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Article history:
Received 1 June 2009
Received in revised form 12 December 2009
Accepted 3 February 2010
Available online 19 February 2010
Keywords:
Marine
Biosurfactant
Bioemulsier
Exopolysaccharides
Glycolipids
Lipopeptides
Applications
a b s t r a c t
Marine biosphere offers wealthy ora and fauna, which represents a vast natural resource of imperative
functional commercial grade products. Among the various bioactive compounds, biosurfactant (BS)/
bioemulsiers (BE) are attracting major interest and attention due to their structural and functional
diversity. The versatile properties of surface active molecules nd numerous applications in various
industries. Marine microorganisms such as Acinetobacter, Arthrobacter, Pseudomonas, Halomonas, Myroides,
Corynebacteria, Bacillus, Alteromonas sp. have been studied for production of BS/BE and exopolysaccharides
(EPS). Due to the enormity of marine biosphere, most of the marine microbial world remains unexplored.
The discovery of potent BS/BE producing marine microorganism would enhance the use of environmental
biodegradable surface active molecule and hopefully reduce total dependence or number of new application
oriented towards the chemical synthetic surfactant industry. Our present review gives comprehensive
information on BS/BE which has been reported to be produced by marine microorganisms and their possible
potential future applications.
2010 Elsevier Inc. All rights reserved.
Contents
1.
2.
Introduction . . . . . . . . . . . . . . . . . . . .
Marine microorganisms The rich natural resource of
2.1.
Polymeric biosurfactant/bioemulsiers . . . .
2.1.1.
Acinetobacter . . . . . . . . . . . .
2.1.2.
Pseudomonas . . . . . . . . . . . .
2.1.3.
Myroides . . . . . . . . . . . . . .
2.1.4.
Halomonas . . . . . . . . . . . . .
2.1.5.
Yeast . . . . . . . . . . . . . . . .
2.1.6.
Streptomyces . . . . . . . . . . . .
2.1.7.
Antarctobacter . . . . . . . . . . . .
2.1.8.
Marinobacter . . . . . . . . . . . .
2.2.
Glycolipid surface active molecules . . . . . .
2.2.1.
Alcaligenes sp. . . . . . . . . . . . .
2.2.2.
Arthrobacter . . . . . . . . . . . . .
2.2.3.
Alcanivorax borkumensis . . . . . . .
2.2.4.
Rhodococcus. . . . . . . . . . . . .
2.2.5.
Halomonas . . . . . . . . . . . . .
2.2.6.
Unidentied marine bacterium MM1 .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
biosurfactant (BS)/bioemulsier (BE), exopolysaccharides (EPS) producers
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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Corresponding author. Institute of Bioinformatics and Biotechnology and Professor of Microbiology, University of Pune, Pune 411007, Maharashtra, India. Tel.: + 91 2025691333;
fax: + 91 2025690087.
E-mail address: directoribb@unipune.ernet.in (B.A. Chopade).
0734-9750/$ see front matter 2010 Elsevier Inc. All rights reserved.
doi:10.1016/j.biotechadv.2010.02.006
2.3.
2.4.
2.5.
2.6.
1. Introduction
The hydrosphere marine environment represents the major
component of the Earth's biosphere. It covers a majority (70%) part
of earth's surface and makes up N90% of the volume of its crust. The
maximum depth of N11,000 m and 365 million km2 is attributed to
the hydrosphere oceanic system. Oceans represent a vast and
exhaustive source of natural products in the globe, harbouring the
most diverse groups of ora and fauna. Marine microorganisms have
developed unique metabolic and physiological capabilities to thrive in
extreme habitats and produce novel metabolites which are not often
present in microbes of terrestrial origin (Fenical, 1993). Therefore,
this rich marine habitat provides a magnicent opportunity to
discover newer compounds such as antibiotics, enzymes, vitamins,
drugs, biosurfactant (BS), bioemulsier (BE) and other valuable
compounds of commercial importance (Jensen and Fenical, 1994;
Austin, 1989; Romanenko et al., 2001; Lang and Wagner, 1993).
Among these various marine bioactive compounds, BS/BE are of great
importance due to their structural and functional diversity and
industrial applications (Banat et al., 1991; Banat, 1995a,b; Rodrigues
et al., 2006). Biosurfactants and bioemulsiers are amphiphilic
compounds containing both a hydrophilic and a hydrophobic moiety
and therefore are able to display a variety of surface activities that
allow solubilisation of hydrophobic substrates (Desai and Banat,
1997). During the last decade they have been under investigation as
potential replacements for synthetic surfactants and are expected to
have many potential industrial and environmental applications
related to emulsication, foaming, detergency, wetting, dispersion
and solubilisation of hydrophobic compounds (Banat et al., 2000;
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Table 1
Major features/aspects of bioactive compounds produced by marine microorganisms in comparison to those for other habitats.
