Biotechnology Advances 47 (2021) 107684

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Biotechnology Advances
journal homepage: www.elsevier.com/locate/biotechadv

Research review paper

Prospects and development of algal-bacterial biotechnology in

environmental management and protection
Jasmine Jill Jia Yi Yong a, 1, Kit Wayne Chew b, 1, Kuan Shiong Khoo a, Pau Loke Show a, **,
Jo-Shu Chang c, d, e, *
a
Department of Chemical and Environmental Engineering, Faculty of Science and Engineering, University of Nottingham Malaysia, Jalan Broga, 43500 Semenyih,
Selangor Darul Ehsan, Malaysia
b
School of Energy and Chemical Engineering, Xiamen University Malaysia, Jalan Sunsuria, Bandar Sunsuria, 43900 Sepang, Selangor, Malaysia
c
Department of Chemical and Materials Engineering, Tunghai University, Taichung 407, Taiwan
d
Research Center for Smart Sustainable Circular Economy, Tunghai University, Taichung 407, Taiwan
e
Department of Chemical Engineering, National Cheng Kung University, Tainan 701, Taiwan

A R T I C L E I N F O A B S T R A C T

Keywords: The coexistence of algae and bacteria in nature dates back to the very early stages when life came into existence.
Algae The interaction between algae and bacteria plays an important role in the planet ecology, cycling nutrients, and
Bacteria feeding higher trophic levels, and have been evolving ever since. The emerging concept of algal-bacterial con­
Consortia
sortia is gaining attention, much towards environmental management and protection. Studies have shown that
Environment
algal-bacterial synergy does not only promote carbon capture in wastewater bioremediation but also conse­
Interactions
Industry 4.0 quently produces biofuels from algal-bacterial biomass. This review has evaluated the optimistic prospects of
Sustainability algal-bacterial consortia in environmental remediation, biorefinery, carbon sequestration as well as its contri­
bution to the production of high-value compounds. In addition, algal-bacterial consortia offer great potential in
bloom control, dye removal, agricultural biofertilizers, and bioplastics production. This work also emphasizes the
advancement of algal-bacterial biotechnology in environmental management through the incorporation of In­
dustry Revolution 4.0 technologies. The challenges include its pathway to greener industry, competition with
other food additive sources, societal acceptance, cost feasibility, environmental trade-off, safety and compati­
bility. Thus, there is a need for further in-depth research to ensure the environmental sustainability and feasi­
bility of algal-bacterial consortia to meet numerous current and future needs of society in the long run.

1. Introduction components from carbon dioxide, thereby supporting heterotrophic

Algae exist as a ubiquitous and diverse group of photosynthetic and
aquatic plants, which represents the fundamental producers and prom­
ising feedstock in the biological community. On the other hand, bacteria
are prokaryotic microorganisms that present themselves in an astro­
nomical amount of numbers in nature. Ecological studies have found
that specific groups of bacteria appear to be in close association with
certain algae, thus they synergistically influence each other’s physiology
and metabolism (Gothandam et al., 2018). Algal-bacterial interaction
describes the mutual relationship between two groups of microorgan­
isms, where algae are the primary producers that synthesize organic
bacteria that decompose organics (Kouzuma and Watanabe, 2015). The
close-knit association that exists between algae and bacteria stems from
the fact that many algae depend on bacteria for ultimate exogenous
sources of cobalamin (vitamin B12), thiamine (vitamin B1), and biotin
(vitamin B7) as a co-factor for B12-dependent methionine synthase
(METH) and growth (Xie et al., 2013). In the laboratory, B12-producing
heterotrophic bacteria was conceived to be able to supply cobalamin to
algae in co-cultures, in return for photosynthate and organic carbon.
Pennisetum purpureum grown with Halomonas sp. showed increased
bacterial growth and a simultaneous significant increase in the up-
regulation of vitamin B12 produced from the bacterial cells (Croft

* Corresponding author at: Department of Chemical and Materials Engineering, Tunghai University, Taichung 407, Taiwan.
** Corresponding author.
E-mail addresses: PauLoke.Show@nottingham.edu.my (P.L. Show), changjs@thu.edu.tw (J.-S. Chang).
1
Co-first author

https://doi.org/10.1016/j.biotechadv.2020.107684
Received 13 September 2020; Received in revised form 24 November 2020; Accepted 22 December 2020
Available online 30 December 2020
0734-9750/© 2021 Elsevier Inc. All rights reserved.
J.J.J.Y. Yong et al.
et al., 2005).
Exchange of micro- and macronutrients takes place to enhance the
growth of both algae and bacteria. Even though the presence of bacteria
in algal cultures was often seen as contamination, the perception has
changed in the past few years and the interactions between algae and
bacteria are considered promising for biotechnological applications
(Yao et al., 2019). Many recent studies have identified that algal-
bacterial symbiosis exhibited a positive effect on algal growth, floccu­
lation processes, spore germination, pathogen resistance, and morpho­
genesis, which are the integral steps towards sustainable algal
biotechnology (Fuentes et al., 2016) (Lian et al., 2018).
The symbiotic relationship of algae and bacteria have been receiving
tremendous emerging attention and are rapidly developing owing to its
benefits and applications in the biotechnology industry. Research re­
ports revealed that the algal-bacterial consortia possess much signifi­
cance in environmental management to improve wastewater
remediation, greenhouse gas mitigation, biomass and biofuel production
(Leong et al., 2019a). An algae-bacteria study using a co-culture of
Chlorella sorokiniana and Azospirillum brasilense showed high removal
rates of ammonium, phosphorus, and contaminants from wastewater
(De-Bashan et al., 2002). Algal-bacterial synergy also possesses the po­
tential for a high tolerance under unfavorable conditions, such as
nutrient limitation, high concentration of dissolved inorganic sub­
stances, extreme pH, and intrinsic temperature (Padmaperuma et al.,
2018) (Lian et al., 2018). Algae are also capable of increasing dissolved
O2 to approximately five times the saturation concentration to be used in
heterotrophic bacterial respiration and metabolism of dissolved organic
compounds in algal-bacterial consortia (Shah and Rodriguez-Couto,
2019).
There have been recent advancements in the use of microalgae for
biofuel production, wastewater treatment, and CO2 removal. However,
there is still a limited number of articles addressing the utilization of
algal-bacterial co-culture in the mitigation of environmental issues. The
major focus is placed on microalgal biofuel production and the removal
of nutrients and pollutants from wastewater using microalgal/bacterial
biomasses. Furthermore, there are inadequate descriptions of the chal­
lenges of algal-bacterial biotechnology as an ecosystem approach, as
well as its future development in environmental management. This re­
view explores the significance of algal-bacterial biotechnology in envi­
ronmental applications, including phycoremediation, carbon
sequestration, biorefinery, dye removal, bloom control, production of
high-value compounds, agricultural biofertilizers, and bioplastics. The
aim of this review is to open new perspectives to future developments of
algal-bacterial consortia by integrating Industry 4.0 technologies that
can greatly optimize cultivation productivity and sustainability. Asso­
ciated challenges and limitations faced in algal-bacterial based tech­
nologies are also discussed. Algal-bacterial consortia are of great
prospects in environmental protection, hence needing a deeper under­
standing of their ecology to promote sustainable patterns of production
in the green industry.
2. Potential applications of algal-bacterial consortia
biotechnology
Over the past decades, endeavors have been undertaken to
comprehend the potential and usefulness of algae and bacteria in
various industries. Their value has been recognized, particularly in
biotechnology for making extraordinary advancements in impacting the
life science sector. All through the history of mankind, there has been a
persistent improvement and development in the field of biotechnology
in line with human advancement. With our rapidly growing scientific
progress, biotechnology has evolved to tackle the practical impediments
and grave ecological concerns correlated with current anthropological
practices. Algal-bacterial symbiosis has been shown to have notable
impacts on the environmental treatment of various contaminants
(Subashchandrabose et al., 2011).
2
Biotechnology Advances 47 (2021) 107684
One of the earliest and conventional algal-bacterial consortia was in
open ponds. The affordable consortia were proven to enhance the
removal of organic nutrients, biochemical oxygen demand, and
contamination from wastewater. However, the conventional systems
were neither mixed nor aerated, hence the lower efficiency than what
could be achieved using systems or reactors developed later (Ludwig
et al., 1951). In the later stages of development, enclosed reactor sys­
tems known as photobioreactors containing algal-bacterial consortia
have evolved from conventional ponds and are gaining focus in treating
wastewater. Modernized reactors utilizing algal-bacterial symbiosis
result in efficient water treatments with the exchange of inorganic and
organic nutrients (Ramanan et al., 2016; Amini et al., 2020). Aerobic
bacteria oxidize organic carbon in sewage and convert it into CO2. The
produced CO2 was then utilized by algae for photosynthesis and con­
version of CO2 to algal biomass occurs. On the other hand, the appli­
cation of algal-bacterial biotechnology in the lysis of algal cells which is
important for biofuel production and algal bloom control was analyzed
and reported (Wang et al., 2020). Research interests spiked on bacterial
action in algal cell lysis and found algicidal bacteria in Cytophaga/Fla­
vobacterium/ Bacteroidetes (CFB) group or the γ-proteobacteria group.
Gram-positive genera employ direct and indirect modes of algicidal
methods that can kill harmful blooms (Roth et al., 2008). Accordingly,
algal-bacterial interaction has the potential to confront the challenges of
algal lysis through proper management and appropriate experimental
design. It can be an inexpensive, non-toxic, and scalable solution (Wang
et al., 2020). The evolution and versatility of the algal-bacterial mutual
relationship can be seen from the development of numerous conven­
tional methods to contemporary biotechnology. Today, it is apparent
that algal-bacterial biotechnology is constantly evolving and is well-
adapted, bringing boons and solutions to persistent environmental
problems. Various algal and bacterial-based biotechnology have
received abundant interest as environment-friendly methods for envi­
ronmental protection and management applications. Table 1 evaluates
the various biotechnological applications of algal-bacterial consortia,
with their advantages and disadvantages.
2.1. The importance of environmental management contributed by algal-
bacterial biotechnology
The mutualistic relationship and interaction between algae and
bacteria play a substantial role in ecosystems and environmental miti­
gation. Algae and bacteria acclimatize their metabolism to suit each
other’s needs, which can impact their physiology, change the chemistry
of their environment, and shape ecosystem diversity (Amin et al., 2015).
Algal-bacterial interactions offer several important solutions for envi­
ronmental impacts including wastewater management, aquaculture,
emissions control, and yield of biofuel. The ever-expanding scientific
and technical prospects make algal-bacterial biotechnology applicable
in various practical fields. Table 2 describes the contribution of the
algal-bacterial consortium in environmental management.
2.2. Algal-bacterial biotechnology applications in environmental
protection
2.2.1. Wastewater treatment
The extensive application of algal-bacterial biotechnology is in
wastewater treatment, as a biological tool for environmental pollutant
control. Algal-bacterial consortium has been utilized in water remedia­
tion for treating municipal wastewaters, industrial wastewater, rubber
effluent, and palm oil effluent. It has also been used in the bioremedi­
ation of contaminants, removal of nutrients, reduction of COD and BOD
since the 1950s (Molinuevo-Salces et al., 2019). Nutrients such as ni­
trogen and phosphorus necessitate the natural growth of non-
diazotrophic algae. Under nutrient-deprived conditions, algae might
face growth stagnation and eventual death because insufficient macro-
and micro-nutrients are incapable of sustaining their development
J.J.J.Y. Yong et al. Biotechnology Advances 47 (2021) 107684

