Journal of Applied Microbiology ISSN 1364-5072

REVIEW ARTICLE

Applications of cyanobacteria in biotechnology

R.M.M. Abed1, S. Dobretsov2 and K. Sudesh3
1 Biology Department, College of Science, Sultan Qaboos University, Al Khoud, Sultanate of Oman
2 Marine Science and Fisheries, College of Agricultural and Marine Sciences, Sultan Qaboos University, Al Khoud, Sultanate of Oman
3 School of Biological Sciences, Universiti Sains Malaysia, Penang, Malaysia

Keywords
bioactive compounds, biofertilizers,
bioplastics, biotechnology, cyanobacteria.
Correspondence
Raeid M. M. Abed, Biology Department,
College of Science, Sultan Qaboos University,
PO Box 36, AL Khoud 123, Muscat, Sultanate
of Oman. E-mail: rabed@mpi-bremen.de
2008 ⁄ 0456: received 15 March 2008, revised
1 June 2008 and accepted 3 June 2008
doi:10.1111/j.1365-2672.2008.03918.x
Summary
Cyanobacteria have gained a lot of attention in recent years because of their
potential applications in biotechnology. We present an overview of the litera-
ture describing the uses of cyanobacteria in industry and services sectors and
provide an outlook on the challenges and future prospects of the field of
cyanobacterial biotechnology. Cyanobacteria have been identified as a rich
source of biologically active compounds with antiviral, antibacterial, antifungal
and anticancer activities. Several strains of cyanobacteria were found to accu-
mulate polyhydroxyalkanoates, which can be used as a substitute for nonbiode-
gradable petrochemical-based plastics. Recent studies showed that oil-polluted
sites are rich in cyanobacterial consortia capable of degrading oil components.
Cyanobacteria within these consortia facilitated the degradation processes by
providing the associated oil-degrading bacteria with the necessary oxygen,
organics and fixed nitrogen. Cyanobacterial hydrogen has been considered as a
very promising source of alternative energy, and has now been made commer-
cially available. In addition to these applications, cyanobacteria are also used in
aquaculture, wastewater treatment, food, fertilizers, production of secondary
metabolites including exopolysaccharides, vitamins, toxins, enzymes and phar-
maceuticals. Future research should focus on isolating new cyanobacterial
strains producing high value products and genetically modifying existing strains
to ensure maximum production of the desired products. Metagenomic libraries
should be constructed to discover new functional genes that are involved in the
biosynthesis of biotechnological relevant compounds. Large-scale industrial
production of the cyanobacterial products requires optimization of incubation
conditions and fermenter designs in order to increase productivity.

to another (Stal 1995). All cyanobacteria carry out oxy-

Introduction
Cyanobacteria are prokaryotic oxygenic phototrophs
found in almost every conceivable habitat on earth (Ferris
et al. 1996; Ward et al. 1997; Nübel et al. 1999, 2000;
Abed and Garcia-Pichel 2001; Garcia-Pichel and Pringault
2001). They exist in different morphologies including uni-
cellular and filamentous forms (Castenholz 2001). While
unicellular types exist as single cells, suspended or ben-
thic, or aggregates, filamentous types may be thin or
thick, single trichome or bundles either with or without a
sheath. Cyanobacteria are able to perform different modes
of metabolism with the capacity to switch from one mode
genic photosynthesis but some cyanobacterial species can
switch to the typical bacterial anoxygenic photosynthesis
using sulfide as electron donor (Cohen et al. 1986).
Under anoxic conditions and during the dark, cyanobac-
teria carry out fermentation (Stal 1997). Some cyanobac-
teria form heterocysts and have the ability to fix
atmospheric nitrogen (Capone et al. 2005). Phylogenetic
analysis of cyanobacteria based on 16S rRNA genes
showed that they are a diverse, monophyletic phylum of
organisms within the bacterial radiation. Research on
cyanobacteria in the last decades focused largely on their
ecology, morphology, physiology and 16S rRNA-based