Feature/aspect
Marine
Other resources
Source of isolation
Ability to cultivate
Maintenance of environmental and
physiological requirement
Novelty in structures and functional properties
Toxicity nature of biosurfactant/bioemulsier
Possibility of culture contamination
Literature availability
Quite large
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RAG-1 was isolated from the Mediterranean Sea (Reisfeld et al., 1972)
and found to produce a surfactant (Rosenberg et al., 1979), which has
been exploited commercially as the product Emulsan. A number of
potential applications of emulsan has been patented by Gutnick and coworkers (1981) and was marketed by Petrorm Inc. for cleaning of oil
contaminated vessels, oil spills and microbial enhanced oil recovery
(MEOR) (Desai and Patel, 1994). Emulsan is a high molecular weight
heteropolysaccharide protein containing repeating trisaccharide of
N-acetyl -D-galactosamine, N-acetylgalactosamine uronic acid and an
unidentied N-acetyl amino sugar. Fatty acids (FA) are covalently linked
to the polysaccharide through o-ester linkages (Belsky et al., 1979;
Rosenberg et al., 1979; Zukerberg et al., 1979; Gorkovenko et al., 1997;
Desai and Banat, 1997; Rosenberg and Kaplan, 1987). Even at very low
concentration (b0.001 to 0.01%), emulsan can emulsify hydrocarbons
efciently and is the most powerful emulsion stabilizer having several
applications in diverse elds (Belsky et al., 1979; Gutnick and Shabtai,
1987; Zosim et al., 1982). Different Acinetobacter sp. produce protein
polysaccharide complexes that possess surface active properties
(Kaplan and Rosenberg, 1986; Shete, 2003). A great deal of studies
have been carried out regarding isolation, emulsifying, chemical, physical
properties, and specicity to hydrocarbon substrate (Rosenberg et al.,
1979). A. calcoaceticus represents an excellent candidate for the
availability of maximum number of reports in the application area.
Table 2
Summary of marine polymeric biosurfactant/bioemulsier produced by different microorganisms.
Source organism
Chemical composition
Application property
Reference
A. calcoaceticus A2
A. radioresistens
A. calcoaceticus BD4
Pseudomonas nautica
Pseudomonas uorescence
Myroids
Streptomyces
Yarrowia lipolytica NCIM 3589
Yarrowia lipolytica IMUFRJ 50682
Rhodotorula glutinis
Carbohydrateprotein complex
Halomonas
Halomonas eurihalina
Antarctobacter
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Table 3
Summary of marine glycolipid and other types of biosurfactant (BS)/bioemulsier (BE) produced by different microorganisms.
BS/BE type
Source organism
Chemical composition
Application property
Reference
Glycolipid
Alcaligenes
Non-toxic glucoselipid
Arthrobacter
Surface activity
Alcanivorax borkumensis
Rhodococcus
Halomonas
Unidentied marine
bacterium MM1
Lipopeptide
Bacillus circulans
Azotobacter chroococcum
Phospholipids
and fatty acids
Glycolipopeptide
Myroides
Corynebacterium kutscheri
a
Bacillus licheniformis BAS50 has been isolated from oil reservoir (at 1500 m) but it can grow at high salt up to 13% and optimum salt around see water level so it resembles marine
bacteria and can grow at a marine environment.
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2.2.4. Rhodococcus
Different species of Rhodococcus are known for production of
glycolipid surface active molecules. Peng et al. (2007) isolated an oildegrading bacterium R. erythropolis strain 3C-9 from seaside soil of the
Island of Xiamen, located on the west bank of the Taiwan Strait. Two
different types (structure and chemical composition) of biosurfactants
as a glycolipid and free fatty acids are produced by R. erythropolis.
2.2.5. Halomonas
These marine strains are mostly known for production of EPS with
emulsifying properties. There are few reports suggesting that the
emulsifying surface active agents are also produced by Halomonas
species. Detailed studies were carried out by Pepi and co-workers
(2005). A bacterial strain, Halomonas ANT-3b was isolated from sea
ice seawater interface at Terra Nova Bay station, Ross Sea, Antarctica
that produces emulsifying glycolipids. This surface active compound is
a mixture of fatty acids and sugars of a molecular weight of 18 kDa
with mesophase (liquid crystal) organization. We isolated another
strain H. hydrothermalis IACS3 from marine water sample of Caspian
Sea (Iran) which produce glycolipid type biosurfactant that reduces
surface tension of distilled water from 71 to 31 mN/m and emulsies
various plant oils efciently (Satpute, 2008).
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Fig. 2. Various roles played by exopolysaccharide (EPS) produced by marine microorganisms in the marine ecosystem.
subsp. jiensis one of the new bacterium produce EPS similar to that of
xanthan containing glucose, galactose, mannose, glucuronic and
galacturonic acids.