Table 1
The modern biotechnology of algal-bacterial consortia.
Technology Application Description Advantage Disadvantage Reference

Phycoremediation Used in wastewater
bioremediation:
- Removal of organic and
nutrients pollutants.
- Removal of toxic
pollutants and heavy
metals such as uranium.
- Biological CO2
sequestration
Hydrothermal - Production of various
liquefaction alternatives of biofuels
(HTL) such as bioethanol,
biohydrogen, and
biodiesel.
Use of transgenic Used as bioinsecticides:
cyanobacterium
- Bacillus thuringiensis
controls mosquitoes and
black flies.
- 12-epi-hapalindole C
isonitrile and hapa-lindole
L eradicated 100% of the
larvae from Chironomus
riparius in 48 h.
Algal-bacterial symbiosis in open
or closed systems is used to treat
wastewater. Algae carry out
photosynthesis using CO2 from
bacterial respiration; the oxygen
requirement of organic carbon
degradation by bacteria is
fulfilled by O2 produced by
cellular respiration of algae,
without the need for additional
aeration.
Sequestration and sedimentation
occur via biosorption of up to
300 mg of uranium per kg dry
mass by algae and bacteria. The
uranyl ions are released by H+
after the algal cell wall material
sequesters them and is then
reduced from U(VI) to U(IV) by
heterotrophic bacteria using
carbon, nitrogen, and phosphorus
from dead algal cells.
CO2 is an essential inorganic
carbon source for algae growth
and CO2 fixation. On a mass basis,
the production of 1 kg of algal
biomass fixes 1.83 kg of CO2.
High biodiesel oil productivities
of microalgae/bacteria using CO2
captured enables conversion of
biomass to biofuel.
A biomass-to-liquid conversion of
symbiotically grown biomass of
algae and bacteria at medium
temperatures (300 ◦ C-350 ◦ C)
and high pressures (5–20 MPa)
via optimization in biomass
loading, residence time, and
reaction temperature. Through
HTL, microalgal lipids in the form
of glycerol are converted into
methanol, aldehydes, allyl
alcohol, and gas products.
Transgenic cyanobacteria
technology can be used to control
pests. Proteins such as lectins,
α-amylase, and Bacillus species
manifest a broad spectrum of
insecticidal activities. Cloning of
δ-endotoxin genes into
cyanobacteria in the breeding
zone will increase toxin
persistence, which results in the
mortality of pests.
- Reduces mechanical - Large carbon footprint (Rawat et al., 2016), (
aeration costs. compared to CAS-BNR Fito and Alemu, 2019)
- Low operational energy systems.
requirements.
- High nutrient removal
percentage.
- Reduces sludge
formation.
- Economically viable. - Production of toxic (Igiri et al., 2018), (
- Eco-friendly. metabolites by microbes. Show et al., 2017), (
- Reduce secondary - Non-biodegradability of Kalin et al., 2005)
pollution. heavy metals.
- Mitigation of greenhouse - Stable production of (Cheah et al., 2015), (
gases algae is required. Subashchandrabose
- High CO2 fixation rates et al., 2011)
that are 10–50 times
faster than terrestrial
plants
- Cost-effective
- No algal biomass drying is - High NOx emissions, due (Goswami et al., 2019), (
needed to high amounts of Sankaran et al., 2018)
- Biocrude oil produced via nitrogen in chlorophyll
HTL has low sulfur and and proteins in algal cells.
ash emissions
- Economically viable
compared to traditional
thermochemical
processes like pyrolysis
and gasification
- Eco-friendly - High industrial scaling (Ram Prasad and N.T,
- Reduce side effects of cost for the production of 2018), (Becher et al.,
agrochemicals microalgae and 2007)
- No synthetic residues cyanobacteria

(Ramanan et al., 2016). Conversely, the plethoric surge of nutrient flow
from industrial, municipal, and agricultural wastewater when released
to water bodies might accelerate extreme blooms of algae and cyano­
bacteria, promoting the persistent growth of harmful algae blooms
(HABs) (Heisler et al., 2008). In this context, the exchange of O2, CO2,
and NH+ 4 ions in algal-bacterial symbiosis is utilized in wastewater
treatment. The rapid bacterial oxidization of organic matter that re­
leases CO2 and NH+ 4 ions matches the rate of algal photosynthesis for O2
generation (Oswald et al., 1957).
A recent study showed that the consortium of a microalga
(C. vulgaris) and a bacterium (Bacillus licheniformis) was capable of
eliminating soluble chemical oxygen demand (sCOD), total dissolved
nitrogen (TDN), and total dissolved phosphorus (TDP) from wastewater
(Ji et al., 2018). At an algal-bacterial ratio of 1:3, the C. vulgaris-B.
licheniformis consortium exhibited the best performance with a decrease
in the concentration of sCOD from 175.78 mg/L to 23.64 mg/L,
indicating a high removal ratio of 86.55%. The efficiency of TDP
removal was recorded at a rate of 80.28%, in which its concentration
decreased from 4.97 mg/L to 0.98 mg/L. Under similar conditions, the
decrease in TDN concentration was measured from 31.23 mg/L to 3.45
mg/L, with an 88.95% removal rate. In another study, C. vulgaris was
used to reduce 71.6% ammonium ion, 28% phosphorus (P), and 61%
COD from industrial wastewater, whereas phototropic bacteria attained
88.2%, 99.6%, and 62.5% removal efficiency for P, NH4-N, and sCOD,
respectively after 24 h (Valderrama et al., 2002; Hülsen et al., 2014).
Table 3 shows the algal-bacterial symbiotic interactions used for nutri­
ents removal in wastewater treatment.
2.2.2. Phycoremediation of heavy metals
Algal-bacterial consortia have been advocated as a promising and
eco-friendly approach for detoxification of recalcitrant industrial
wastewater that is rich in toxic heavy metals. The discharge of large

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Table 2
The importance of algal-bacterial biotechnology in environmental management.
Aspect/Field Contribution Reference

Wastewater - To remove nutrients from wastewater to prevent eutrophication that can pose a (Qi et al., 2019), (Nguyen et al., 2019)
treatment considerable impact on water quality.
- Enable minimum sludge formation in the effluent.
- Contribution of algal-bacterial symbiosis in bio flocculation for microalgae
harvesting.
Aquaculture - Algal-bacterial consortium acts as an oxygen supply and reduces the amount of (Fang et al., 2017), (Natrah et al., 2014), (Toi et al., 2013)
toxic gases including hydrogen sulfide, carbon dioxide, nitrite, and ammonia.
- Acts as natural food base/feed and dietary requirements, sustaining production
of aquaculture, hence contribute to increased food production.
- To improve nitrogen assimilation via co-ingestion of algae and bacteria by
shrimp Artemia franciscana as a feed source.
Greenhouse gas - Reduce the emission of fuel gases due to the fast consumption of CO2 by the (Anbalagan et al., 2017)
mitigation algal-bacterial consortia.
- Photosynthetic microalgae release O2 during the production of biomass, which
helps in CO2 bio-mitigation.
Agriculture - Algal-bacterial interactions help in metal bioremediation to detoxify heavy (Hassan et al., 2017), (Secundo et al., 2016)
metals and chemicals in polluted soil.
- Vital to enhancing and stabilizing agricultural soil health/ properties.
- Remediate contaminated soils and sustain plant growth.
Marine - Algicidal bacteria against eukaryotic microalgae are used to control (Mayali and Azam, 2004)
environment phytoplankton that forms harmful algal blooms (HAB)/ red tides in marine
ecosystems.
- Able to decline/control red tides via decomposition of red tide organisms by
bacteria.
Energy source - Algal-bacterial based consortium can help improve the production of biomass (Beer et al., 2009), (Yao et al., 2019), (Olivares et al., 2019)
and biofuel owing to its higher conversion rate of solar energy into biomass
relative to terrestrial crops.
- Anaerobic digestion of microalgal biomass generates biogas:
Organic Matter + H2 0