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Journal compilation ª 2008 The Society for Applied Microbiology, Journal of Applied Microbiology 106 (2009) 1–12 1
Cyanobacterial biotechnology
phylogeny but relatively little has been done on their
potential uses in biotechnology.
The overwhelming available knowledge on the diversity
and physiology of cyanobacteria serves as an excellent
base for exploring their applications in biotechnology. In
the last few years, cyanobacteria have gained much atten-
tion as a rich source of bioactive compounds and have
been considered as one of the most promising groups of
organisms to produce them (Bhadury et al. 2004; Dahms
et al. 2006). These cyanobacterial metabolites include
antibacterial (Jaki et al. 2000), antifungal (Kajiyama et al.
1998), antiviral (Patterson et al. 1994), anticancer (Ger-
wick et al. 1994), antiplasmodial (Papendorf et al. 1998),
algicide (Papke et al. 1997) and immunosuppressive
agents (Koehn et al. 1992). Screening of cyanobacteria for
antibiotics has opened a new horizon for discovering new
drugs. Some cyanobacteria have also been found to intra-
cellularly accumulate polyhydroxyalkanoates (PHA),
which are comparable in properties to polyethylene and
polypropylene (Steinbüchel et al. 1997). These biodegra-
dable plastics could replace oil-derived thermoplastics in
some fields. Recent research on cyanobacteria has demon-
strated that they form ideal consortia with chemotrophic
bacteria and can effectively be used to cleanup oil-
contaminated sediments and wastewaters (Abed and
Köster 2005). Cyanobacteria have many more biotechno-
logical applications that await possible uses in maricul-
ture, food, fuel, fertilizers, colorants and production of
various secondary metabolites including toxins, vitamins,
enzymes and pharmaceuticals. Considering the diverse
potential uses of cyanobacteria in biotechnology, an over-
view of these uses is presented here.
Cyanobacterial bioactive compounds
Cyanobacteria have been identified as a new and rich
source of bioactive compounds (Abarzua et al. 1999;
Shimizu 2003; Bhadury et al. 2004; Dahms et al. 2006).
Isolated compounds belong to groups of polyketides,
amides, alkaloids, fatty acids, indoles and lipopeptides
(Abarzua et al. 1999; Burja et al. 2001; Table 1). The liter-
ature review showed that to date up to 19 cyanobacterial
strains produce more than 20 different bioactive com-
pounds. Most of the bioactive compounds isolated from
cyanobacteria tend to be lipopeptides, i.e. they consist of
an amino acid fragment linked to a fatty acid portion.
The range of biological activity of secondary metabolites
isolated from cyanobacteria includes antibacterial, anti-
fungal, antialgal, antiprotozoan, and antiviral activities
(Table 1). Only few cyanobacteria produce bioactive com-
pounds that show a broad spectrum of biological activi-
ties. For example, the cyanobacterium Phormidium sp.
has been reported to inhibit growth of different Gram-
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R.M.M. Abed et al.
positive and Gram-negative bacterial strains, yeasts, and
fungi (Bloor and England 1989). Another example is
Lyngbya majuscula (Burja et al. 2001) that produces
numerous chemicals including nitrogen-containing com-
pounds, polyketides, lipopeptides, cyclic peptides and
many others (Shimizu 2003). The biological activity of
these compounds is also diverse and includes protein
kinase C activators and tumour promoters, inhibitors of
microtubulin assembly, antimicrobial and antifungal com-
pounds and sodium-channel blockers.
Secondary metabolites with antibacterial activity are
widely produced by cyanobacteria (Dahms et al. 2006;
Table 1). These compounds are effective against Gram-
positive and ⁄ or Gram-negative bacteria. Both toxic and
nontoxic strains of cyanobacteria are producers of anti-
bacterial compounds that are distinct from cyanotoxins
(Østensvik et al. 1998). Antifungal compounds include
fisherellin A, hapalindole, carazostatin, phytoalexin, toly-
toxin, scytophycin, toyocamycin, tjipanazole, nostocycla-
mide and nostodione produced by cyanobacteria
belonging to Stigonematales, Nostocales and Oscillatoriales
(Table 1). Additionally, cyanobacteria produce a broad
spectrum of antialgal compounds, which may be used to
control algal blooms. Cyanobacteria probably use these
compounds in order to out-compete other micro-organ-
isms. Antialgal compounds produced by cyanobacteria
inhibit growth of algae, their photosynthesis, respiration,
carbon uptake, enzymatic activity and induce oxidative
stress (Dahms et al. 2006). In contrast to the large
amount of antibacterial and antialgal compounds isolated
from cyanobacteria there are only a few compounds that
show antiviral properties (Table 1), although 2–10% of
extracts of different cyanobacterial species have been
shown to have antiviral activity (Cohen 1999). These
include acetylated sulfoglyco-lipids from Oscillatoria raoi
(Reshef et al. 1997) and spirulan from Spirulina platensis
(Hayashi et al. 1991). The compounds isolated from
Lyngbya lagerhaimanii and Phormidium tenue has been
shown to have anti-HIV activity (Rajeev and Xu 2004).
Gamma linolenic acid (GLA) found rich in S. platensis
and Arthrospira sp. is medically important since it is
converted in the human body into arachidionic acid and
then into prostaglandin E2. This compound has a lower-
ing action on blood pressure and plays an important role
in lipid metabolism. Some of the marine cyanobacteria
constitute potential sources for large-scale production of
vitamins, such as vitamins B and E (Plavsic et al. 2004).
Piechula et al. (2001) demonstrated that some cyanobac-
teria can produce thermostable enzymes. Out of 21 endo-
nucleases from Phormidium spp. 4 enzymes that catalyse
the hydrolysis of DNA and RNA were active in a wide
range of temperatures from 15 to 60C. Prenyltransferases
enzymes catalysing the consecutive condensation of
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R.M.M. Abed et al. Cyanobacterial biotechnology