2.6.6. Pseudoalteromonas producing exopolysaccharide
Chemically diverse types EPS are produced by Antarctic isolates
which mostly contains charged uronic acid residues and some
contains sulfate groups. Polymers produced by some of the isolates
from this microbial population are large polymers of approximately
5.7 MDa in size (Mancuso Nichols et al., 2005c). EPS producing
Pseudoalteromonas sp. strain CAM025 and CAM036 were also isolated
from seawater and sea ice in southern ocean (Mancuso Nichols et al.,
2004). Marine bacterium CAM025, isolated from sea ice sample shows
30 times higher yield at 2 C and 10 C than at 20 C. Most of the
marine bacterial isolates are psychrotolerant where their optimum
growth occurs at about 20 C. However, these bacteria may produce
optimum EPS at lower temperature (Mancuso Nichols et al., 2005a).
Recently, interesting work is reported by Gutirrez et al. (2008). This
research group has described the production of polymeric emulsifying
agent from Pseudoalteromonas with the absorption property to metal
ions. The high-molecular-weight glycoprotein exopolymer PE12
contains identical monosaccharide with exopolymers of bacteria.
PE12 also contains uronic acid, protein similar to that of Pseudoalteromonas/Alteromonas species. EPS producing Pseudoalteromonas strain
HYD 721is isolated from deep-sea hydrothermal vent of East Pacic
Rise which contains repeating unit of the polymer with a branched
oligosaccharide (Rougeaux et al., 1999a). Novel types of EPS
producing bacteria have been isolated from deep-sea hydrothermal
vent (Rougeaux et al., 1996).
In the year of 1996, Raguns et al.'s group isolated two
Pseudoalteromonas species from deep-sea hydrothermal vents produce same types of polymer containing glucose, galactose, mannose,
glucuronic and galacturonic acids. Studies with transmission electron
microscopy (TEM) of immunogold-labeled indicated that the EPS
production in Marine Pseudomonas sp. strain S9 is closely associated
with the cell surface. EPS production takes place during the bacterial
growth and energy-nutritional starvation condition (Wrangstadh
et al., 1990). A previous study also reported secretion of EPS by
Pseudomonas sp. S9 during complete nutrient starvation and energy.
When the cells were starved in agitated conditions, EPS secretion
occurs and different adhesion pattern to hydrophobic surfaces is
observed (Wrangstadh et al., 1986).
2.6.7. Rhodococcus producing exopolysaccharides
Urai et al. (2007) isolated marine R. erythropolis PR4 that produces
a large quantity of acidic EPS (FR1 and FR2). FR2 composed of
D-galactose, D-glucose, d-mannose, D-glucuronic acid, and pyruvic
acid at a molar ratio of 1:1:1:1:1, and contained 2.9% (w/w) stearic
acid and 4.3% (w/w) palmitic acid attached via ester bonds. It was
proposed that EPS plays a signicant role in the hydrocarbon tolerance
of this bacterium.
2.6.8. Zoogloea producing exopolysaccharides
These are Gram-negative, straight or slightly curved, plump rods,
non-spore or cyst forming free living bacteria in organically polluted
fresh and waste water bodies. Kwon et al. (1994) has reported the
production of polysaccharide extracellularly from Zoogloea sp. This
marine bacterium produces water-soluble polysaccharide: WSP (cellfree liquid medium) and cell-bound polysaccharide: CBP (cell surface)
containing glucose, galactose and mannose as main sugars and uronic
acid. Zoogloea sp. KCCM10036 produces two types of EPS (CBP: cellbound polysaccharide and WSP: water-soluble polysaccharide).
Emulsions formed by cell-bound polysaccharide are highly stable
when compared with other commercially available EPS such as
xanthan gum, Tween series, Triton. Both types of EPS from Zoogloea
sp. possess excellent occulating activity (Lim et al., 2007). Zoogloea
sp. secretes water-soluble and the other cell-bound EPS possessing
non-Newtonian, pseudoplastic uid behaviour. Solutions of these EPS
were also reported to have low activation energies with good
rheological behaviour over broader pH range (212) and temperature
(2080 C) in the presence of NaCl (Jang et al., 2002).
2.6.9. Cyanobacteria producing exopolysaccharide
Mucilaginous envelope coating is common around marine Cyanobacteria living in hypersaline environments and therefore, most of
these cyanobacterial strains can produce EPS in the surrounding
media (Campbell and Golubic, 1985). Fifteen EPS producing unicellular cyanobacterial strains have been isolated from saline environments. Cyanothece EPS contains uronic acids, six to eight
monosaccharides, with one or two acidic sugars. Other chemical
groups like acetyl, pyruvyl, and/or sulfate groups are also detected.
