Indian Journal of Biotechnology

Vol 9, October 2010, pp 397-402

Bioprospecting potential of fast growing endophytic bacteria from leaves of

mangrove and salt-marsh plant species
S Gayathri1, D Saravanan1, M Radhakrishnan1, R Balagurunathan2 and K Kathiresan3*
1
Department of Microbiology, Sri Sankara Arts & Science College, Kanchipuram 631 561, India
2
Department of Microbiology, Periyar University, Salem 636 011, India
3
Centre of Advanced Study in Marine Biology, Annamalai University, Parangipettai 608 502, India

Received 10 August 2009; revised 2 February 2010; accepted 9 April 2010

In the present study, bioprospecting aspects of endophytic bacteria, isolated from leaves of 5 mangroves and 2 salt-marsh
plant species, were explored. In total, 104 bacterial isolates were isolated from the leaf samples of mangrove plants of
Pichavaram, Tamil Nadu. Thirty six fast growing isolates were selected and screened for biological activities. Of 36 isolates,
28 (77%) showed antimicrobial activity, and 94.4% of isolates exhibited pectinase, 58.3% protease and 52.7% inulinase and
invertase activities. Plant growth promoting activity, such as, ammonia and acetoin production, was shown by 22 (61.1%)
and 25 (69.4%) endophyte isolates, respectively; whereas 26 isolates (72.2%) showed nitrogen fixing activity. Six isolates
(16.6%) showed phosphate solubilization activity and 7 isolates (19.4%) produced indole acetic acid (IAA). Further,
malachite green and phenol degrading activities were observed in 12 (33.3%) and 20 (55.5%) endophytic bacterial isolates,
respectively. More than 20 endophytic bacteria were sensitive to antibiotics like streptomycin and trimethoprim, while
31 isolates (86.1%) were resistant to vancomycin and bacitracin. Thirty four (94.4%) and 31 (86.6%) endophytic isolates
exhibited growth onto a medium containing upto 7.5 and 10% NaCl concentrations, respectively. The present study has
proved that the mangroves are the sources of endophytic bacteria with bioprospecting potential, which deserves further
studies.

Keywords: Biological activity, endophytic bacteria, mangroves, Pichavaram

Introduction used to quickly narrow down the search for

Microbes that colonize living, internal tissues of
plants without causing any immediate, over-negative
effects are termed as endophytes1. It is noteworthy
that, of the nearly 300,000 plant species that exist on
the earth, each individual is the host to endophytes2. It
is estimated that there may be as many as 1 million
different endophyte species; however, only handful of
them are described3. The growth stimulation of host
plant by the endophytic microbes can be a
consequence of nitrogen fixation, the production of
phytohormones, biocontrol of phytopathogens in the
root zone or by enhancing availability of nutrients and
minerals4. Recently, many new endophytic bioactive
metabolites, possessing a wide variety of biological
activities, have been reported. About 51% of
biologically active substances isolated from
endophytic fungi were unknown in the past2.
Since the number of plant species in the world is so
great, creative and imaginative strategies must be
——————
*Author for correspondence:
Mobile: 094420 68003; Fax: 04144-243555
Email: kathirsum@rediffmail.com
endophytes displaying activity. Several reasonable
hypotheses govern this plant selection strategy, which
includes plants from unique environmental settings,
especially those with an unusual biology having an
ethnobotanical history, or plants that are endemic that
has an unusual longevity, and plants growing in areas
of great biodiversity5.
Mangroves are salt tolerant plants existing at the
interface between land and sea in the tropical and
subtropical latitudes. The mangrove forests are one
among the world’s most productive ecosystems with
great ecological and economic significance6. Extracts
of mangroves have shown activity against human,
animal and plant pathogens7. Recent studies on
endophytic fungi and secondary metabolite, such as,
xyloketals, eniatin G and marinamide, have been
reported from mangroves8-10. Although the presence
of endophytic fungi in leaves of mangroves is
known11,12, their endophytic bacteria and their
bioprospecting potential is largely unknown for the
Indian mangroves. Hence, the present attempt was
made to isolate, characterize and explore the
398 INDIAN J BIOTECHNOL, OCTOBER 2010
biological activity of endophytic bacteria from
mangrove and salt-marsh plants.