Table 1 Bioactive compounds from cyanobacteria

Species of cyanobacteria Bioactive compounds Biological activity References

Family oscillatoriaceae
Lyngbya majuscula Malyngolide Antibacterial Burja et al. (2001)
Lyngbyatoxins PKC activator (Shimizu 2003)
Debromoaplysiatoxin Inflammatory
Curacin A Microtubulin assembly inhibitors
Kalkitoxin Sodium channel blocker
Cyclic polypeptide Anti-HIV activity Rajeev and Xu (2004)
L. lagerheimii Sulpholipid Anti-HIV activity Rajeev and Xu (2004)
Oscillatoria raoi Acetylated sulfoglyco-lipids Antiviral Reshef et al. (1997)
Phormidium tenue Galactosyldiacylglycerols Antialgal Murakami et al. (1991)
Anti-HIV Rajeev and Xu (2004)
Phormidium spp. Thermostable enzymes Catalysis of reactions Piechula et al. (2001)
Spirulina platensis Spirulan Antiviral Hayashi et al. (1991)
Gamma linolenic acid Predecessor of arachidonic acid Cohen (1999)
Vitamin B and E Antioxidants and co-enzymes Plavsic et al. (2004)
Family hyellaceae
Hyella caespitose Carazostatin Antifungal Burja et al. (2001)
Family nostocaceae
Nostoc spongiaeforme Nostocine A Antialgal Hirata et al. (1996)
N. commune Nostodione Antifungal Bhadury and Wright (2004)
Nostoc sp. Nostocyclamide Antifungal Moore et al. (1988)
N. linckia Cyanobacterin LU-1 Antialgal Gromov et al. (1991)
N. insulare Norharmane Antibacterial Volk and Furkert (2006)
N.sphaericum Indolocarbazoles Antiviral Cohen (1999)
Anabaena circinalis Anatoxin-a Inflammatory Rajeev and Xu (2004)
Family Scytonemaceae
Scytonema hofmanni Cyanobactericin Antialgal Abarzua et al. (1999)
S. ocellatum Tolytoxin Antifungal Patterson and Carmeli (1992);
Phytoalexin Patterson and Bolis (1997)
S. pseudohofmanni Scytophycins Antifungal Burja et al. (2001)
Family microchaetaceae
Tolypothrix tenuis Toyocamycin Antifungal Banker and Carmeli (1998)
T. tjipanasensis Tjipanazoles Antifungal Bonjouklian et al. (1991)
Family stigonemataceae
Fischerella muscicola Fisherellin Antialgal, antifungal Dahms et al. (2006)
Hapalosiphon fontinalis Hapalindole Antifungal Burja et al. (2001)
Family merismopediaceae
Gomphosphaeria aponina Aponin Antialgal Bhadury and Wright (2004)
Family chroococcaceae
Microcystis aeruginosa Kawaguchipeptin B Antibacterial Dahms et al. (2006)
Synechocystis sp. Naienones A-C Antitumoural Nagle and Gerwick (1995)
Synechococcus elongates Thermostable enzyme Catalysis of reactions Ohto et al. (1999)
Thermosynechococcus elongatus BP-1 Thermostable polyphosphate kinase Production of dipeptides Sato et al. (2007)

homoallylic diphosphate of isopentenyl diphosphates at
temperatures above 60 C have been isolated from the
thermophilic cyanobacterium Synechococcus elongates
(Ohto et al. 1999). Thermostable polyphosphate kinase
from the thermophilic cyanobacterium Thermosynecho-
coccus elongatus BP-1 was successfully employed in an ATP
regeneration system that could be used at high tempera-
tures for the effective production of d-amino acid dipep-
tides (Sato et al. 2007). These enzymes and other heat
stable bioactive compounds are of great interest in
biotechnology. So far none of the isolated compounds have
demonstrated antilarval activity and inhibition of settle-
ment of larvae and algal spores (Dobretsov et al. 2006).
Cyanobacteria have been used to synthesize isotopically
labelled compounds such as sugars, lipids and amino
acids, which are nowadays commercially available (Patter-
son 1996). This is achieved by growing the cyanobacetria
in photobioreactors and allowing them to photosyntheti-
cally transform simple labelled compounds such as 14CO2,
13
CO2, 33H2O and 15NO3 into complex organics.