EPS from such organisms are promising candidates for various
industrial applications (De Philippis et al., 1998). Other EPS from
cyanobacterial strain Cyanothece 16Som2 which was isolated from a
Somaliland saltpan contains various sugars like glucuronic acid,
galacturonic acid, galactose, glucose, mannose, xylose and fucose.
Nitrogen limitation condition favours the enhanced production of EPS
from Cyanothece sp. (De Philippis et al., 1993). EPS from Cyanothece
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sp. ATCC 51142 has high content of calcium and plays key role in
removing metals ions (Shah et al., 2000). Another report on EPS
producing Cyanothece sp. 113 isolated from salt lakes in China (Chi
et al., 2007) reported novel type of -D-1,6-homoglucan. Other
marine cyanobacterial strains like Schizothrix sp and Oscillatoria sp.
isolated from marine stromatolites, Bahamas have been reported for
production of EPS (Kawaguchi and Decho, 2000).
2.6.10. Vibrio producing exopolysaccharides
Vibrio is a Gram-negative marine bacterium and ubiquitous in
nature. Marine biofouling material were used to isolate EPS from V.
alginolytics to carry out and physicochemical studies (Jayaraman and
Seetharaman, 2003). Jodi et al. (2000) investigated the factor(s)
responsible for the opaque and translucent phenotypes, the cell
organization within both colony types in V. parahaemolyticus. They
showed that both type cultures secrete EPS containing four sugars like
glucose, galactose, fucose and N-acetylglucosamine. They also noted
that the copious amount of EPS produced by the opaque strain lls the
intercellular space within the colony, resulting in increased structural
integrity and the opaque phenotype. The structure of the EPS
produced under laboratory conditions by Vibrio diabolicus (isolated
from a deep-sea hydrothermal vent of East Pacic Rise) showed the
presence of linear tetrasaccharide repeating unit (Rougeaux et al.,
1999b). Raguns et al. (1996) reported production of EPS containing
different sugars like glucose, galactose, mannose, glucuronic and
galacturonic acids from the Vibrio species isolated from deep-sea
hydrothermal vents. An interesting patent is led by Guezennec et al.
(2002) on genus Vibrio belonging to V. diabolicus for production and
uses of water-soluble polysaccharides. Molecular weight of the EPS
from this marine Vibrio is 800,000 Da and contains osamines (30 5%)
with uronic acids (32 5%) and monosaccharide like glucuronic acid
(11.2%), N-acetylglucosamine (18%), and N-acetylgalactosamine (7.9%).
The EPS is similar to that of heparin and hence nds important applications in pharmaceutics.
2.6.11. Other exopolysaccharide producing bacteria
Other than the above mentioned (2.6.1 to 2.6.10) marine bacterial
population, there are many other marine bacteria producing EPS.
However, the number of reports available on each bacterial strain is
limited. This shows the diversity in production of bioactive molecules
by microorganisms in the marine biosphere. For example, Hyphomonas strain MHS-3 isolated from shallow water sediments in Puget
Sound found to produce adherent form of exopolysaccharide (EPS)
capsule associated with Glycine max lectin, Arachis hypogaea lectin
and Bauhinia purpurea lectin (BPA). The capsule of Hyphomonas strain
MHS-3 EPS has an effective adhesive property (Quintero and Weiner,
1995). EPS namely, Marinactan from Flavobacterium uliginosum shows
antitumor activity against sarcoma-180. Basically this bacterium
requires typical marine sea water condition for growth. The novel
type of Marinactan is composed of glucose, mannose and fucose with
the proportion of 7:2:1 (Umezawa et al., 1983). It is possible to
determine the effect of surface on the yield and composition of EPS
produced by bacteria. This fact is well explained from the work on two
marine Desulfovibrio sp. Indl SRB-sulfate-reducing bacteria (SRB)
isolated from severe corrosion failures. Bacterial isolates were grown
in static batch cultures with and without the presence of carbon steel
surfaces. EPS of these Desulfovibrio sp. contained similar kind of
carbohydrates, proteins and nucleic acids in with different quantities.
Studies also showed that surface provided for the growth had effect
on the type of EPS produced by Desulfovibrio sp. (Zinkevich et al.,
1996). Highly mucoidal Mediterranean Sea isolate, Haloferax mediterranei is an extreme halophilic archaebacterium which secretes
heteropolysaccharide extracellularly. The EPS contains glucose,
galactose, unidentied sugar, amino sugars and uronic acids. The
rheological properties of EPS like pseudoplasticity and viscosity are
seen effectively at extreme ranges of pH, temperature and salinity.
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new polymer-secreting bacterium from a deep-sea hydrothermal vent, Ah.
macleodii subsp jiensis, and preliminary characterization of the polymer. Appl
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