Materials and Methods
Sample Collection
For the isolation of endophytic bacteria, healthy
leaf samples of five species of mangroves
[Rhizophora apiculata Blume, Avicennia marina
(Forssk.) Vierth., Excoecaria agallocha L., Ceriops
decandra (Griff.) Ding Hou & Aegiceras
corniculatum (L.) Blanco] and two species of salt-
marsh plants [Suaeda maritime Forssk. ex J. F. Gmel.
& Sesuvium portulacastrum (L.) L.] were collected
from mangrove forest, Pitchavaram (latitude 11°27’N;
longitude.79°47’E), Tamil Nadu. All the samples
were collected in sterile plastic bags and transported
aseptically to the laboratory. For the pretreatment of
leaf samples and isolation of endophytic bacteria, the
method described by Sun et al13 was adopted. Fresh
leaf samples were washed in running tap water,
followed by 2 min wash in 70% ethanol. Then the leaf
samples were washed in 2% sodium hypochlorite for
1 min. Finally, leafs were washed in distilled water
for 2 min and dried.
Isolation and Characterization of Endophytic Bacteria
After pretreatment, mangrove leaves were crushed
with sterile distilled water using mortar and pestle.
About 1 mL of crushed sample was serially diluted
and 0.1 mL of aliquot from 10-2 to 10-5 dilutions were
taken and spread onto nutrient agar medium using
sterile glass L-rod. Plating was done in duplicates and
all the plates were incubated at 28°C for 5 d. After
incubation, morphologically different bacterial
colonies were selected and streaked on nutrient agar
plates and incubated at 28°C for 48 h. From the total
isolates, based on the differences in cultural
morphology, such as, colour, texture, consistency and
size, limited numbers of representative isolates were
selected from all the samples for further
investigations. All the selected isolates were
subcultured in nutrient agar slants and preserved in a
refrigerator at 4°C. Phenotypic characteristics, such
as, Gram’s reaction, endospore staining, capsule
staining, motility, catalase and oxidase activity of all
the isolates were performed by adopting standard
procedures.
Screening of Endophytic Bacteria for Antimicrobial Activity
Antimicrobial substance from endophytic bacteria
was produced by submerged fermentation using No. 3
medium (polypeptone 1%; glucose 1%; potassium
dyhydrogen phosphate 0.1%; magnesium sulphate
0.05%; pH 7.0)14. About 10% of inoculum was
transferred into production medium and incubated in
centrifugation at 10,000 rpm for 10 min.
Antimicrobial activity of culture supernatant
(100 µL/well) was tested by agar well diffusion
method using Staphylococcus aureus, Bacillus sp.,
Escherichia coli, Vibrio parahaemolyticus,
V. anguillarum and Fusarium sp. as test organisms.
All the cultures were obtained from Christian Medical
College, Vellore. Cultures of V. parahaemolyticus and
V. anguillarum were obtained from Central Brackish
Water Aquaculture (CIBA), Chennai. Medium used
for V. parahaemolyticus and V. anguillarium was
supplemented with 2% NaCl. All the plates were
observed for zone of inhibition after incubation at
37°C for 24 h. In case of fungi, plates were incubated
at 28ºC for 3 d.
Screening of Endophytic Bacteria for Enzymatic Activity
All the endophytic bacterial isolates were screened
for various enzymes, such as, amylase, lipase,
protease, inulinase, invertase, chitinase, pectinase,
cellulase and urease by plate method14-17. All the
isolates are spot inoculated on respective enzyme
screening media and incubated at 28°C for 5-7 d.