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Journal compilation ª 2008 The Society for Applied Microbiology, Journal of Applied Microbiology 106 (2009) 1–12 3
Cyanobacterial biotechnology
Cyanobacterial bioplastics
(polyhydroxyalkanoates, PHAs)
In conditions of excess essential nutrients, many micro-
organisms usually assimilate and store nutrients for future
consumption. Various storage materials have been identi-
fied in micro-organisms, which include glycogen, sulfur,
polyamino acids, polyphosphate, and lipid. PHAs are
lipoidic material accumulated by a wide variety of micro-
organisms in the presence of abundant carbon sources
(Anderson and Dawes 1990). The assimilated carbon
sources are biochemically processed into hydroxyalkano-
ate monomer units, polymerized, and then stored in the
form of water insoluble inclusions (or granules) in the
cell cytoplasm (Fig. 1a). PHA is a crystalline thermoplas-
tic with properties comparable to that of polypropylene
(Doi 1990). PHA has been the focus of attention for the
past three decades as a potential substitute for nonbio-
degradable petrochemical-based plastics. This is because
PHA is an ideal biodegradable material that can be com-
pletely mineralized into water and carbon dioxide by the
action of naturally occurring micro-organisms. In addi-
tion, PHA is also a biocompatible material and is being
studied for its application in the biomedical and biophar-
maceutical fields (Williams et al. 1999; Sudesh 2004). The
idea of producing PHA from CO2 is thought to contrib-
ute to a carbon neutral process for making plastics. The
idea is also commercially attractive because then the pro-
duction of PHA will be based on a readily available and
free carbon and energy source (sunlight).
(a) (b)
PHA granules
500 nm
(c) PHA (d)
granules
10 µm
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R.M.M. Abed et al.
Some cyanobacteria, like S. platensis, can accumulate
PHA under phototrophic and ⁄ or mixotrophic growth
conditions with acetate (Fig. 1c,d). The PHA granules can
be stained by Nile blue A (Ostle and Holt 1982) (Fig. 1a)
and they appear as discrete electron transparent granules
in the cell cytoplasm (Fig. 1b). Cyanobacteria are of
particular interest as PHA producers because of their
minimal nutrient requirements for growth and capability
of accumulating PHA by oxygenic photosynthesis. Like
higher plants, cyanobacteria fix CO2 from the atmosphere
and turn it into PHA under nitrogen limiting conditions.
Most of the known cyanobacteria that are capable of syn-
thesizing PHA usually accumulate PHAs amounting to
less than 6 wt% of their cell dry weight (CDW) (Vincen-
zini et al. 1990; Stal 1992; Arino et al. 1995; Carr 1996).
Spirulina platensis and Synechocystis sp. PCC 6803 have
been reported to accumulate a maximum of 6 wt%
(Campbell et al. 1982) and 7 wt% (Sudesh et al. 2002)
of CDW of PHA under mixotrophic conditions. Poly(3-
hydroxybutyrate) [P(3HB)] is the most common type of
PHA synthesized by most bacteria. P(3HB) is also the
most common type of PHA synthesized by cyanobacteria.
Vincenzini et al. (1990) found that when Spirulina was
cultivated under photoautotrophic conditions, the
P(3HB) content was low (0Æ3 wt% of CDW), while under
mixotrophic growth conditions in the presence of acetate,
P(3HB) level amounted to about 3 wt% of CDW. It is
known that the synthesis of P(3HB) is primarily regulated
by 3-ketothiolase, which is inhibited by high concentra-
tions of free coenzyme A (CoA). High concentrations of
Figure 1 Polyhydroxyalkanoates (PHAs) in
1 cm micro-organisms. (a) Transmission electron
micrograph (TEM) of PHA granules in a
cyanobacterial cell. The granules can be read-
ily dissolved in chloroform and precipitated in
methanol. The resulting extract resembles tra-
ditional thermoplastics such as polypropylene.
(b) PHA film cast from solvent. (c) PHA
granules accumulated by Spirulina platensis
stained with Nile blue A. The PHA granules
PHA
appear as bright orange particles in the cell
cytoplasm when viewed under fluorescence
microscope. (d) Transmission electron micro-
5 µm graph showing the discrete PHA granules
accumulated in the cytoplasm of S. platensis.
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R.M.M. Abed et al. Cyanobacterial biotechnology