→CH4 + CO2 + NH3

- Sustainable production of biofuels from algal-bacterial biomass.

amounts of these trace metals into natural waterways jeopardizes the
environment and human health. The most common heavy metals such as
lead, cadmium, mercury, selenium, arsenic, chromium, copper, nickel,
zinc, and silver hail from chemical industries. These pollutants that are
derived from agricultural and economic activities are carcinogenic to
human and aquatic life. Heavy metals are easily absorbed by living
creatures due to their high solubility in the aquatic environment and
eventually leads to the death of aquatic life and even prolonged toxicity
of the environment from chemical contaminants (Hong et al., 2020; Bar
akat, 2011). To alleviate the negative impacts of the metal-contaminated
wastewaters on the environment, a green bioremediation technology
known as phycoremediation is introduced to treat wastewater prior to
its release to other water bodies.
Phycoremediation is microbial remediation where algae and bacteria
are deployed for the removal of toxic and non-toxic substances in
wastewater via biotransformation processes or metabolic uptake (Rao
et al., 2019). Algae possess large surface to volume ratios and high
metal-binding affinity which increase the adsorption ability of heavy
metals on their cell surface. The efficient metal uptake and storage
systems also boost the adsorption and removal of metal pollutants from
wastewater (Ahmad et al., 2020). Algal cell walls are made up of
extensive matrix polysaccharides, proteoglycans, and cellulose that
contain negatively-charged (amino, carboxyl, hydroxyl, and sulfide)
groups that can act as binding sites for metals (Domozych, 2019; Sha­
mim, 2018). The biosorption of heavy metals is carried out by algae
through metal adsorption onto the cell surface, entrapment by cellular
components, cation exchange or complexation, and active transport
across the cell membrane (Kanamarlapudi et al., 2018; Derco and Vrana,
2018). Similarly, bacteria can exhibit metal adsorption behaviors and
bind aqueous cations of heavy metals to their acidic functional groups
on the cell wall (Ali Redha, 2020). Thus, the cultivation of an algal-
bacteria consortium in wastewater serves as a strategy for the
treatment of heavy metal polluted effluent.
Safonova et al. (2004) found that the consortium of Chlorella sp. ES-
13, Chlorella sp. ES-30, Scenedesmus obliquus ES-55, Phormidium sp. ES-
90 and Rhodococcus sp. Ac-1267, Kibdelosporangium aridum 754 resul­
ted in efficient degradation of pollutants. The consortium exhibited a
considerable decline in metal pollutants in industrial wastewater with
efficient removal of zinc up to 90%, phenol up to 85%, copper up to
62%, nickel up to 62%, manganese up to 70%, and iron up to 64%
(Safonova et al., 2004). The algal-bacterial consortium in another study
found that complete degradation of cadmium and 80% removal of
copper were attained within 5 min at an initial pH of 4.0 using a mixture
of cyanobacteria (i.e., Chroococcus sp., Pseudoanabaena sp., Leptolyngbya
sp.), chlorophycae (i.e., Scenedesmus sp., Tetraerdon sp., Chlorella sp.,
and Chlorococcus sp.) diatoms (i.e. Navicula sp., Nitzschia sp., Cyclotella
sp.) and bacteria (Loutseti et al., 2009). Table 4 summarizes the per­
formance of algal-bacterial consortia for the removal of heavy metals in
wastewater treatment.
2.2.3. CO2 sequestration
Global warming arising from the emission of greenhouse gases has
been the biggest environmental concern in the past decade. In Earth’s
atmosphere, the key greenhouse gases are carbon dioxide, methane,
nitrous oxide, hydrofluorocarbons, and perfluorocarbons (Maccarthy
et al., 2018). CO2, one of the main contributors to the greenhouse effect,
accounts for serious global warming due to its annual emission of over
36 billion tonnes. The atmospheric CO2 concentration is now over 400
ppm, which is at the highest levels in over 800,000 years (Ritchie and
Roser, 2017) (Show et al., 2017). The high level of CO2 emission is
primarily driven by anthropogenic activities such as increasing fossil
fuel consumption, deforestation, agriculture, and industrial processes. In
2012, International Energy Agency (IEA) stated that to attain the ±2 ◦ C
goal parallel to a stable CO2 concentration at 450 ppm, the current

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Table 3
Algal-bacterial symbiosis used in wastewater nutrient treatment.
Algae Bacteria Source of Wastewater Nutrients & Removal Efficiency Reference
− l
C. vulgaris B. licheniformis Synthetic wastewater sCOD 86.55% (175.78 mg L ) (Ji et al., 2018)
TDP 80.28% (4.97 mg L− l)
TDN 88.95% (31.23 mg L− l)
− l
C. vulgaris Pseudomonas putida Synthetic municipal NH+4 -N 85% (190 mg L ) (Mujtaba et al., 2017), (Shen et al.,
wastewater PO34-P 66% (40 mg L− l) 2017)
COD 86% (490 mg L− l)
C. vulgaris Activated sludge bacteria Municipal and industrial COD 79.7–83.8% (1130 mg L− l) (Zhu et al., 2019), (Ye et al., 2018),
wastewater NH3-N 75.5% (260 mg L− l) (Sepehri et al., 2020)
PO3−
4 -P 87.5% (1.50 mg L )
− l

Total phosphorus (TP) 100% (28.5

mg L− l)
Total organic carbon (TOC) 85.6%
(285 mg L− l)
C. vulgaris Rhizobium sp. Synthetic wastewater TOC 60.8% (127 mg L− l) (Ferro et al., 2019)
TN 69.1% (21.7 mg L− l)
TP 98.9% (0.07 mg L− l)
C. vulgaris B. licheniformis Municipal wastewater Nitrogen 88.82% (~75 mg L− l) (Ji et al., 2019)
Ammonium 84.98% (~60 mg L− l)
Orthophosphate phosphorus
84.87% (~6.2 mg L− l)
COD 82.25% (~275 mg L− l)
− l
C. sorokiniana Activated sludge bacteria Domestic wastewater NH+4 − N 82–98% (2500 mg L ) (Fan et al., 2020)
P 92–98% (2500 mg L− l)
COD 88–90% (2500 mg L− l)
Selenastrum Activated sludge bacteria Printing and dyeing COD 70–85% (Lin et al., 2019), (Ahmad et al.,
bibraianum industrial wastewater NH+4 84.9–89.7% 2017)
TP 30.2–37.7%
Navicula sp. Ammonia oxidizing bacteria (Comamonada- Synthetic wastewater COD 95% (600 mg. L− l) (Meng et al., 2019)
ceae and Nitroso-monadaceae) TP 31–42%
− l
NH+4 -N > 99% (50 mg. L )
Scenedesmus Filamentous bacteria Milk whey processing COD 93% (982 mg L− l) (Marazzi et al., 2020)
acuminatus wastewater TDN 88% (52 mg L− l)
TP 69% (17 mg L− l)
− l
NH+4 -N 88% (31 mg L )
Chlorella sp. Beijerinckia fluminensis Vinegar production COD 76.7% (740 mg L− l) (Huo et al., 2020)
wastewater TN 78.7% (20.5 mg L− l)
TP 74.8% (7.4 mg L− l)