Screening of Endophytic Bacteria for Production of Plant
Growth Promoting Substances
Nitrogen fixing activity of endophytic bacteria was
studied by inoculating them into Jenson’s medium
and incubating at 28°C for 5 d. Presence of growth on
Jenson’s medium indicates nitrogen fixation.
Phosphate solubilizing activity of endophytic bacteria
was studied by the method described by Pandey
et al18 using Pikovskaya’s agar medium. After
incubation, presence of clear halo around the growth
indicates phosphate solubilization. For the screening
of ammonia production, all the endophytic bacterial
isolates were inoculated into 5 mL of sterile peptone
broth separately and incubated at 28°C for 48 h. After
incubation, about 0.5 mL of Nessler’s reagent was
added into all the tubes. The production of ammonia
was indicated by the appearance of pink colour. The
Voges-Proskauer test was used as a qualitative
method for the detection of acetoin. Endophytic
cultures were inoculated into 2 mL of MR-VP broth
individually and incubated at 28ºC for 5 d. After
incubation, to 1 mL of bacterial culture, 3 mL of
freshly prepared 5% α-naphthol in absolute ethanol
and 1 mL of 40% KOH were added and the mixture
was stirred vigorously. The formation of a red colour
 GAYATHRI et al: MANGROVE ENDOPHYTIC BACTERIA 399

was indicative of the presence of acetoin19. For the of which 36 isolates were selected for further
screening of indole acetic acid (IAA), about 0.1 mL of investigations based on their fast growth.
18 h old culture was inoculated into each 5 mL of
nutrient broth supplemented with 3 mg/mL of L- Characterization of Endophytic Bacteria
tryptophan and incubated at 28°C for 5 d. After Of 36 isolates selected, 9 were pigmented and 27
centrifugation at 10,000 rpm for 10 min, a drop of were non-pigmented isolates. Regarding cell shape
orthophosphoric acid and 2 mL of Solawaski’s and Gram staining, 2 were Gram-negative cocci, 11
reagent was added into the supernatant and kept for Gram-negative rods, 3 Gram-positive cocci and 20
20 min. The development of pink colour indicated the Gram-positive rods. In total, 13 isolates showed
production of IAA20. positive result for endospore staining, possibly
belonging to the genus Bacillus.
Screening of Endophytic Bacteria for Biodegradation Activity
Endophytic bacteria were spot inoculated onto Antimicrobial Activity
screening medium supplemented with malachite green Regarding antimicrobial activity, 28 isolates (77%)
(0.01%) and incubated at 28°C for 5 d. Clear zone showed inhibitions against any one of the test strains.
around the bacterial spot indicated dye degradation21. Out of them, 20 (55.5%) isolates exhibited broad
For phenol degradation, the endophytic bacterial spectrum activity. Only 5 isolates were active against
isolates were spot inoculated on mineral agar medium Gram-positive bacteria and fungi, and 7 isolates were
supplemented with phenol (0.05%) as sole carbon active against Gram-negative bacteria. Number of
source. All the plates were incubated at 28ºC for 5 d. endophytic bacteria showing antimicrobial activity is
Presence of growth on the mineral agar medium given in Fig. 1.