intracellular acetyl-CoA favours the synthesis and accu-
mulation of P(3HB). Thus, when acetate is present in the
growth medium, the intracellular acetyl-CoA concentra-
tion increases at the expense of free CoA pool, resulting
in enhancement of P(3HB) biosynthesis. The role of
P(3HB) in cyanobacteria is to provide cells with a mecha-
nism of removal of excess reducing equivalents resulting
from a disruption of the balanced formation of ATP and
NADPH from photosynthesis (De Philippis et al. 1992).
Table 2 shows a summary of known PHA-producing
cyanobacteria, the various carbon sources tested and the
maximum amount of PHAs accumulation obtained to
date.
Cyanobacterial consortia for bioremediation
purposes
Many studies have reported the ability of cyanobacteria
to oxidize oil components and other complex organic
compounds such as surfactants and herbicides (Yan et al.
1998; Radwan and Al-Hasan 2000; Raghukumar et al.
2001; Mansy and El-Bestway 2002). Among these cyano-
bacteria were the nonaxenic cultures of Microcoleus chtho-
noplastes and Phormidium corium, degrading n-alkanes
(Al-Hasan et al. 1998), Oscillatoria sp. and Agmenellum
quadruplicatum oxidizing naphthalene to 1-naphthol
(Cerniglia et al. 1979, 1980a), Oscillatoria sp. strain JCM
that oxidized biphenyl to 4-hydroxybiphenyl (Cerniglia
et al. 1980b) and Agmenellum quadruplicatum that
metabolized phenanthrene into trans-9,10-dihydroxy-
9,10-dihydrophenanthrene and 1-methoxy-phenenthrene
(Narro 1985). However, recent studies have demonstrated
that it is indeed not the cyanobacteria that are responsible
for the degradation of these compounds but the associ-
ated aerobic organotrophic bacteria (Radwan et al. 2002;
Abed and Köster 2005; Sánchez et al. 2005). In spite of
that, the presence of cyanobacteria alongside with the
aerobic organotrophs facilitated the degradation process
and both groups constituted ideal consortia for degrada-
tion of petroleum and other complex organic compounds
(Abed and Köster 2005). While aerobic bacteria directly
degrade these components, cyanobacteria play an equally
important indirect role in biodegradation by (i) providing
them with oxygen (byproduct of photosynthesis), which
is needed for the breakdown of aromatic rings, (ii) provi-
ding them with simple organics (produced by photosyn-
thesis and fermentation) and (iii) providing them with
fixed nitrogen (through nitrogen fixing strains), which is
often limited in different environments. Abed and Köster
(2005) demonstrated the capability of an Oscillatoria-
Gammaproteobacteria consortium to degrade phenan-
threne, dibenzothiophene, pristine and n-octadecane. The
degradation rate of these compounds was enhanced in
the presence of the cyanobacterium. Similarly, Microcoleus
chthonplastes was found to form consortia with organo-
trophic bacteria, some of which were able to fix atmo-
spheric nitrogen while others could degrade aliphatic
heterocyclic organo-sulfur compounds as well as alkyl-
ated monocyclic and polycyclic aromatic hydrocarbons
(Sánchez et al. 2005). Bacteria immobilized in the sheaths
coating macroalgae were shown to degrade petroleum
compounds (Radwan et al. 2002). These indigenous
consortia could be ideally used in the bioremediation of
polluted sites without the need to add new bacteria or
fertilizers to the field. Al-Awadhi et al. (2003) grew these
consortia on gravel particles and used them successfully
to clean up oil pollution.
Cyanobacteria and their associated bacteria have also
been successfully used in wastewater treatment. For exam-
ple, cultures of Oscillatoria sp. BDU 30501, Aphanocapsa

Table 2 PHA-producing cyanobacteria and the types and contents of their PHA compounds

Species of cyanobacteria Types of PHA Carbon sources tested PHA content (w ⁄ w) References

Chlorogloea fritschii P(3HB) CO2 ND Jensen and Sicko (1971)

C. fritschii P(3HB) Acetate ND Jensen and Sicko (1971)
Spirulina platensis P(3HB) CO2 6% Campbell et al. (1982)
S. maxima P(3HB) CO2 0Æ70% De Philippis et al. (1992)
S. maxima P(3HB) Acetate 2% De Philippis et al. (1992)
Gloeothece sp. P(3HB) CO2 ⁄ acetate 6% Stal (1992)
Oscillatoria limosa P(3HV) CO2 ⁄ acetate 6% Stal (1992)
Trichodesmium thiebautii P(3HB) Natural conditions 0Æ20% Siddiqui et al. (1992)
Gloeothece sp. PCC6909 P(3HB) CO2 ND Arino et al. (1995)
Synechococcus MA19 P(3HB) CO2 7Æ50% Miyake et al. (1997)
Synechocystis sp. PCC6803 P(3HB) Acetate 10% Hein et al. (1998)
S. platensis P(3HB) CO2 ⁄ acetate 10% Jau et al. (2005)
Synechocystis sp. PCC6803 P(3HB) Fructose ⁄ acetate 38% Panda and Mallick (2007)
Nostoc muscorum P(3HB) Acetate ⁄ glucose 45Æ60% Sharma et al. (2007)

ND, not defined.

