Production of Biodiesel and Biogas from
Algae: A Review of Process Train Options
Patrick E. Wiley1,2*, J. Elliott Campbell1,2, Brandi McKuin1,2
ABSTRACT: Algae are an attractive biofuel feedstock because of their
fast growth rates and improved land use efficiency when compared with
terrestrial crops. Process train components needed to produce algal
biofuels include (1) cultivation, (2) harvesting, and (3) conversion into
usable fuel. This paper compares various process train options and
identifies knowledge gaps presently restricting the production of algal
biodiesel and algae-derived biogas. This analysis identified energy-
intensive processing and the inability to cultivate large quantities of
lipid-rich algal biomass as major obstacles inhibiting algal biodiesel
production. Anaerobic digestion of algal biomass requires fewer process
train components and occurs regardless of lipid content. In either scenario,
the use of wastewater effluent as a cultivation medium seems necessary to
reduce greenhouse gas emissions and maximize water use efficiency.
Furthermore, anaerobically digesting algal biomass generated from low-
technology wastewater treatment processes represents an appropriate
technology approach to algal biofuels that is poorly investigated. Coupling
these processes can improve global health by improving sanitation, while
providing a cleaner burning biogas alternative to indoor biomass cooking
systems typical of less-developed areas. Water Environ. Res., 83, 326
(2011).
KEYWORDS: algae, biofuel, biogas, biodiesel, appropriate technology,
wastewater, cultivation, anaerobic digestion.
doi:10.2175/106143010X12780288628615
Introduction
The production of renewable fuels is becoming increasingly
important as the supply of petroleum reserves diminish and
environmental consequences resulting from fossil fuel combustion
become more severe. Fuels produced from biomass have the
potential to reduce reliance on petroleum resources and reduce
greenhouse gas (GHG) emissions. However, Fargione et al. (2008)
and Searchinger et al. (2008) reported that land use practices, such
as clearing carbon-rich forests for biofuel production, might
actually increase GHG emissions when compared with emissions
released from fossil fuel combustion. Additionally, the use of
arable land for biofuel production could negatively affect the
global food supply (Johansson and Azar, 2007).
Algae have been identified as a promising renewable energy
feedstock because of their high photosynthetic efficiency and
improved land use (Mandal and Mallick, 2009; Miao and Wu,
1
School of Engineering, University of California at Merced, Merced,
California.
2
Sierra Nevada Research Institute, University of California at Merced,
Merced, California.
* School of Engineering, University of California at Merced, 5200 North
Lake Road, Merced, CA 95348; email: pwiley@ucmerced.edu.
326
2006). Lardon et al. (2009) reported that photosynthetic
efficiencies of algae range from 3 to 8%, compared with 0.5%
for many terrestrial crops. This effective conversion of solar
energy results in biomass productivities that are significantly
greater than terrestrial biofuel feedstocks (Johnson and Wen,
2009; Mandal and Mallick, 2009). In addition to rapid biomass
productivity, algae thrive in areas lacking arable land and can be
cultivated using saline, brackish, or wastewaters that have few
competing uses (Aaronson and Dubinsky, 1982; Oswald, 1995;
Pienkos and Darzins, 2009; Shen et al., 2009; Smith et al., 2010).
Considerable research has focused on producing biodiesel from
lipids that accumulate in algal cells. The U.S. Department of
Energy (USDOE) Aquatic Species Program (1978–1996) exam-
ined over 3000 strains of organisms in an effort to identify those
having the most promise as a biodiesel feedstock (Sheehan et al.,
1998). Numerous other publications have identified species of
green algae that develop proportions of lipids exceeding 40% of
their total mass (Illman et al., 2000; Mandal and Mallick, 2009;
Mata et al., 2010; Scragg et al., 2002).
Algal biomass also can be used to produce biogas through
anaerobic digestion. Anaerobic digestion is a process involving
the degradation of organic matter by bacteria in the absence of
molecular oxygen (Chen et al., 2008; Metcalf & Eddy, 2003).
Methanogenic archea present in anaerobic digesters convert algal
carbon and other organic matter into carbon dioxide (CO2),
methane gas (CH4), and organic acids (Bdour et al., 2009;
Golueke et al., 1957; Kelleher et al., 2002). Anaerobic digesters
require few energy inputs and can tolerate solids with high water
content. Converting algal biomass into energy requires the
following three distinct chronological processes:
(1) Cultivating the algae,
(2) Harvesting the biomass, and
(3) Converting the biomass into usable fuel.