indicated phenol degradation22. Table 1—Number of bacterial endophytes recovered from
coastal plants
Screening of Endophytic Bacteria for Salt (NaCl) Tolerance
Endophytic bacteria were inoculated onto Name of plant Pigmented Non- Total colonies
nutrient agar medium supplemented with different pigmented recovered
concentrations of NaCl (0-10%). All the plates were Mangroves
incubated at 28°C for 5 d and bacterial growth was Rhizophora apiculata 4 22 26
observed at every 24 h. Avicennia marina 5 12 17
Excoecaria agallocha 2 11 13
Screening of Endophytic Bacteria for Antibiotic Susceptibility Ceriops decandra 0 10 10
Antibiotic susceptibility pattern of endophytic Aegiceras corniculatum. 2 4 6
bacterial isolates against different antibiotic classes Salt-marshes
was determined by adopting Kirby-Bauer disc Suaeda maritima 2 14 16
Sesuvium portulacastrum 10 6 16
diffusion method23. Broth culture of endophytic
bacteria was prepared using nutrient agar and adjusted Total 23 81 104
to 0.5 McFarland standard. All the cultures were
inoculated into nutrient agar plates using sterile cotton
swab. Standard antibiotic discs, viz., streptomycin
(amino glycoside; 10 µg), trimethoprim (synthetic;
5 µg), vancomycin (glycopeptide; 30 µg) and
bacitracin (peptide; 10 U) were placed on nutrient
agar plates and incubated at 37°C for 24 h. After
incubation, antibiotic susceptibility pattern was
determined by measuring the zone of inhibition.

Results
Isolation of Eendophytic Bacteria
Nutrient agar plates inoculated with mangrove and
salt-marsh plant leaf samples showed morphologically
different bacterial colonies. In total, 104 bacterial Fig. 1—Number of endophytic bacteria showing antimicrobial
colonies were recovered from the samples (Table 1), activity
400 INDIAN J BIOTECHNOL, OCTOBER 2010
Enzymatic Activity
The enzymatic activity of endophytic isolates
revealed that majority of the isolates (94.4%)
exhibited pectinase activity, followed by protease
(58.3%), inulinase (52.7%) and invertase (52.7%).
The detailed results are shown in Fig. 2.
Production of Plant Growth Promoters
Of 36 endophytic isolates screened for plant
growth promoting substances, maximum of 22
(61.1%) and 25 (69.4%) isolates showed positive
results for ammonia and acetoin production,
respectively. Only 6 isolates showed phosphate
solubilization activity and 7 isolates exhibited IAA
production.
Biodegrading Activity
Of 36 isolates tested, 12 isolates (33.3%) showed
positive results for malachite green degradation and
20 isolates (55.5%) showed phenol degrading activity.
Antibiotic Susceptibility
Of the isolates studied for antibiotic susceptibility,
80.5, 63.8 and 86.1% of endophytic bacteria were
sensitive to streptomycin, trimethoprim and
vancomycin, respectively. About 31 (86.1%) isolates
were resistant to bacitracin. Antibiotic susceptibility
pattern of endophytic bacteria are given in Fig. 3.
Tolerance to Sodium Chloride
All the 36 endophytes showed good growth at 0 to
5% NaCl concentration. Thirty four (94.4%) isolates
grew upto 7.5% NaCl and 31 (86.1%) isolates
tolerated upto 10% NaCl concentration. Overall, the
growth rate of endophytes decreased with increasing
concentration of NaCl.
Fig. 2—Number of endophytic bacteria showing different
enzymatic activity
Discussion
In the present study, mangrove and salt-marsh
leaves yielded in total 104 bacterial colonies. Several
reports on endophytic fungi in mangroves8,9,24,
especially from the present study area of Pichavaram
mangrove forest, are available from R. apiculata,
R. mucronata, A. corniculatum, Avicennia marina,
A. officinalis, Brugulera cylindrica, C. decandra,
E. agallocha and Lumnitzera racemosa11,12. However,
there is no considerable report on endophytic bacteria
from the mangroves. In general, endophytic bacteria
occur at lower population densities than rhizospheric
bacteria25. Based on the visible morphological
differences, 36 bacterial isolates, in total, were
selected from seven plant samples for further
investigations.