emissions of 43 Gt CO2 should be reduced to 14 billion tons (Ul-Islam
and Wiley, 2019). Therefore, several key implementations are required,
including the improvement of energy efficiency (43%), renewable en­
ergy (28%), and CO2 capture and storage (CCS) technology (22%) (IEA,
2012).
Nowadays, various technologies have been proposed in light of the
rapidly increasing energy demand and the trend towards renewable
energy. Among the CO2 capture technologies suggested, the direct use of
CO2 in photosynthetic algal or bacterial cultivation is found to be pro­
spective because algae can consume atmospheric CO2, CO2 released
from power plants, CO2 from soluble carbonate. At the same time, the
derived biomass can act as feedstock to produce biofuel. The CO2-con­
verted chemicals and energy products are promising for they have high
market potential. They are also beneficial because microalgae/cyano­
bacteria have about 10 to 50-folds higher CO2 fixation rates when
compared to terrestrial plants. The rapid evolution of algae strains can
be achieved by high-throughput technologies because algae are unicel­
lular organisms that duplicate by division (Show et al., 2017) (Sub­
ashchandrabose et al., 2011). Algal-bacterial consortia with high biofuel
productivities are capable of CO2 capture and sequestration, ultimately
converting them from algal biomass to biofuel. On a mass basis, 1 kg of
algal biomass production consumes 1.83 kg of CO2 (Cheah et al., 2015).
Additionally, bacteria also play an essential role in the carbon cycle.
Acetogenic bacteria, Clostridium autoethanogenum, fixes CO2 (in the
presence of H2) and CO via the Wood–Ljungdahl pathway, the linear
pathway for CO2 fixation (Liew et al., 2016). The β-proteobacteria Ral­
stonia eutropha could capture and convert CO2 into useful products by
fixing carbon in the form of polyhydroxyalkanoate (bioplastics)
(Bringham et al., 2011). In algal-bacterial consortia, their interactions
are symbiotic, in which the exchange of substrates CO2 and O2 are
required for both algae growth and CO2 fixation. A study has shown an
increase in carbon fixation rate by 20.3% in Thalassiosira pseudonana and
heterotrophic bacteria Pelagibacter sp. HTCC1062 (SAR11) co-cultures
compared to T. pseudonana monocultures. The high carbon capture is
used for the algal photosynthetic activity and growth, which subse­
quently resulted in enhanced biogas production (Moore et al., 2020).
Gao et al. (2018) also reported that an algal-bacterial consortium of
C. vulgaris and activated sludge bacteria exhibited an optimal CO2
removal efficiency of 63.48% (Gao et al., 2018). In view of current de­
velopments in algal-bacterial consortia and understanding their envi­
ronmental, commercial, and economic benefits, it would become a
practicable technology in the near future in CO2 removal and green­
house gas mitigation. The application of algae and bacteria in the
sequestration of CO2 can be an ultimate potential alternative in miti­
gating energy and environmental issues. Table 5 shows the CO2 removal
efficiency by algal-bacterial consortia.
2.2.4. Biofuel production
In this world where fuel energy demand is soaring along with the
increasing population, depletion of fossil fuels is imminent due to its
profound and prevalent usage for anthropogenic activities and economic
development. Every year, the burning of fossil fuels is responsible for an
average emission of 29 gigatonnes of CO2 and is hitting a total of 55.3
gigatonnes until now (Andersen et al., 2019). The greenhouse gas
emission from fossil fuels is the source of global warming and climate
change. Thus, carbon-neutral bioenergy is required to meet the eco­
nomic and environmental sustainability to curb climate damage issues
(Lee et al., 2019).

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J.J.J.Y. Yong et al. Biotechnology Advances 47 (2021) 107684

Table 4
Algal-bacterial consortia in the removal of heavy metals in wastewater.
Algae Bacteria Source of Wastewater Heavy Metals & Reference
Removal Efficiency

C. sorokiniana DBWC2, Chlorella sp. DBWC7 Klebsiella pneumoniae ORWB1, Acinetobacter Artificial wastewater Copper 78.1% (6.79 mg (Makut et al.,
calcoaceticus ORWB3 L− l) 2019)
Chromium 76.3% (24.7
mg L− l)
Cadmium 55.2% (1.3
mg L− l)
Nickel 70% (18.6 mg
L− l)
Lead 65.1% (12.4 mg
L− l)
C. vulgaris Phototrophic bacteria (PBB) Piggery wastewater Zinc 98% (0.78 mg. L− l) (García et al.,
2019)
Chlorella sp., Nannochloropsis sp., Scenedesmus bijugatus, Bacteria from activated sludge in a municipal Municipal wastewater Copper 85.1% (0.02 mg. (Sharma et al.,
Chlamydomonas reinhardtii, Oscillatoria wastewater treatment plant L− l) 2020)
Chromium 75.2% (0.31
mg. L− l)
Lead 98.2% (0.47 mg.
L− l)
Cadmium 99.6% (0.004
mg. L− l)
Zinc 57.7% (0.40 mg.
L− l)
Nickel 99.4% (0.20 mg.
L− l)
Chlorella sp., Phormidium sp. Wastewater bacteria Tannery wastewater Nickel 74.54% (Sahoo et al.,
Chromium 80% 2019)
Ulva lactuca Gram-negative bacteria Industrial wastewater Lead 100% (1000 mg. (Aly and Amasha,
L− l) 2019)
C. vulgaris-BH1 Exiguobacterium profundum-BH2 Artificial metal-rich Copper 78.67% (729 (Batool et al.,
wastewater mg. L− l) 2019)
Chromium 56.4% (913
mg. L− l)
Nickel 80% (331 mg.
L− l)
Spirulina sp. LEB 18 Aquaculture effluent bacteria Aquaculture Bromine 19.35% (250 (Cardoso et al.,
wastewater μg L− l) 2020)
C. vulgaris, Scenedesmus quadricuda, Spirulina platensis Wastewater bacteria Urban wastewater Cadmium 86% (0.7 ± (Abdel-Razek
0.015 mg. L− l) et al., 2019)
Nickel 95% (0.5 ± 0.07
mg. L− l)
Lead 87% (1.3 ± 0.04
mg. L− l)
C. vulgaris Proteobacteria Piggery wastewater Zinc 83% (0.16 ± 0.02 (García et al.,
mg. L− l) 2017)
Scenedesmus sp. Activated sludge bacteria Sewage wastewater Chromium 100% (1500 (Lei et al., 2018)
mg. L− l)
Copper 59% (1500 mg.
L− l)
Nickel 65% (1500 mg.
L− l)
Lead 83% (1500 mg.
L− l)
Zinc 95% (1500 mg.
L− l)

Table 5
Algal-bacterial consortia in the sequestration of CO2.
Algae Bacteria CO2 Removal Efficiency Reference

Tetraselmis chui, Nannochloropsis gaditana Algal pond bacteria 89–97% (Anbalagan et al., 2017)
C. vulgaris Activated sludge bacteria 69.93–88.27% (Zhang et al., 2017)
C. vulgaris Anaerobic bacteria 190.9 ± 8.6 mg/Ld− 1 (Yadav et al., 2020)
C. vulgaris Activated sludge bacteria 53.24–63.48% (Gao et al., 2018)
Scenedesmus obliquus Activated sludge bacteria 51.46–62.29% (Gao et al., 2018)
Spongiochloris sp. Aerobic heterotrophic bacteria 2.9205 g L− 1d− 1 (Abid et al., 2017)
1 1
Thalassiosira pseudonana Pelagibacter sp. HTCC1062 (SAR11) 0.0249–0.0939 pmol C cell− h− (Moore et al., 2020)
Chlorella sp. Cupriavidus necator ~20% (Yi et al., 2020)
Acutodesmus obliquus Photobioreactor bacteria 61.6–90.8% (Toledo-Cervantes et al., 2018)
C. sorokiniana Sulfurihydrogenibium yellowstonense, Escherichia coli 20 ± 0.7 WAUa/mg (Salbitani et al., 2020)
C. vulgaris Nitrifying bacteria 90% (156 mg) (Sepehri et al., 2020)
a
Wilbur-Anderson Unit.

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J.J.J.Y. Yong et al. Biotechnology Advances 47 (2021) 107684

Fig. 1. Transesterification of triacylglycerol with alcohol into glycerol fatty acid methyl esters in the presence of a catalyst.

