also supply essential vitamins to plants (Pirttila duction of antimicrobial products, a number of low-mole-
et al., 2004). Moreover, a number of other beneficial effects cular-weight compounds, active at low concentrations
on plant growth have been attributed to endophytes and against a range of human, animal and plant pathogenic

c 2007 Federation of European Microbiological Societies FEMS Microbiol Lett 278 (2008) 1–9
Published by Blackwell Publishing Ltd. All rights reserved
Bacterial endophytes 5

Table 1. Natural products that have been derived or produced from various endophytic bacteria
Organism Plant association Active agent Activity Reference
Taxomyces andreanae Taxus brevifolia Taxol Anticancer Strobel et al. (1993)
Pseudomonas viridiflava Grass Ecomycins B and C Antimicrobial Miller et al. (1998)
Streptomyces griseus Kandelia candel p-Aminoacetophenonic acids Antimicrobial Guan et al. (2005)
Streptomyces NRRL 30562 Kennedia nigriscans Munumbicins Antibiotic Castillo et al. (2002)
Munumbicin D Antimalarial
Streptomyces NRRL 30566 Grevillea pteridifolia Kakadumycins Antibiotic Castillo et al. (2003)
Serratia marcescens Rhyncholacis penicillata Oocydin A Antifungal Strobel et al. (2004)
Paenibacillus polymyxa Wheat Fusaricidin A–D Antifungal Beck et al. (2003)
Lodge pine Li et al. (2007)
Green beans

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Arabidopsis thaliana
Canola Beatty & Jensen (2002)
Cytonaema sp. Quercus sp. 103 Cytonic acids A and D Antiviral Guo et al. (2000)
Streptomyces sp. Monstera sp. Coronamycin Antimalarial antifungal Ezra et al. (2004)

bacteria, have been isolated from bacterial endophytes Endophytic microorganisms with the
(Table 1). One member of the plant-associated fluorescent potential to improve phytoremediation
pseudomonads, Pseudomonas viridiflava, which has
been isolated on and within the tissues of many grass Siciliano et al. (2001) showed that plants grown in soil
species (Miller et al., 1998), was found to produce two contaminated with xenobiotics naturally recruited endo-
novel antimicrobial compounds called ecomycins. Ecomy- phytes with the necessary contaminant-degrading genes.
cins represent a family of novel lipopeptides and are Indeed, in field sites contaminated with nitro-aromatics,
made up of some unusual amino acids including homo- genes encoding for nitro-aromatic compound degradation
serine and b-hydroxy aspartic acid. It was found that these were more prevalent in endophytic strains than within
compounds were able to inhibit the human pathogens rhizospheric or soil microbial communities. Van Aken et al.
Cryptococcus neoformans and Candida albicans (Miller (2004) also showed that a phyto-symbiotic strain of
et al., 1998). While viral inhibitors have been isolated Methylobacterium, which was isolated from hybrid Poplar
from Cytomaema sp. of fungi, such as cytonic acids A and trees (Populus deltoids x nigra), was capable of biodegrading
D, these were found to inhibit the human cytomegalovirus numerous nitro-aromatic compounds such as 2,4,6-trinitro-
(Guo et al., 2000). However, comprehensive screens for toluene. In the absence of a natural biodegradation ability,
antiviral compounds from bacterial endophytes have yet genetically engineered strains can be constructed and tailor-
to be reported. made for the desired application. Lodewyckx et al. (2001),
Bioplastics are biomaterials that are receiving increasing demonstrated that endophytes of yellow lupin, genetically
commercial interest. Lemoigne (1926) first described a constructed for nickel resistance, were able to increase the
bioplastic, poly-3-hydroxybutyrate (PHB) produced by nickel accumulation and tolerance of inoculated plants.
Bacillus megaterium. They are polyesters, produced by An application of bacterial endophytes with considerable
a range of microorganisms cultured under different nutrient biotechnological potential was described by Barac et al.
and environmental conditions. The most widely produced (2004), who showed that engineered Burkholderia cepacia
microbial bioplastics are poly-3-hydroxyalkanoate (PHA) G4 could increase plant tolerance to toluene, and decrease
and PHB. Genomic analysis indicates that many species the transpiration of toluene to the atmosphere. Because
of bacteria have the potential to produce bioplastics (Kalia toluene is one of the four components of BTEX contamina-
et al., 2003). tion, this has the potential to improve phyto-remediation by
Herbaspirillum seropedicae, a diazotrophic endophyte, can decreasing toxicity and increasing degradation of the xeno-
colonize a variety of higher plants and utilize a diverse range biotic (Barac et al., 2004). Table 2 outlines a number of
carbon sources. Catalán et al. (2007) have shown that studies that demonstrate the potential role of endophytes in
H. seropedicae accumulates significant levels of PHB, when phytoremediation.
grown on a range of individual carbon sources. The design Germaine et al. (2006) inoculated pea plants with a
and development of bacteria and higher plants able to Pseudomonas endophyte capable of degrading the organo-
accumulate PHAs may also help to streamline cost-effective chlorine herbicide, 2,4-dichlorophenoxyacetic acid (2,4-D).
production and to produce novel heteropolymers for When inoculated plants were exposed to 2,4-D, they showed
a range of applications (Aldor & Keasling, 2003). no accumulation of the herbicide into their tissues and

FEMS Microbiol Lett 278 (2008) 1–9

c2007 Federation of European Microbiological Societies
Published by Blackwell Publishing Ltd. All rights reserved
6 R.P. Ryan et al.