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Cyanobacterial biotechnology R.M.M. Abed et al.

sp. BDU 16 and a halophilic bacterium Halobacterium US
101 were used to treat a factory effluent and resulted in
reduction of calcium and chloride to levels that did not
inhibit survival and multiplication of fish (Uma and
Subramanian 1990). Phormidium valderianum BDU 30501
was used to reduce phenol concentrations (Shashirekha
et al. 1997) while Oscillatoria boryana BDU 92181 was
used to eliminate melanoidin pigment from distillery
effluents (Kalavathi et al. 2001).
Cyanobacterial alternative energy sources
Cyanobacteria have been used to produce hydrogen gas
that constitutes an alternative future energy source to the
limited fossil fuel resources (Dutta et al. 2005). The
advantages of using biological hydrogen as a fuel are its
eco-friendly nature, efficiency, renewability and the
absence of carbon dioxide emission during its production
and utilization (Lindbald 1999). Cyanobacteria produce
hydrogen either as a byproduct of nitrogen fixation, when
nitrogenase-containing heterocystous cyanobacteria are
grown under nitrogen limiting conditions, or by the
reversible activity of hydrogenases enzymes. Heterocystous
cyanobacteria are thus more efficient in hydrogen produc-
tion than nonheterocystous types (Pinzon-Gamez et al.
2005). More than 14 cyanobacterial genera including
Anabaena, Calothrix, Oscillatoria, Cyanothece, Nostoc,
Synechococcus, Microcystis, Gloeobacter, Aphanocapsa,
Chroococcidiopsis and Microcoleus are known for their
ability to produce hydrogen gas under various culture
conditions (Lambert and Smith 1977; Sveshnikov et al.
1997; Masukawa et al. 2001; Table 3). Anabaena spp. are
able to produce significant amounts of hydrogen. Nitro-
gen-starved Anabaena cylindrica cells produce the highest
amount of hydrogen (30 ml of H2 per lit culture per
hour) (Jefferies et al. 1978). Gloeocapsa alpicola showed
increase in hydrogen production under sulfur starvation
(Antal and Lindblad 2005) whereas S. platensis could pro-
duce hydrogen under complete dark and anoxia (Aoyama
et al. 1997).
Large-scale hydrogen production by Spirulina and
Anabaena spp. has been tried using different types of bio-
reactors that included vertical column reactor, tubular
type and flat panel photobioreactor (Dutta et al. 2005).
These reactors were designed to make use of solar light for
illumination, to maximize the area for incident light (high
surface to volume ratio) and to allow sterilization and
hydrogen collection with convenience and ease. Neverthe-
less, the reactors are subjected to continuous modification
in order to increase their productivity and to decrease
costs of maintenance and production. So far, these modifi-
cations succeeded in bringing down the cost of biologically
produced hydrogen to $25 per m3 compared to $170 per
m3 for the hydrogen produced by splitting of water (Block
and Melody 1992). Increased production of hydrogen was
also achieved by genetically modifying the nitrogenase
enzyme in hydrogen-producing strains. The ongoing
research on hydrogen production by cyanobacteria focuses
on finding new strains with higher potential to produce
hydrogen, optimizing the mass production of hydrogen in
bioreactors and modifying the physiology and the genetic
system of H2-producing cyanobacteria to ensure maxi-
mum production of hydrogen.
Cyanobacteria as biofertilizers
Heterocystous cyanobacteria and several nonheterocys-
tous cyanobacteria are known for their ability to fix

Table 3 Cyanobacterial species that are capable of producing hydrogen and the growth conditions at which maximum production occur

Species of cyanobacteria Growth conditions Maximum hydrogen production Reference

Anabaena sp. PCC 7120
Anabaena cylindrica IAMM-I
Nostoc commune IAMM-I 3
Anabaena variabilis AVM13
Anabaena variabilis PK84
Anabaena variabilis ATCC 29413
Synechococcus PCC6803
Synechococcus PCC6301
Microcystis PCC 7820
Gloeobacter PCC 7421
Synechocystis PCC 6308
Synechocystis PCC 6714
Aphanocapsa montana
Gloeocapsa alpicola CALU 743
Chroococcidiopsis thermalis
Air, 20 lE m)2 s)1 2Æ6 lmol mg)1 chl a h)1 Masukawa et al. (2001)
Air, 20 lE m)2 s)1 2Æ1 lmol mg)1 chl a h)1 Masukawa et al. (2001)
Air, 20 lE m)2 s)1 0Æ25 lmol mg)1 chl a h)1 Masukawa et al. (2001)
Air and 1% CO2, 100 lE m)2 s)1 68 lmol mg)1 chl a h)1 Happe et al. (2000)
Air and 2% CO2, 113 lE m)2 s)1 32Æ3 lmol mg)1 chl a h)1 Tsygankov et al. (1998)
73%Ar, 25%N2, 2% CO2, 90 lE m)2 s)1 46Æ16 lmol mg)1 chl a h)1 Sveshnikov et al. (1997)
Air, photon fluence rate 20 lE m)2 s)1 0Æ26 lmol mg)1 chl a h)1 Moezelaar et al. (1996)
Air, photon fluence rate 20 lE m)2 s)1 0Æ09 lmol mg)1 chl a h)1 Howarth and Codd (1985)
Air, photon fluence rate 20 lE m)2 s)1 0Æ16 lmol mg)1 chl a h)1 Moezelaar et al. (1996)
Air, photon fluence rate 20 lE m)2 s)1 1Æ38 lmol mg)1 chl a h)1 Moezelaar et al. (1996)
Air, photon fluence rate 20 lE m)2 s)1 0Æ13 lmol mg)1 chl a h)1 Howarth and Codd (1985)
Air, photon fluence rate 20 lE m)2 s)1 0Æ07 lmol mg)1 chl a h)1 Howarth and Codd (1985)
Air, photon fluence rate 20 lE m)2 s)1 0Æ40 lmol mg)1 chl a h)1 Howarth and Codd (1985)
Sulfur free 4% CO2; 25 lmol 0Æ58 lmol mg)1 protein Antal and Lindblad (2005)
photons m)2 s)1
Ar and 1% CO2 0Æ7 lmol mg)1 chl a h)1 Serebryakova et al. (2000)