Biofuels including algal fuels are now a sought-after preference as a trophic chain levels (Medlin and Cembella, 2013). A huge amount of
renewable and sustainable replacement for fossil fuels. Biofuels gener­ HABs affects the coastal ecosystems as the degradation of high biomass
ated from algal biomasses contribute a considerably lower emission of blooms causes anoxic events, depleting oxygen levels, and leading to
greenhouse gases, are non-polluting, accessible, sustainable, and reli­ toxic impacts on fisheries (Burkholder, 2002). Mass mortalities of fish
able as they can be obtained from renewable sources (Liu et al., 2013). and other marine invertebrates occur when mucilage-producing or
The high carbohydrate and lipid content of algae, particularly micro­ toxin-producing algae invade their gills (Zingone and Oksfeldt Ene­
algae possess great capabilities in renewable energy production as a voldsen, 2000). Bacteria play an important role in controlling the
higher amount of energy can be obtained from algae compared to other blooms and dying out of HABs. Biological control of HABs using bac­
biomasses. High photosynthetic conversion of solar energy into chemi­ terial strains was reported to be an eco-friendly solution without causing
cal energy comparative to terrestrial biofuel feedstock makes algae the secondary pollution. Bacteria were found to use specific mechanisms for
prospective candidate for the synthesis of biofuel. Algal remnants after
extraction of biofuel can be utilized to yield biodegradable methane,
ethanol, biodiesel, and biogas (Yao et al., 2019) (Chia et al., 2018). The Table 6
Symbiotic algae and bacteria involved in microalgal-based biofuel production.
oxygen elimination during respiration of bacteria symbionts is necessary
for the activation of a Fe-dependent hydrogenase in algae which acts as Microalgae Bacteria Biofuels Conversion Reference
an efficient catalyst to enhance the production of molecular hydrogen Chlorophyta Rhodocyclaceae Lipid contents: 31.2 (Meng et al.,
(Lakatos et al., 2014) (Stripp and Happe, 2009). Photosynthetic algae to 59.6 mg g− 1 SS 2019)
can capture and convert solar energy into electricity with nil external Chlorella sp., Xanthomonadaceae, Biodiesel yield of (Liu et al.,
Scenedesmus sp. Rhodobacteraceae 66.21 ± 1.08 mg g− 1 2018a)
input of organics via the synergistic relationship between algae and
SS with large
bacteria in the form of microalgal fuel cells (Gajda et al., 2015). quantities of poly-
Transesterification of algae-produced triacylglycerol and other lipids unsaturated FAMEs
into fatty acid and methyl esters can yield biodiesel (Chew et al., 2017). Chlamydomonas Pseudomonas sp. Hydrogen 154 mL (Ban et al.,
reinhardtii strain D L− 1 2018)
Fig. 1 shows the transesterification of algal lipid (triacylglycerols) into
FACHB-265
biodiesel. The symbiotic association between Auxenochlorella proto­ Chlamydomonas Escherichia coli, Hydrogen 125 mL (Ban et al.,
thecoides and Escherichia coli led to an increase in algal growth and reinhardtii Pseudomonas sp. L− 1 2018)
double neutral lipid content yield compared to axenic growth (Higgins FACHB-265 strain D
and VanderGheynst, 2014). Biogas can also be produced from algal Chlamydomonas Pseudomonas putida Hydrogen 23.1 mL (Fakhimi
reinhardtii 704 291 L− 1 et al., 2019)
residue biomass through anaerobic digestion as the biochemical
Chlorella sp., Xanthomonadaceae FAMEs: (Liu et al.,
composition of microalgae (e.g., iron, cobalt, zinc) meets the nutrient Scenedesmus sp. Methyl palmitate 2018b)
requirement of anaerobic microbiota (Richmond, 2008). Moreover, 24.06%
microorganisms, such as Cellulomonas uda and Clostridium thermocellum, Methyl palmitoleate
are known for their cellulolytic properties that can enhance biofuels and 22.44%
Tetraselmis striata Pelagibaca Triacylglycerol: (Park et al.,
biohydrogen production (Rambabu et al., 2020; Swathy et al., 2020). bermudensis Palmitic acid 35% 2017)
Table 6 shows the symbiotic algae and bacteria involved in microalgal- Linolenic acid 20%
based biofuel production. Stearic acid 10%
C. vulgaris Brevundimonas Lipid content: 272 (Xie et al.,
mg/L SCOD ADE 2018)
(43.21%)
2.3. Minor applications or by-products of algal-bacterial biotechnology
Desmodesmus spp. Nitrifying bacteria FAME: (Hernández-
Saturated 38.78% García et al.,
Algal-bacterial biotechnology is vastly applicable in a plethora of Monounsaturated 2019)
industries from commercial to environmental, owing to its extensive 11.90%
biological diversity. Additionally, it possesses intrinsic advantages such Polyunsaturated
62.01%
as low production cost, enormous growing capacity, and environmental- Leptolyngbya sp. Wastewater-borne Lipid content: 13% (Tsolcha
friendly nature. Algal-bacterial interactions also contribute to certain bacteria (w/w) et al., 2017)
environmental events which are discussed in the further sections. Fig. 2 Saturated FA: 85.4%
shows the applications of symbiotic algal-bacterial consortia. Chlorella sp., Rhodobacter sp., Lipid content: 17% (Cho et al.,
Acutodesmus Rhodococcus sp. (17.2 mg/L d− 1) 2017)
sp.,
2.3.1. Bloom control Scenedesmus sp.
Harmful algal blooms (HABs) have become a global environmental Scenedesmus sp. A. brasilense Unsaturated FA: (Contreras-
problem, affecting aquatic life and the quality of potable water, and 89.1% Angulo et al.,
human health. Harmful algae are of high taxonomic diversity and can Saturated FA: 10/9% 2019)
Total FA: 56.8%
result in various toxins levels and relative mechanisms at different