Table 2. A non-exhaustive list of pollutants that have been associated with bacterial endophyte phyto-remediation strategies
Compound Plant association Organism Reference
Mono- and dichlorinated Wild rye (Elymus dauricus) Pseudomonas aeruginosa strain R75 Siciliano et al. (1998)
benzoic acids and Pseudomonas savastanoi strain CB35
2,4-D Poplar (Populus) and willow (Salix) P. putida VM1450 Germaine et al. (2006)
Methane Poplar tissues Methylobacterium populi BJ001 Van Aken et al. (2004)
(Populus deltoidesnigra DN34)
TNT, RDX, HMX Poplar tissues Methylobacterium populi BJ001 Van Aken et al. (2004)
(Populus deltoidesnigra DN34)
MTBE, BTEX, TCE Populus cv. Hazendans and Pseudomonas sp Germaine et al. (2004),
cv. Hoogvorst Porteous-Moore et al. (2006)
Toluene Poplar (Populus) B. cepacia Bu61(pTOM-Bu61) Taghavi et al. (2005)

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TCP and PCB Wheat Herbaspirillum sp. K1 Mannisto et al. (2001)
Volatile organic compounds Yellow lupine (Lupinus luteus L.) Burkholderia cepacia G4 Barac et al. (2004)
and toluene

TNT, 2,4,6-trinitrotoluene; 2,4-D, 2,4-dichlorophenoxyacetic acid; TNT, 2,4,6-trinitrotoluene; RDX, hexahydro-1,3,5-trinitro-1,3,5-triazene; HMX,
octahydro-1,3,5,7-tetranitro-1,3,5-tetrazocine; NDAB, aliphatic nitramine 4-nitro-2,4-diazabutanal; BTEX, benzene, toluene, ethylbenzene, and
xylene; TCP, 2,3,4,6-tetrachlorophenol; PCB, polychlorinated biphenyl.

experienced little or no signs of phytotoxicity, whereas populations with the capacity to degrade a pollutant and does
uninoculated plants showed significant accumulation of not require long-term establishment of the inoculant strain.
2,4-D and displayed signs of toxicity including a reduction It is has been demonstrated that endophytic bacteria
in biomass, leaf abscission and callus development on their efficiently expressing the necessary catabolic genes can
roots. Large rhizosphere populations were also observed promote the degradation of xenobiotic compounds or their
during these experiments, which were responsible for en- metabolites as they are accumulated or while being translo-
hanced degradation of 2,4-D in soil. cated in the vascular tissues of the host plant. With
The possible advantages of using endophytic microorgan- phytoremediation playing an ever-increasing role in the
isms to improve xenobiotic remediation were summarized clean-up of contaminated land and water, it is envisaged
by Newman & Reynolds (2005), a major advantage being that endophytes will play a major role in enhancing both the
where genetic engineering of a xenobiotic degradation path- range of contaminants that can be remediated and the rate
way is required, bacteria are easier to manipulate than plants. of their degradation.
In addition, quantitative gene expression of pollutant
catabolic genes within the endophytic populations could be
The endophyte niche and emerging
a useful monitoring tool for assessing the efficiency of the
remediation process. The unique niche of the interior plant
pathogens
environment provides the xenobiotic degrader strain with an Various opportunistic bacterial pathogens including
ability to reach larger population sizes due to reduced Burkholderia, Enterobacter, Herbaspirillum, Ochrobactrum,
competition. Another important advantage of using endo- Pseudomonas, Ralstonia, Staphylococcus and Stenotrophomo-
phytic pollutant degraders is that any toxic xenobiotics taken nas have been identified as colonizers of the plant rhizo-
up by the plant may be degraded in planta, thereby reducing sphere (Berg et al., 2005). Many facultative endophytes are
phytotoxic effects and eliminating any toxic effects on recruited from the large pool of soil and rhizospheric species
herbivorous fauna residing on or near contaminated sites. and bacteria adapted for living in planta may include
The endophyte niche is a hot spot for horizontal gene opportunistic human/animal pathogens. Some bacteria as-
transfer (HGT) as demonstrated by Taghavi et al. (2005). In sociated with human infections have been isolated from the
this study, the degradative plasmid, pTOM-Bu61, was found interior of alfalfa (Ponka et al., 1995). This research area
to have transferred naturally to a number of different needs further investigation to establish the risks, if any,
endophytes in planta. This HGT activity promoted more associated with the development of the endophytic niche
efficient degradation of toluene in poplar plants. HGT for biotechnological applications.
in planta is likely to be widespread, as studies in pea with
Pseudomonas endophytes harbouring the plasmids pWWO
and pNAH7 also show high rates of transfer into a range of
Concluding remarks
autochthonous endophytes (Ryan et al., 2007b; E. Keogh & Exploitation of endophyte–plant interactions can result in
D.N. Dowling, unpublished data). This approach may have the promotion of plant health and can play a significant role
practical applications in equipping the natural endophyte in low-input sustainable agriculture applications for both

c 2007 Federation of European Microbiological Societies FEMS Microbiol Lett 278 (2008) 1–9
Published by Blackwell Publishing Ltd. All rights reserved
Bacterial endophytes 7

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Acknowledgements
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