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R.M.M. Abed et al.
atmospheric nitrogen (Capone et al. 2005). The fertility
of many tropical rice field soils has been mainly attrib-
uted to the activity of nitrogen-fixing cyanobacteria. An
estimation showed that more than 18 kg N ha)1 year)1
was added to the soils by cyanobacteria (Watanabe and
Cholitkul 1979). Inoculation of cyanobacteria to increase
the fertility of soils has been successfully attempted. For
example, Azolla was used as an organic fertilizer in rice
cultivation in many countries (Kaushik and Venkata-
raman 1979). Addition of Azolla was found to support
the growth of soil micro-organisms including heterotro-
phic N2 fixers. Recently, nitrogen-fixing cyanobacteria
have been reported to dominate desert crusts worldwide
(Garcia-Pichel and Pringault 2001). This is believed to
contribute significantly to the fertility of desert soils and
may eventually facilitate vegetation of deserts.
Cyanobacteria as a healthy food source
Strains of Spirulina, Anabaena and Nostoc are consumed
as human food in many countries including Chile,
Mexico, Peru and Philippines. Arthrospira platensis (mis-
identified as S. platensis) is grown in large scale using
either outdoor ponds or sophisticated bioreactors but
marketed in the form of powder, flakes, tablets and cap-
sules. It is used as a food supplement because of its rich-
ness in nutrients and digestibility. It contains more than
60% proteins and is rich in beta-carotene, thiamine and
riboflavin and is considered to be one of the richest
sources of vitamin B12 (see cyanobacterial active com-
pounds). Nostoc commune is rich in fibres and proteins
and can play an important physiological and nutritional
role in the human diet. Aphanizomenon sp. is collected
from natural blooms in the Lake Klamath (Oregon, USA)
to be used as healthy food (Carmichael and Gorham
1980). Marine nitrogen-fixing cyanobacteria have also
been tested to feed fishes in aquacultures. The Tilapia fish
showed high growth rates when fed with marine cyano-
bacteria in indoor and outdoor cultures (Mitsui et al.
1983). Phormidium valderianum has been used in India to
serve as a complete aquaculture feed source based on its
nutritional value and nontoxic nature. In view of the cy-
anobacterial significance as a food source, very little
research has been performed and published about this.
Cyanobacterial emulsifiers
Halophilic cyanobacteria produce large amounts of
exopolysaccharides (EPS), which can be relevant to oil
recovery by decreasing its surface tension and increasing
its solubility and mobility. EPS, when gelated under alka-
line conditions, was employed to remove dyes from
textile effluent. The halophilic cyanobacterium Aphano-
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Journal compilation ª 2008 The Society for Applied Microbiology, Journal of Applied Microbiology 106 (2009) 1–12
Cyanobacterial biotechnology
capsa halophytica was used for the production of EPS and
its yield was increased to tenfold by immobilizing the
cells on light-diffusing optical fibres (Matsunaga et al.
1996). The cyanobacterium Cyanothece sp. ATCC 51142
had a maximum EPS production at 4Æ5% (w ⁄ v) NaCl and
pH 7 (Shah et al. 1999). Other EPS-producing cyanobac-
teria are the halotolerant Anabaena sp. ATCC 33047 (Mo-
reno et al. 1998) and Synechococcus sp. (Matsunaga et al.
1991). It should be pointed out that it is necessary in
future research to investigate the potential of additional
cyanobacteria in EPS production.
Conclusion and future directions
Our review demonstrates the versatility of cyanobacteria
to biotechnological applications. They are potent sources
of bioactive compounds, biofertilizers, bioplastics, energy,
food and have currently been used in drugs discovery,
medical diagnostics, and bioremediation. However, further
research needs to focus on the satisfactory axenic culturing
of these micro-organisms in order to facilitate their
exploitation. Additionally, new methods need to be devel-
oped to allow the cultivation of previously ‘uncultivable’
strains. The methods should consider the organism’s
requirements in the field and these conditions should be
mimicked in the laboratory. The cultivation efforts should
be directed towards unique environments, particularly
those with extreme conditions of salinity, temperature,
pH, UV and light intensity. These environments are likely
to contain novel strains with strong potential in biotech-
nology. To circumvent cultivability problem, metagenom-
ics could serve as an alternative approach (Handelsman
2004). This approach involves construction of metage-
nomic clone libraries from nucleic acids extracted directly
from environmental samples. The clone libraries are then
screened for the presence of functional genes that are
involved in the biosynthesis of certain biotechnologically
significant compounds (Zhang et al. 2005). These genes
are transferred to cultivable hosts, which can be directly