While much has been published regarding these components
individually, few studies evaluate them collectively as an algal
biofuel process train. Furthermore, appropriate technology
approaches to algal biofuels suitable for less-developed areas
have received insufficient attention. This review paper identifies
and describes various technologies available for each step in the
process train and discusses the feasibility of various process
train combinations. The goal of this paper is to identify
knowledge gaps currently inhibiting algal biofuel production
on both industrial- and community-level scales. This discussion
will concentrate on end-use applications involving biodiesel
production from algal lipids and anaerobic digestion of algal
biomass.
Water Environment Research, Volume 83, Number 4

Table 1—Comparing HRP wastewater systems, raceway ponds, flat panel PBRs, and OMEGA process for the
cultivation of algal biomass.
Biomass Biomass Capital and
Cultivation concentration productivity operating
system (g/L) (g/m2?d) costs
Wastewater HRP 0.1 to 0.4 10 to 20 Low
Raceway pond 0.5 to 1 10 to 25 Low
Flat-panel PBR 1 to 2 25 to 50 High
Tubular PBR 1.5 to 2.0 25 to 50 Very high
OMEGA system 1 to 2 Undetermined Undetermined
Cultivation of Algal Biomass
The manufacturing efficiency of algae biodiesel operations is
contingent on the lipid productivity and growth rate of algae cells.
Mata et al. (2010) identified over 40 strains of microalgae capable
of accumulating lipid contents ranging from 2 to 75% by mass.
Other authors have reported lipid concentrations of algae in the
Chlorella and Scenedesmus genera in excess of 50% of the total
cell mass (Illman et al., 2000; Mandal and Mallick, 2009; Scragg
et al., 2002). However, the development of such high proportions
of lipids appears to be triggered during conditions of nutrient
limitation, particularly in circumstances of nitrate starvation (Beer
et al., 2009; Lardon et al., 2009; Sheehan et al., 1998; Woertz et
al., 2009). During these stressful periods, algae shift biosynthetic
pathways and produce lipids called triacylglycerols (TAGs),
which accumulate in the cytoplasm for the purpose of energy and
carbon storage (Beer et al., 2009; Hu et al., 2008; Sheehan et al.,
1998). However, intentionally cultivating algae in stressful
conditions inhibits cell division, which can lead to decreased
overall lipid productivity (Lardon et al., 2009; Sheehan et al.,
1998; Woertz et al., 2009).
Unlike algal biodiesel production, anaerobic digestion process-
es are capable of producing methane-rich biogas regardless of
lipid content. Sialve et al. (2009) estimated that anaerobically
digesting algal biomass containing between 2 and 22% lipid
would produce a theoretical methane yield ranging from 0.47 to
0.80 m3 CH4/kg VS. However, experimental data concerning the
anaerobic digestion of algae report actual yields ranging between
0.17 and 0.45 m3 CH4/kg VS (Sialve et al., 2009). Despite the
discrepancy between experimental data and theoretical prediction,
methane productivity from algae digestion is comparable with
experimental values reported for pig waste (0.19 m3 CH4/kg VS),
sugar beets (0.21 m3 CH4/kg VS), wastewater sludge (0.23 m3
CH4/kg VS), and clover grass (0.34 m3 CH4/kg VS) (Amon et al.,
2007; Hansen et al., 1998; Sosnowski et al., 2003). Furthermore,
the wide range of acceptable substrate constituents for anaerobic
digestion may result in greater operational flexibility when
compared with algal biodiesel production.
Numerous cultivation systems designed to generate algal
biofuel feedstocks have been proposed. This section evaluates
the performance of wastewater treatment ponds, raceway ponds,
and various photobioreactor (PBR) configurations for the
cultivation of algal biomass. A comparison of the systems can
be found in Table 1.
April 2011
Wiley et al.
Surface-to-
Evaporative volume
losses ratio References
High Low Azov and Shelef, 1982; Golueke
et al., 1957; Mara, 2008;
Oswald, 1995
High Low Chen, 1996; Lee, 2001; Pulz,
2001; Shen et al., 2009
Low High Lee, 2001; Mata et al., 2010; Shen
et al., 2009
Low Very high Lee, 2001; Mata et al., 2010; Shen
et al., 2009
Low High Trent, 2009
Wastewater Treatment Ponds. Wastewater treatment ponds
are aerobic, secondary biological treatment systems that effec-
tively reduce biochemical oxygen demand (BOD) and eliminate
enteric bacteria (Bahlaoui et al., 1997; Hosetti and Frost, 1995).