7
J.J.J.Y. Yong et al.
Fig. 2. The contributions and applications of algae-bacteria symbiosis.
the termination and decomposition of HABs, such as direct attack using
cell-to-cell contact by the production of N-acyl-homoserine lactone
signals or indirect attack by producing extracellular algicidal sub­
stances. These extracellular compounds can be proteins, amino acids,
peptides, hydroxylamine, lipid peroxidation, and biosurfactants (Shu­
nyu et al., 2006) (Zheng et al., 2015). In the removal of HABs, bacteria
are capable of accelerating algal cell aggregation through the induction
of extracellular polymeric substances (EPS) by algal cells and the pro­
duction of bioflocculants that can cause direct accumulation and settling
of algal cells (Sun et al., 2018). An article has documented that het­
erotrophic bacterial communities such as Porphyrobacter and Fla­
vobacteriaceae are responsible for the algal aggregation in the
development of Microcystis aeruginosa bloom in natural waters (Shen
et al., 2011). Roseobacter species is a key ecological player in open wa­
ters rich in dimethylsulfoniopropionate (DMSP)-producing algal
blooms, in which they can degrade DMSP (Gonzalez et al., 2000). A
study has shown that 17 strains out of 20 algae-lysing bacteria were able
to reduce water blooms via the production of algicides in aquatic envi­
ronments. The algae-lysing bacteria manifested anti-Microcystis activ­
ity, which is an efficient anti-cyanobacterial/algae compound that can
depress algal bloom (Imamura et al., 2001). The engagement of algae
and bacteria in dynamic symbiosis also manifested that roseobacticides
Phaeobacter gallaeciensis can produce algicidal compounds against Emi­
liania huxleyi and may help to terminate algal bloom (Seyedsayamdost
et al., 2011). Furthermore, the distribution of Flavobacterium sp. on the
Gymnodinium nagasakiense red tide showed strong inhibitory effects on
the dinoflagellate during its algorithmic growth phase by producing a
basic compound with a molar mass of less than 500 Da. The results of
this study asserted that certain bacteria are effective against bloom algae
and may eventually be used for the elimination of algal blooms (Fukami
et al., 1991). Similarly, strains of isolated algicidal Saprospira spp. were
found to attack the cells of diatoms Chaetoceros calcitrans and other
unicellular algae such as Isochrysis sp. and Chaetoceros sp. (Sakata,
1990). The author suggested that the findings of marine algicidal bac­
teria provide insightful comprehension in the studies related to algal
bloom regulation in marine ecosystems.
2.3.2. Dye removal
Synthetic dyes are commonly and widely used in various commercial
8
Biotechnology Advances 47 (2021) 107684
industries including textile, plastics printing, paint, pharmaceuticals,
food, and cosmetics. Despite the infinite applications, the improper
management and disposal of dyes can pose detrimental environmental
threats due to its harmful chemical properties and toxic nature (de
Bashan and Bashan, 2010). These dyes retain xenobiotic attributes,
hence the resistance to biodegradation. Immediate approaches are
needed to tackle the environmental impacts as incessant production and
industrial usage of dyes are increasing. After several biological measures
of wastewater dye removal including enzymatic decomposition and
aerobic-anaerobic biodegradation, it was found that the biotechnolog­
ical approach using microalgae can be more beneficial compared to the
other biological methods. Many algae are capable of effective decolor­
ization and microbial degradation of dyes in effluents (El-Sheekh et al.,
2009). Studies have shown that immobilized thermophilic cyanobac­
teria Phormidium sp. could decolorize dyes by adsorbing the dye on the
algal cell surface, showing a relatively high dye removal rate at 45 ◦ C
and 50 ◦ C incubation temperatures (Ertuǧrul et al., 2008). The ability of
different algae, namely C. vulgaris, Lyngbya lagerlerimi, Nostoc lincki,
Oscillatoria rubescens, Elkatothrix viridis, and Volvox aureus for dye
decolorization is in the range of approximately 4 to 95% for methyl red-
orange II, G-Red (FN-3G), basic cationic, and basic fuchsin dyes (El-
Sheekh et al., 2009). Carbajo Arteaga et al. (2018) revealed that the
microalgae C. vulgaris also portrayed a high removal efficiency for ani­
line blue dye at a concentration of 25 ppm (Carbajo Arteaga et al.,
2018). Furthermore, for microalgae-based textile wastewater bioreme­
diation, five microalgae strains namely, Anabaena flos aquae, Anabaena
variabilis, Nostoc elepsosporum, Nostoc linkia, and C. vulgaris were eval­
uated for red color removal of the effluent. The rapid dye absorbance
percentage by the microalgae were recorded as 50.81%, 100%, 79.03%,
88.71%, 96.16%, respectively proving their high removal rates of syn­
thetic colouring in wastewater (Ghazal et al., 2018). Another study
showed that the biodegradation and biosorption of tributyltin (TBT), an
antifouling agent used in paints that exceed acute toxicity levels, by
C. vulgaris in six consecutive treatment cycles would result in over 90%
of TBT removal at different contamination levels (Luan et al., 2006).
2.3.3. Biopolymers and bioplastics
The demand for plastics and plastic-based products has increased
largely, exerting strain on the ecosystems and generating major plastic
J.J.J.Y. Yong et al.
pollution worldwide. These petrochemical-based products that accu­
mulate in the environment can destroy most marine wildlife by suffo­
cation and also their habitats due to their non-biodegradable nature.
Therefore, green and sustainable alternatives to conventional plastics
are of grave attention, as they would mitigate plastic accumulation and
reduce environmental pollution. Microbial biomass metabolites such as
lipids, polysaccharides, and starch can be converted into plastics. It was
found that algae-based biomass is a potential source for biopolymer
synthesis due to their high photosynthetic efficiency and growth rate,
high amount of carbohydrates, and high carbon fixation to produce
bioplastics (Chia et al., 2020). Bioplastics, such as Poly­
hydroxyalkanoate (PHA), are known for their thermoplasticity, enan­
tiopurity, rapid biodegradability, and biocompatibility (Leong et al.,
2019a, 2019b; Zhang et al., 2019; Leong et al., 2014). Exploitation in the
production of bio-based polymers is mainly due to the valuable and
atypical sulfated polysaccharide metabolites in algae which are not
present in any other organism, namely, alginate, ulvan, and laminarin.
Algae or cyanobacteria are an excellent feedstock for plastics and
polymers synthesis as they have a strong ability to grow in diverse en­
vironments as well as promote carbon mitigation (Noreen et al., 2016).
Among PHA, Poly(3-hydroxybutyrate) (PHB) is also a 100% biode­
gradable bioplastic produced by several bacteria as an intracellular
carbon storage material. The properties of pure PHB such as hydro­
phobicity, thermoplastic processability, biodegradability, and biocom­
patibility are seen as excellent raw materials for biodegradable plastics
(Balaji et al., 2013). A research study focusing on Chlorella and Spirulina
in the development of algal-based bioplastics observed that the effective
glycerol plasticization was accomplished at a ratio of 4:1 of biomass to
glycerol (Zeller et al., 2013). The potential conversion of algae biomass
into bioplastics by blending algae populace such as Nannochloropsis and
catfish algae was described in a study, showing that the prospects of
biopolymers production from algal biomass can be exploited to create a
plastic-free future in all industries (Wang et al., 2016).
2.3.4. Soil biocontrollers and biofertilizers in agriculture
Nowadays, modern agriculture is heavily dependent on synthetic
inputs, resulting in negative effects on vegetation, soil structure, and
even water quality. Biofertilizers or biocontrol agents are natural fer­
tilizers containing live microorganisms that colonize the rhizosphere of
plants and promote growth via enhancement of nutrient uptake besides
improving soil fertility. They are significant in replacing chemical fer­
tilizers in intensive crop production. Microbiological fertilizers are now
seen as a promising approach for environmental-friendly organic agri­
cultural practice (Mącik et al., 2020). Studies have identified that
microalgae (Chlorella sp. and Scenedesmus sp.) and purple phototrophic
bacteria (PPB) are capable of removing and recovering nutrients from
anaerobic digestion effluent which can be used as biofertilizers in ni­
trogen and carbon cycling (Huy et al., 2020). The rhizosphere bacterial
community in the soil act as biostimulants which contribute to plant
growth, proving that biofertilizers execute similar effect as chemical
fertilizers but in an eco-friendly manner. The effectiveness of microalgae
and PPB as organic fertilizers can be seen from their profound effect on
C- and N-cycling capacities and also the improvement in crop and soil
performances (Zarezadeh et al., 2019). The effect of microalgae and
bacterial community as fertilizers are investigated and have been re­
ported to show higher mass and nutrient contents (phosphorus and ni­
trogen) followed by an increased plant yield (Mulbry et al., 2005).
Moreover, the ability of PPB-based fertilizers was demonstrated through
the enhanced growth of Stevia rebaudiana plants, showing an increased
yield of 69.2% with a combination of foliar spray method. The irrigation
of the rhizosphere with PPB promoted the soil dehydrogenase activity
and induced plant shoot biomass (Wu et al., 2013). In another study, the
inoculation of purple phototrophic dinitrogen-fixing bacteria, Rhodop­
seudomonas palustris strains in paddy soils were found to increase the rice
grain yields by 29%, as the bacterial community strongly promoted
nitrogenase activity and growth of rice straw (Harada et al., 2005).
9
Biotechnology Advances 47 (2021) 107684
Algal-bacterial biomass obtained from wastewater treatment processes
can potentially be used as agricultural fertilizers due to the presence of
nitrogen and phosphorus accumulated in the biomass. The biofertilizers
were used in as many as two million hectares for the cultivation of rice in
India (Su et al., 2011; Muñoz and Guieysse, 2006). The use of algal-
bacterial biofertilizer can enhance and strengthen on-farm sustainabil­
ity, productivity, and profitability owing to their elemental abilities of
nitrogen fixation, high yield, and alteration in physicochemical prop­
erties of soil (Bansal et al., 2018).
2.3.5. High-value compounds
In recent years, due to initiatives aimed at renewable sources of
energy, and the technological and economic advantages algae have over
oleaginous plants, they are being tested as producers of biofuels exper­
imentally (Brennan and Owende, 2010). The interaction between algae
and bacteria can foster the development of algae and also the production
of high-value molecules. Studies have demonstrated the roles played by
bacterial communities in the enhancement of carbon storage in micro­
algae, which is obtained in algal growth specifically under heterotrophic
conditions (Fuentes et al., 2016). C. vulgaris was co-immobilized with
plant growth-promoting bacterium A. brasilense and the alga showed an
increase in the accumulation of fatty acids, lipids, and activity of acetyl-
CoA carboxylase (ACC) (Leyva et al., 2014). The bacterium can augment
the growth of algal cells, with an increase in the size of algal cells and its
metabolic compounds such as lipids, fatty acids, and photosynthetic
pigments (De-Bashan et al., 2002). The joint immobilization of
A. brasilense with C. vulgaris and C. sorokiniana in alginate beads were
also found to improve the total carbohydrates and starch content after
an incubation period of 96 h. The study concluded that the bacteria
A. brasilense is an important biological factor in altering the composition
of algae, enhancing algal growth, producing high volumetric produc­
tivity, and carbohydrate yield, mainly under dark and heterotrophic
conditions (Choix, 2012). The effect of A. brasilense on the enzymatic
activity of ADP-glucose pyrophosphorylase (AGPase) using D-glucose or
Na-acetate as the carbon source was studied and it was observed that
this microalgal-bacterial interaction enhanced AGPase activity and
resulted in higher starch accumulation in C. vulgaris (Choix et al., 2014).
Moreover, a Rhizobium sp. isolated from a bacterial biofilm was
capable of producing acyl-homoserine lactones (AHLs) and stimulating
the growth of Botryococcus braunii which is competent in the synthesis of
compounds such as polysaccharides and carotenoids (Rivas et al., 2010).
The cyanobacterium S. platensis also has great potential in the produc­
tion of high-value natural components. C-phycocyanin (C-PC) extracted
from S. platensis is a widely used pigment in the food and cosmetics