used for the production of the desired compounds.
Numerous compounds that have antibacterial and
antialgal activity have been isolated from different cyano-
bacteria, however, only limited compounds have been
screened for their antifouling, antiviral and antitumour
activities (Table 1). Recently, several potent antifouling
(Fusetani 2004), antiviral and antitumour compounds
(Proksch et al. 2002) have been isolated from macroalgae,
sponges and tunicates. These compounds are often pres-
ent in small quantities and in order to obtain sufficient
amounts it is needed to harvest and extract many marine
organisms (Dobretsov et al. 2006). In many cases, it is
the associated micro-organisms and not the host who are
the true sources of bioactive compounds. For example,
7
Cyanobacterial biotechnology
the major compounds of the sponge Dysidea herbacea are
actually produced by its cyanobacterial symbiont Oscilla-
toria spongeliae (Unson and Faulkner 1993). Unlike
eukaryotes, cyanobacteria can produce compounds much
more rapidly and in large amounts (Dahms et al. 2006).
Furthermore, they can be genetically and chemically easily
modified in order to increase compounds yield and bio-
activity. Therefore, screening of cyanobacteria for novel
bioactive compounds should be an important future
direction.
The efficient production of PHA (bioplastics) using
cyanobacteria is technologically challenging. Nevertheless,
it remains as an attractive approach considering the fact
that the carbon source comes directly from atmospheric
CO2. In contrast, the more efficient production of PHA
by bacteria relies on the use of valuable carbon sources
such as sugars from starch and fatty acids from vegetable
oils. Because of the growing pressure to reduce CO2 emis-
sion, the demands for plant products such as starch and
vegetable oils are on the rise for use as starting materials
for the production of biofuels and biobased materials.
Therefore, the demands for plant products can be
expected to increase, which inevitably will require more
fertile land to be used for agricultural activities. In such a
scenario, using cyanobacteria to produce PHA may
become more promising because the large-scale cultiva-
tion of cyanobacteria does not require fertile land.
While the biotechnological potential of cyanobacteria is
attracting an increasing attention, most of the commercial
compounds were isolated from freshwater cyanobacteria.
Marine environments with different environmental condi-
tions ranging from shallow euphotic zone to deep-sea
hydrothermal vents are likely to be a good source for a
variety of cyanobacterial species that may have high bio-
technological significance.
Although production of hydrogen by cyanobacteria
provides a promising environmentally-friendly source of
energy, it still requires optimization in order to lower the
production costs and to increase the yield. The design of
the bioreactors is essential for large-scale production and
must be transparent to allow adequate entry of light. The
hydrogen productivity tends to decrease at higher light
intensity because photosynthesis diverts the hydrogen
production pathway, hence the light regime must be con-
trolled carefully. Liquid circulation time and aeration has
an influence on hydrogen productivity since the hydrogen
producing enzymes are oxygen sensitive. Therefore, aera-
tion, although needed, must be adjusted to avoid any
inhibition of the hydrogen production pathway.
More research on the ecological applications of cyano-
bacteria needs to be performed. Recently they have been
shown to play a significant role in the stabilization of
coastal sediments by reducing hydrodynamic erosional
8 Journal compilation ª 2008 The Society for Applied Microbiology, Journal of Applied Microbiology 106 (2009) 1–12
R.M.M. Abed et al.
forces (Noffke et al. 2003). During low hydrodynamic
disturbance, cyanobacteria enhance deposition of sedi-
ments by baffling, trapping, and binding while during
periods of intensive hydraulic reworking, they shelter
their substrata against erosion or they permit flexible
deformation of sandy sediments. The exact role of cyano-
bacteria in fighting erosion has to be investigated, consid-
ering different types of sediments and different
environmental settings.
Cyanobacteria have been studied for a long time for
their interesting morphology, diversity and physiology but
pioneering work in the last decades has raised the level of
these microbes to be viewed with favour in biotechno-
logy-relevant fields. Therefore, it is essential not only to
understand and describe the diversity of cyanobacteria in
yet unexplored habitats but also to gainfully exploit them
for various industrial applications. This requires com-
bined efforts of taxonomists, molecular biologists, bio-
chemists, engineers and industry-related scientists as well
as politicians and policy makers.
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