Endophytes are the chemical synthesizers within
plants3. Many of them are capable of synthesizing
bioactive compounds that can be used by plants for
defense against pathogens and some of these
compounds have been proven useful for drug
discovery. Up to now, most of the natural products
discovered from endophytes are antibiotics,
anticancer agents, antivirals, antidiabetic agents and
other bioactive compounds having different functional
roles3. Mangrove endophytic fungi are known to
produce bioactive metabolites like anthraquinones,
eniatin G, xyloketals26. A great part of the compounds
are in the group of antibiotics; 66, 30 and 5% of the
compounds are inhibiting Gram-positive, Gram-
negative and mycobacteria, respectively. Majority of
the antibiotic compounds are active against Gram-
positive bacteria27. However, in the present study,
culture filtrate from 20 (55.5%) endophytic bacteria,
out of 36 studied, showed broad spectrum antibiotic
activity. Detection of active compounds from these
isolates and further studies are needed for their proper
utilization.
Fig. 3—Antibiotic susceptibility pattern of endophytic bacteria
 GAYATHRI et al: MANGROVE ENDOPHYTIC BACTERIA
Next to antibiotics, enzymes are the most important
products. In the present study, while screening the
endophytic bacterial isolates for 9 different
extracellular enzymes, 15 isolates showed protease,
inulinase, invertase, pectinase and cellulase activity;
13 isolates exhibited amylase and 12 isolates the
urease activity. Kumaresan and Suryanarayanan28
have reported the assemblage of endophytes in young,
mature and senescent leaves of R. apiculata and its
possible role in mangrove litter degradation. Maria
et al16 have also reported enzymatic activity of
mangrove-derived endophytic fungi. The results of
the present study showed the production of several
enzymes by endophyte bacteria which might play role
in mangrove litter degradation.
Endophytic bacteria residing within plant tissues
have been reported to be promoting the plant growth
directly or indirectly through production of
phytohormones, biocontrol of host plant diseases
and/or improvement of plant nutritional status18. They
possess the capacity to solubilize phosphates as
shown with the endophytic bacteria of soybean in
phosphate assimilation4. This is also evident in the
present study as endophytic bacterial isolates showed
four plant growth promoting activities, particularly
ammonia and acetoin production by 22 and 25
isolates, respectively. Volatile substances, such as,
2,3-butanediol and acetoin, produced by bacteria are
responsible for plant growth promotion, which is a
newly discovered mechanism29.
Endophytes are recently reported in the degradation
of environmental toxins. The bacteria degrading
recalcitrant compounds were found to be more
abundant among endophytic populations in the
rhizosphere of plants in contaminated soils30. The
improvement of phytoremediation of water soluble,
volatile organic pollutants by selected endophytic
bacteria was experimentally proved by Barac et al31.
In the present study, 12 and 20 isolates, out of 36
endophytic bacterial isolates, showed the degradation
of two important environmental pollutants malachite
green and phenol, respectively. Usually dye degrading
microorganisms were from dye contaminated soils.
Interestingly, in the present study, endophytic bacteria
from coastal plants showed malachite green
degradation. Further the dye degrading endophytes
isolated in this study will be a potential candidate
for dye degrading enzymes. Moreover, the detection
of recalcitrant degrading endophytic bacteria from
coastal plants will be a biological marker for
401
monitoring the pollution of coastal ecosystems by
recalcitrant molecules.
Emergence of antibiotic resistance among the
pathogenic microorganisms limits treatment options.
Antibiotic resistant genes, in addition to clinical
pathogens, are also present in environmental isolates
which are horizontally transferred to other
microorganisms. In the present study, antibiotic
susceptibility pattern of endophytic bacteria were
studied using four different antibiotics. Most of the
isolates (23-32) were sensitive to streptomycin,
trimethoprim and vancomycin, but 31 isolates, out of
36 studied, were resistant to bacitracin.
Occurrence of halophilic bacteria is well known in
coastal marine biotopes including mangrove and salt-
marsh ecosystems. But there is no report on halophilic
endophytes in the coastal plants other than the report
of Kamalraj et al32 who reported the effect of NaCl on
endophytic fungal assemblage in the leaves of a
mangrove C. roxburghiana. In that study, none of the
endophytic fungi showed growth above 300 mM NaCl
concentration. However, in the present study, salt
tolerance of endophytic bacteria was reported upto
10% NaCl concentration, but the growth rate of
endophytes decreased with the increase in the salt
concentration.