industries (Chew et al., 2019). Therefore, the control of algae and bac­
teria interactions is fundamental in promoting algal growth and the
accumulation of useful, high-valuable compounds.
3. Future developments of algal-bacterial biotechnology in
environmental protection
Algal-bacterial technology is gaining prominence as the current
fragmentary comprehension of ecology and evolution of algal-bacterial
relationships are slowly and surely being exploited in algal-based
biotechnology. This includes the cultivation of algal-bacterial feed­
stock, biorefineries, and environmental technologies. In the bio-
industry, controlled algal-bacterial co-cultures that are becoming prev­
alent are harnessed to increase productivity, product diversity, and
mitigate contamination issues (Padmaperuma et al., 2018). Tradition­
ally, the growth of algae was initiated in simpler open ponds or raceway
ponds. Over the years, larger-scale cultivation of microalgae and even
co-cultivation of algae and bacteria are done in either open or closed
systems, the two most common methods. Open co-cultivation systems
include open ponds and tanks whereas controlled closed systems use
different types of bioreactors. Generally, open pond cultivation systems
have been associated with contamination issues by unwanted species,
J.J.J.Y. Yong et al.
light deficiency, temperature fluctuation, and scale-up space re­
quirements despite being an energy-saving alternative. However, it can
be a more suitable measure for the treatment of aquaculture wastewater
compared to closed glass bioreactors (Han et al., 2019). The develop­
ment of photobioreactor (PBR) technology has made algal cultivation
and production of high-value metabolites commercially viable. Olaizola
(2000) reported the use of proprietary computer-driven technology to
control outdoor photobioreactor for the exploitation of microalgae and
it has been proven to improve the productivity of biomass processing
and drying as well as pond management (Olaizola, 2000). Closed
cultivation systems, PBRs, are found to be more efficient with respect to
overall conditions and growth parameters such as light availability,
culture mixing, pH, temperature, and mass transfer in a rather
economically viable way (Panahi et al., 2019; Chang et al., 2017). On
top of that, PBRs appear to be a highly attractive approach for waste­
water treatment and heavy metals removal compared to open systems
owing to their favorable advantages, including high areal productivity,
low contamination, less CO2 losses, higher photosynthetic efficiency,
and high bio-oil yield efficiency in a controlled environment. However,
the limiting issue is the high capital expense, however productive
cultivation in large volumes can be easily achieved (Show et al., 2017)
(Pal et al., 2019). Intensive research efforts are highly in demand to
make the most of algal-bacterial consortia in closed systems and develop
a cost-competitive growth model that can be used in the sustainable
trend of environmental protection (Kumar and Singh, 2019; Hannon
et al., 2010).
In addition, the algal wheel is also a new upcoming technology that
applies symbiotic algal-bacterial consortium in fully engineered waste­
water treatment systems. Pre-screened wastewater is channeled into
secondary treatment tanks where algal-bacterial growth systems under
appropriate nutrient-laden conditions exchange respiratory gases and
organic nutrients. The rotating wheel of this system, driven by air
bubbles, provides intentional exposure to solar energy for algal photo­
synthetic uptake of nutrients. The algae biofilm formed on the surface of
the rotating wheel undergoes biological aeration during photosynthesis.
Conversion of biomass then happens when heterotrophic bacteria
perform various biological processes. Fig. 3 illustrates the algal wheel
used for wastewater treatment. This inventive system capitalizes on the
symbiosis between algae and bacteria where the by-products of one
group are the inputs for the other. Several advantages are achieved in
Fig. 3. The algal wheel utilizes the symbiosis of algae-bacteria in the application of wastewater treatment.
10
Biotechnology Advances 47 (2021) 107684
such a system, including being cost-effective, low energy requirement,
low environmental footprint, the high endurance of flow rate variability,
noiseless and odorless systems in the treatment of domestic or com­
mercial wastewater (Bansal et al., 2018) (OneWater Inc., 2019).
In the upcoming development of algal-bacterial biotechnology, the
concept of “Industrial Revolution 4.0” plays a critical and epoch-making
role in flourishing the industry with smart technologies. Industry 4.0 is a
new manufacturing approach that focuses heavily on intelligent ma­
chine learning, real-time data exchange, interconnectivity, and auto­
mation, including Cyber-Physical Systems, Internet of Things (IoT),
Smart Factory, and Internet of Services (Frank et al., 2019). Recent
advances in technologies coupled with smart machines and remote
monitoring have created new possibilities to further enhance the
applicability and adaptability of algal-bacterial consortia. This creates
leaps in environmental-friendly algal-based biotechnology. The inte­
gration of Industry 4.0 technologies like automation using sensors and
actuators can assist not only in the smooth process flow in the cultiva­
tion, harvesting, and extraction of microbial biomasses, but also help in
energy saving and reduction in environmental impact. The construction
of optical density sensors using near-infrared (NIR) light transmittance
sensors can be utilized to sustain a high biomass yield rate, where data
recorded are continuously transferred to, interpreted, displayed, and
filed using the LabView software, an online monitoring system that
provides the data of biomass density and algal growth rate. This algal
density control system employs the continuously-measured optical
density values and allows real-time control of the biomass density in the
algal culture (Sandnes et al., 2006). In this context, a network of plug-
and-play sensors not only provides real-time monitoring of algal-
bacterial co-culture productivity and tracking of algal growth, but the
cultivation and harvesting systems can be automated to reduce oper­
ating energy and expenses (ElFar et al., 2020). The technologies of In­
dustry 4.0 can be coalesced a step further in this algal-bacterial
biotechnology by implementing a simulation, or known as Digital Twin,
of the algal-bacterial culture in the plants using real-time sensor data
acquisition. This simulation-based virtual replica technology can be
used to optimize algal-bacterial consortia operation, allow concurrent
prediction of future cellular yield, and centralized analysis of biomass
production to address the product requirements and minimize wastes.
For instance, a fully-realized algal biorefinery would integrate an
automated serial downstream extraction of algal-bacterial biomasses for
J.J.J.Y. Yong et al.
a controlled cellular yield of specific compounds or by-products driven
by current demand in lieu of the conventional linear production stock­
piling that awaits demand (Uhlemann et al., 2017). A smart meter, in the
context of a Smart Grid connected to a network operator, can also be
incorporated to measure the energy consumption of affected algal pro­
cesses or facilities, enhancing intelligent energy and cost management
which further diminishes the ecological footprint and impact. Such en­
ergy management systems can reduce waste through algal waste opti­
mization and increase sustainable performance in large-scale algal
production facilities (Gabriel and Pessl, 2016).
With the rapid development and adoption of Industry 4.0 technolo­
gies in algal-based biotechnology, new levels of consistent optimization
and productivity can be achieved, leading to almost zero downtime in
production such as harvesting and biomass production which facilitate
the generation of biofuels, remediation processes, and eco-friendly
biofuels production. This will help meet the world demand for renew­
able sources fast and smoothly. Holistically, it is important to analyze
Industry 4.0 as a great approach in algal-bacterial biotechnology for a
clean and sustainable environment to open up new horizons and po­
tential benefits, continue to revolutionize fields of biotechnology over
the next few years.
4. Challenges of algal/bacterial-based biotechnology
4.1. Potential of algal-bacterial biotechnology in the current green
industry
The prospect of algal-bacterial symbiosis was tested for the green
industry, including the removal of nutrients and heavy metals in
wastewater treatment, CO2 mitigation, and biofuel generation. The
algal-bacterial consortia reduce land footprints and the demand for fresh
water, hence low competition with agricultural lands and water for feed
production (Morello and Pate, 2010). However, sparse knowledge of
algae and bacteria interactions lead to several challenges. The harvest of
algae biomass from liquid cultures for biofuel production is a costly
process that requires high energy consumption; solely the dewatering
step takes up nearly 30% of the total production cost of biomass (Yew
et al., 2019). Efforts and investment in biotechnological development
are needed for algae growth and harvesting strategies to be economi­
cally feasible, in an energy-efficient manner and at high scalability.
Another challenge faced in microbial biomass applications is that the
integration of specific bacteria has to be well-controlled in certain
microalgal production processes (Yao et al., 2019). Algal and bacterial
strain selection requirements must consider cultivation, harvesting,
processing, and conversion into renewable biofuels. Additionally, in the
biological treatment of wastewater, algae, and bacteria symbiotic rela­
tionship has shown potential in the high removal efficiency of pollutants
and oxidize them into CO2 and water. However, on the downside, the
algal-bacterial consortia frequently lead to a high concentration of algae
in the water, resulting in secondary pollution to the receiving water
bodies (van Wyk et al., 2009). Other limitations observed in algal-
bacterial wastewater treatment include the reduction of bacterial
growth due to high pH and O2 concentration from algal photosynthesis.
This is because the antibacterial substances released by algae tend to
hinder the growth of bacteria. Pathogenic bacteria can weaken the algal
cell walls, resulting in cell disruption and death (Sukla et al., 2018). If
filamentous bacteria grow rapidly at excessive concentrations, algal-
bacterial wastewater systems will face challenges like loss of nitrifica­
tion, bulking and foaming, eventually creating solid separation prob­
lems and pose major risks to the operation and environment (D’Antoni
et al., 2017). The poor removal efficiency of microalgae’s cells from
effluent and sludge disposal are also issues faced in algal-bacterial
consortia that entail appropriate measures. Thus, attention has to be
focused on space requirement, nutrients requirement, algal-bacterial
strain selection and contamination from unwanted species (Tiron
et al., 2014).
11
Biotechnology Advances 47 (2021) 107684
4.2. Competition with other sources of food additives
Algal biomass can be a sustainable source for additives in the food
industry as the extraction of biomolecules and compounds such as fatty
acids, pigments and antioxidants serve as an alternative for chemical
additives and supplements (Chacón-Lee and González-Mariño, 2010).
The open-pond cultivation system is a cost-effective way of producing
such biomass of photosynthetic organisms (Tang et al., 2020). The po­
tential nutritional or bioactive content of different algae can be used for
the production of food additives, colorants, and thickening agents like
hydrocolloids (e.g., alginates, agars, carrageenans) in various food
products namely pasta, bread, yogurt, cheese, and in beverages (Wells
et al., 2017). It was found that the optimization of algal culture medium
results in the efficient production of β-carotene, an important carotenoid
pigment due to the presence of provitamin-A, which can be applied as a
food colorant in cheese, pastry, and ice cream (Koyande et al., 2019).
Haematococcus pluvialis and Chlorella zofingiensis are also algal sources
for astaxanthin which possesses strong antioxidant capacities compared
to Vitamin C, E, or other carotenoids (Ambati et al., 2014). The appli­
cability of additives in food products is determined by techno-functional
properties such as emulsification, antioxidant activity, gelation, foam­
ing, water, and fat absorption capacities. These properties were reported
for some algae proteins and hydrolysates but many of them still remain
unclear (Caporgno and Mathys, 2018). However, despite all the poten­