The coastal plants like mangroves and salt-marshes
have the great potential but are under-exploited source
for endophytic bacteria that produce substances of
potential antimicrobial, enzymatic, plant growth
promoting and biodegrading activities. Therefore,
further studies are needed for possible commercial
utility of these potential biochemicals.
Acknowledgement
The authors sincerely thank Professor K R
Venkatesan, Principal, Sri Sankara Arts & Science
College, Kanchipuram for his encouragement and
providing the research facilities. The authors also
thank the Director, CAS in Marine Biology,
Annamalai University and Vice-Chancellor and
Registrar of Periyar University, Salem for their
valuable help. One of the authors (RB) thanks DST,
Government of India (SR/SO/HS-42/2005), New
Delhi for financial assistance.
References
1 Bacon C W & White J F, Microbial endophytes (Marcel
Dekker Inc., New York) 2000, 341-388.
2 Strobel G & Daisy B, Bioprospecting of microbial
endophytes and their natural products, Microbiol Mol Biol
Rev, 67(2003) 491-502.
402 INDIAN J BIOTECHNOL, OCTOBER 2010
3 Guo B, Wang Y, Sun X & Tang K, Bioactive natural
products from endophytes: A review, Appl Microbiol
Biotechnol, 44, 2 (2008) 136-142.
4 Rosenblueth M & Romero E M, Bacterial endophytes and
their interactions with hosts, Mol Plant-Microbe Interact, 19
(2006) 827-837.
5 Balagurunathan R & Radhakrishnan M, Exploiting the less
explored—Microbial endophytes, Adv Biotechnol, 6 (2007)
20-23.
6 Kathiresan K, A review of studies on Pichavaram mangrove,
Southeast India, Hydrobiologia, 43 (2000) 185-205.
7 Bandaranayke W M, Bioactivities, bioactive compounds and
chemical constituents of mangrove plants, Wetlands Ecol
Manage, 10 (2002) 421-452.
8 Lin Y, Wu X, Feng S, Jiang G, Luo J et al, Five unique
compounds: Xyloketals from mangrove fungus Xylaria sp.
from the South China Sea coast, J Org Chem, 66 (2001)
6252-6256.
9 Lin Y C, Wang J & Wu X Y, A novel compound eniatin G
from the mangrove fungus Halosarpheia sp. (Strain 732)
from the South China Sea, Aust J Chem, 55 (2002) 225-227.
10 Feng Z H U & Youngcheng L, Marinamide, a novel alkaloid
and its methyl ester produced by the application of mixed
fermentation technique to two mangrove endophytic fungi
from the South China Sea, Chin Sci Bull, 51(2006)
1426-1430.
11 Suryanarayanan T S, Kumaresan V & Johnson J A, Foliar
fungal endophytes from two species of the mangrove
Rhizophora, Can J Microbiol, 44 (1998) 1003-1006.
12 Kumaresan V & Suryanarayanan T, Occurrence and
distribution of endophytic fungi in a mangrove community,
Mycol Res, 105 (2001) 1388-1391.
13 Sun L, Lu Z, Bie X, Lu F & Yang S, Isolation and
characterization of a co-producer of fengycins and surfactins,
endophytic Bacillus amyloliquefaciens ES-2, from
Scuttellaria baicalensis Georgi, World J Microbiol
Biotechnol, 22 (2006) 1259-1266.
14 Cho K M, Hong S Y, Lee S M, Kim Y H, Khang G G et al,
Endophytic bacterial communities in Ginseng and their
antifungal activity against pathogens, Microb Ecol, 54 (2007)
341-351.
15 Sahu M K, Sivakumar K & Kannan L, Degradation of
organic matters by the extracellular enzymes of
actinomycetes isolated from the sediments and molluscs of
the Vellar estuary, J Aqua Biol, 20 (2005) 142-144.
16 Maria G L, Sridhar K R & Raviraja N S, Antimicrobial and
enzyme activity of mangrove fungi of South West coast of
India, J Agric Technol, 1 (2005) 67-80.
17 Cheesbrough M, Medical laboratory manual (Tropical
Health Technology, Doddington, Cambridgeshire, England)
2003, 20-35.
18 Pandey A, Das N, Kumar B, Rinu K & Trivedi P, Phosphate
solubilization by Penicillium spp. isolated from soil samples
of Indian Himalayan region, World J Microbiol Biotechnol,
24 (2008) 97-102.
19 Houdt R V, Moons P, Buj M H & Michiels C W, N-acyl-L-
homoserine lactone quarum sensing controls butanediol
fermentation in Serratia plymuthica RVH1 and S.
marcescens MG1, J Bacteriol, 188 (2006) 1570-1572.
20 Hamdali H, Hafidi M, Virolle M J & Achdouch Y, Rock-
phosphate solubilizing actinomycetes: Screening for plant
growth promoting activities, World J Microbiol Biotechnol,
24 (2008) 2565-2575.
21 Gopalakrishnan V, Radhakrishnan M & Balagurunathan R,
Microbial degradation of textile dyes —An inventory, in
Proc Natl Symp on Recent Trends in Microbial
Biotechnology (Sri Sankara Arts & Science College, Tamil
Nadu) 2005, 57-64.
22 Rigo M & Alegre R M, Isolation and selection of phenol
degrading microorganisms from industrial wastewaters and
kinetics of the biodegradation, Folia Microbiol, 49 (2004)
41-45.
23 Bauer A W, Kirby M, Sherris J C & Turck M, Antibiotic
susceptibility testing by a standardized single disc method,
Am J Clin Pathol, 45 (1966) 493-497.
24 Chen G Y, Lin Y C & Wen L, Two new metabolites of a
marine endophytic fungus (No. 1893) from an estuarine
mangrove on the South China Sea coast, Tetrahedron, 59
(2003) 4907-4909.
25 Hallmann J, Hallmann A Q, Mahaffee W F & Kloepper J W,
Bacterial endophytes in agricultural crops, Can J Microbiol,
43 (1997) 895-914.
26 Cheng Z S, Pan J H, Tang W C, Chen Q C & Lin Y C,
Biodiversity and biotechnological potential of mangrove-
associated fungi, J For Res,20 (2009) 63-72.
27 Berdy J, Bioactive microbial metabolites, J Antibiot, 58
(2005) 1-28.
28 Kumaresan V & Suryanarayanan T S, Endophyte assemblage
in young, mature and senescent leaves of Rhizophora
apiculata: Evidence for the role of endophytes in mangrove
litter degradation, Fungal Diversity, 9 (2002) 81-91.
29 Ryu C, Farag M A, Hu C, Reddy M S, Wei H et al, Bacterial
volatile promotes growth in Arabidopsis, Proc Natl Acad Sci
USA, 100 (2003) 4927-4932.
30 Siciliano S D, Fortin N, Mihoc N, Wisse G, Labelle S et al,
Selection of specific endophytic bacterial genotypes by
plants in response to soil contamination, Appl Environ
Microbiol, 67 (2001) 2469-2475.
31 Barac T, Engineered endophytic bacteria improve
phytoremediation of water soluble, volatile, organic
pollutants, Nat Biotechnol, 22 (2004) 583-588.
32 Kamalraj S, Sridevi S, Gangadevi V, Venkatesan A &
Muthumary J, Effect of NaCl on biochemical changes and
endophytic assemblages in the leaves of a mangrove, Ceriops
roxburghiana Arn., Indian J Sci Technol, 1 (2008) 1-7.