tials, algal-based usage in the food industry faces challenges regarding
food safety hazards that may include allergens, toxins, pathogens, and
even heavy metal contaminants. Extracts from products consisting of
Spirulina sp., Aphanizomenon flos-aquae and Chlorella sp. appear to be
contaminated with microcystins and noludarin toxins (Heussner et al.,
2012). Moreover, another safety aspect is the exposure to biological and
non-biological contamination when algae such as Spirulina and Chlorella
are cultivated in open systems. Since algae are primary producers at the
base of the food chain, they may transfer any accumulated heavy metals
to higher levels of the trophic chain, imposing adverse effects on higher
trophic level organisms and subsequently the environment (van der
Spiegel et al., 2013). Unfortunately, the utilization of algal or algae-
derived products as food substitutes remains uncompetitive in the
market due to the lack of economic scaling for microalgae cultivation
and processing, immature technology and understudied algal-based
biotechnology (Caporgno and Mathys, 2018).
4.3. Societal acceptance of algal biotechnology for environmental
protection
The question of social acceptance of algal biotechnology in envi­
ronmental protection is a lot harder to address. A number of attributes
that are to be engineered into algae, such as high lipid content, appear
implausible to increase their competitive capacities in the natural
environment; but this reference requires further experimental exami­
nation and validation. Environmental concerns arise as there may be a
loss of algal diversity or potential dominance by a single algal species
(Shurin et al., 2016). In recent years, research groups are developing
new techniques in algal biofuel production to boost algal growth rates
and oil-extraction rates genetically and metabolically (Chen et al.,
2019). However, genetically altered algae generate dispute within sci­
ence and society over worries about the environment. A study had found
that respondents are unconvinced about the need for alteration of nat­
ural algae strains to increase productivity, with the argument that there
are unknown consequences of using genome editing (Villarreal et al.,
2020). The general public perception of algal biotechnology, especially
in Western nations, is often negative. The impression of algae can con­
jure images of green tides or harmful algae blooms (HABs) which can be
displeasing (Jacquin et al., 2014). Social perception can be seen as a
hurdle for algal development, especially in environmental protection.
Consistent explorations are essential in overcoming the challenges of
algae-based biotechnology to receive reassurance and acceptance from
J.J.J.Y. Yong et al.
the general public.
4.4. Cost feasibility and environmental impact trade-off
A trade-off must be met between cost-effectiveness and environ­
mental impact, as environmental problems would affect the cost and
industrial scale-up feasibility. Reduction of production costs depends on
the algal growth rate and its harvest efficiency for biofuel generation.
The control of contamination also affects the cost of production and
operational efficiency as algal culture renewal is needed (Ang, 2004).
The major challenge for algal-bacterial biotechnology in industrial
biofuel and biogas production is the high cost but low yield. For
instance, algal-bacterial based production of biofuel has a comparably
higher potential than terrestrial crops due to the difference in land re­
quirements, but the inability of algal technology to reach a commercial
scale remains a huge challenge. It becomes cost-intensive for large scale
operations because of the enormous investments in operating expenses,
which could lead to a four-fold increase in total wastewater treatment
costs (Lavrinovičs and Juhna, 2018). Moreover, the environmental
sustainability of algal-bacterial-based wastewater treatment technolo­
gies was questioned since it was found that algae and bacteria could
synthesize N2O, a greenhouse gas that possesses a global warming po­
tential that is 298 times higher than CO2 (Olivares et al., 2019). Algal-
bacterial consortia could increase water footprint in water-stressed
areas owing to the high evaporation rates in photobioreactors. A
calculation of an evaporation rate of up to 2.275 m3 m− 2 yr− 1 in an algae
raceway pond with a depth of 0.25 m was demonstrated in Arizona,
showing a water footprint of 33.1 m3 GJ− 1, which represents more than
600 years of rainfall for the particular geographical area to enable
production of 1 GJ m− 2 of biomass energy (Guieysse et al., 2013).
4.5. Safety and compatibility of algal-bacterial biotechnology for a safe
environment
Algae can accumulate toxins, pesticides and heavy metals, and there
could be potential contamination by pathogens especially for cultivation
in open ponds. An open-culture environment is prone to uncontrolled
growth conditions and is less protected and unsafe due to unwanted
invasion by competing bacteria, protozoa, molds, and other algae spe­
cies (Hamed, 2016). Toxic chemicals used in the harvesting process of
algae could pose serious safety problems if aluminum ions and poly­
acrylamide are accumulated in the water or food chain in aquaculture
(Han et al., 2019). Another challenge in the mutualistic relationship of
bacteria and algae is the great control of integration of bacterial com­
munity in terms of the desired composition, ways to maintain the bal­
ance in various operation modes, types of reactors, and fluctuations in
outdoor conditions (Lian et al., 2018). Nevertheless, limited information
is available regarding the uncertainties of algal-bacterial consortia in the
management of a safe environment. Further research is needed to
deepen our understanding of the complex interactions that can
contribute largely to biotechnological outcomes.
5. Sustainability and prospects
Algal-bacterial biotechnology is receiving persistent efforts on
extensive scientific research and development. Some industrialized
countries are beginning to invest heavily in algae and bacteria-based
biotechnology for environmental protection, including wastewater
treatment and reduction of CO2 emission to cope with air pollution. The
top worldwide producers of algae Dunaliella are Taiwan, Australia, Inner
Mongolia in China, and Israel, with over 80% of green algae producers.
Other algal species including Spirulina and Chlorella production are
predominant in China, India, USA, Germany, and Japan for potential
commercial exploitation of biofuels (Usher et al., 2014). Efforts were
made to establish the feasibility of reliable production of biofuels in
California, Hawaii, and New Mexico. In the next few years, the
12
Biotechnology Advances 47 (2021) 107684
consumption of coal and fossil fuels for electricity generation will
skyrocket inexorably. Algal-based technology can outstretch the useful
energy from fossil fuel combustion and at the same time minimize car­
bon emissions by recycling waste CO2 from power plants into clean-
burning and sustainable biofuels. Algae are competent in converting
the waste form of carbon (CO2) into energy-dense lipids, which are
natural oil (Sheehan et al., 1978). Theoretically, algae can utilize up to
9% of the incoming solar energy to sequester approximately 513 tons of
CO2 and produce 280 tons of dry biomass per ha− 1 yr− 1 (Bilanovic et al.,
2009). Production of alternative biofuels from algae and bacteria is
likely to lead to a more neutral CO2 emission compared to fossil fuel
sources due to the uptake of CO2 during algal growth (Usher et al.,
2014). However, cost-competitive production to an economic level re­
mains an obstacle. The European Algae Biomass Association (EABA) has
deduced that the production of algae-based biofuels and bioenergy from
laboratory level to an industrial scale could take another 10 to 15 years.
(Kovalyova, 2009).
Several studies were conducted to understand the practicability and
applicability of algal-bacterial symbiosis for the treatment of various
types of wastewaters. The symbiotic relationship between photosyn­
thetic algae and heterotrophic bacteria regarding the exchange of nu­
trients and substrates facilitates both bacterial growth and algal
photosynthesis (Choudhary et al., 2015). Algae also protect bacteria
against a harsh environment, for instance, it is found that Cladophora in
Lake Michigan serves as an environmental reservoir for Salmonella
(Byappanahalli et al., 2009). As a result, algae and bacteria consortia are
proven to enhance nutrients removal and subsequently increase biomass
productivity. The removal of nitrogen, phosphorus, and COD in syn­
thetic wastewater in a photobioreactor with algal-bacterial mixed cul­
tures was studied. It was observed that more than 90% N and P and 80%
of COD could be removed from the wastewater (Ashok et al., 2014). The
capability of amalgamation and incorporation of an algal-bacterial
symbiotic relationship into conventional activated sludge systems
makes algae-based technologies attractive, as this leads to the reduction
of aeration costs, which accounts for more than 50% of the overall en­
ergy of wastewater treatment plants (Arashiro, 2016). The feasibility of
algal-bacterial cultures in wastewater treatment and simultaneous pro­
duction of biomass was found to be comparable with the existing
microalgae cultivation (Leong et al., 2019a, 2019b). It was verified that
algal-bacterial synergy makes a significant contribution to environ­
mental management such as metal phycoremediation. In this context,
mutualistic algae and bacteria species can degrade and assimilate metals
from metal-rich wastewater. The detoxification of hazardous organic
pollutants including acetonitrile, salicylate, phenol, black oil, phenan­
threne by algal-bacterial consortia has been documented with high
removal rates (Muñoz and Guieysse, 2006). Algae- and bacteria-driven
technologies portray viable and sustainable ecological approach for
environmental protection. Algal-bacterial co-culture is, however, an
underexplored section for sustainable biotechnological application.
Careful optimization and assessment are required to elucidate the effi­
ciency of algal-bacterial consortia in environmental management in the
quest for economic feasibility and scalability to enable wider industrial
usage instead of lab-scale experiments.
6. Conclusion
In the present scenario, algal-bacterial consortia offer striking ap­
proaches and prospects in environmental management. This review
emphasizes the environmental benefits of algal-bacterial symbiosis,
ranging from phycoremediation, greenhouse gas mitigation, bloom
control to the production of high-value products, bioplastics, and
renewable biofuels as a substitute for fossil fuels. PBRs and algal wheel
technology that applies algal-bacterial symbiosis appear to be efficient
in wastewater bioremediation due to high areal productivity, low energy
requirement, and low environmental footprint. Moreover, there exists a
large potential advancement of algal-bacterial biotechnology by
J.J.J.Y. Yong et al.
integrating Industry 4.0 technologies such as smart grid and simulation
to achieve sustainable performances in the algal-bacterial co-cultivation
and reduce ecological impacts. There are, however, several hurdles to
overcome as the scale of the impacts of algal-bacterial biotechnology
stay obscure due to large gaps in literature and research. While the
potentials are wide, the achievements in operation remain modest. The
use of algal-bacterial biotechnology is still limited due to the high cost
for expansive commercial application, general societal acceptance,
algal-bacterial strains selection, and its compatibility in a safe environ­
ment. Nevertheless, the environmental stakes are attractive enough to
drive stronger holistic research and development of algal-bacterial
biotechnology to better understand their interactions and possible ap­
plications to fully utilize them in the ecosystems. Algal-bacterial
biotechnology holds both promising opportunities and challenges to
the planet for the sustainability of energy and the environment. As long
as the related technological, market and policy constraints are resolved,
the application of algal-bacterial symbiosis could guide us towards a
more sustainable and greener industry in the future decades.
Acknowledgment
This work was supported by the Fundamental Research Grant
Scheme, Malaysia [FRGS/1/ 2019/STG05/UNIM/02/2] and MyPAIR-
PHC-Hibiscus Grant [MyPAIR/1/2020/STG05/UNIM/1]. This research
was supported by Xiamen University Malaysia Research Fund, Malaysia
(Grant number: XMUMRF/2021-C7/IENG/0033). The authors also
gratefully acknowledge the financial support received from Taiwan’s
Ministry of Science and Technology under grant number MOST 109-
3116-F-006-016-CC1, 109-2621-M-029-001, and 107-2221-E-006-112-
MY3.
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