Algae are essential to the successful operation of wastewater
treatment pond systems, because they assimilate nutrients and,
through photosynthesis, produce dissolved oxygen that is
immediately available to bacteria for the oxidization of wastes
(de-Bashan et al., 2002; Oswald, 1990; Oswald et al., 1978;
Shilton et al., 2008). In contrast, electromechanical wastewater
treatment processes, such as activated sludge, require energy
intensive and highly mechanized aeration systems to maintain
aerobic conditions. Shilton et al. (2008) estimated aeration
processes to consume up to 85% of the total electrical energy
required by electromechanical wastewater treatment plants.
Therefore, wastewater ponds represent an inexpensive and
effective treatment process requiring few electrical inputs and
mechanical components. This makes wastewater ponds an
appropriate technology that is ideal for communities with limited
financial and material resources (Kivaisi, 2001).
Unicellular algae in the Chlorella and Scenedesmus genera
commonly dominate wastewater pond ecosystems, both of which
are suitable for biodiesel and biogas production. However,
wastewater environments do not present algae with nutrient-
limited conditions, preventing them from accumulating high
proportions of TAGs. Woertz et al. (2009) reported lipid
concentrations within Chlorella to between 4.9 and 11.3% when
grown in municipal wastewater, which translates into 1.9 to 4.3 MJ
of lipid energy per kilogram of biomass (assuming a lower heating
value [LHV] of 38.3 MJ/kg for algal lipids; Lardon et al., 2009).
In contrast, anaerobically digesting this material would generate
between 6.1 and 16 MJ of methane energy per kilogram of
biomass (assuming an LHV of 35.6 MJ/m3 for methane; Sialve et
al., 2009). Therefore, the lipid content of wastewater algae needs
to be somewhere between 16 and 42% to match the energy content
of biogas.
The economic feasibility of algae cultivation may be improved
by integrating biomass production and wastewater treatment.
However, the productivity of algal biomass is contingent on the
process configuration. For example, high-rate ponds (HRPs) aim
to maximize algal yields, which consequently increases aeration
and reduces detention times (Azov and Shelef, 1982; Green et al.,
1996). The HRP systems are shallow (0.5 to 1.0 m) to allow
327
Wiley et al.
adequate sunlight penetration through the water column, which is
necessary to stimulate algal productivity. In contrast, facultative
wastewater ponds typically are deeper than HRPs (1.5 to 2 m) and
consist of an anaerobic bottom layer and an oxygen-producing
photosynthetic upper portion containing algal biomass (Katseno-
vich et al., 2008; Mara, 2008). While both systems provide
adequate wastewater treatment, the decreased solar exposure
characteristic of facultative ponds necessitates longer detention
times and yields less algal biomass than HRPs. As algal
productivity forms the core of this paper, the discussion regarding
wastewater ponds will be confined to HRP systems.
Raceway Ponds. A raceway pond is a continuously operated,
closed-loop recirculation channel used to cultivate algal biomass
(Chisti, 2007). These systems are built as individual ponds or as
groups of ponds arranged in a series connection and typically are
constructed of concrete or compacted earth (Shen et al., 2009). A
paddlewheel is used to drive water around the circuit, keeping the
algae in a well-mixed suspension (Chaumont, 1993; Sheehan et
al., 1998; Shen et al., 2009). Algal species control is difficult in
raceway ponds, because they are easily contaminated with other
organisms (Mata et al., 2010; Packer, 2009). Minimizing
contamination can be achieved using highly selective conditions
that favor dominance of one particular species or cultivating algae
that spontaneously appear and dominate in ponds (Benneman et
al., 1987; Lee, 2001). Another potential drawback of raceway
ponds is evaporative losses resulting from their large surface area.
This can be especially problematic and expensive if specially
prepared media is required to cultivate algae exhibiting desirable
attributes, such as high lipid content. While these issues exist for
HRP wastewater systems, they are of less importance. This is
because the highly nutritious growth media leaves few options for
species selection and control, as Chlorella sp. and Scenedesmus
sp. likely will dominate this environment. Evaporative losses are
less concerning, because the growth media is a waste product with
limited competing uses.
Raceway ponds are advantageous, because they are relatively
inexpensive to construct and operate when compared with PBR
systems and are very easy to scale up (Lardon et al., 2009; Shen et
al., 2009). They are more productive than HRP wastewater
systems and have few electrical and mechanical components. For
these reasons, the closeout report of the USDOE Aquatic Species
Program concluded, ‘‘there is little prospect for any alternatives to
the open pond designs, given the low cost requirements associated
with fuel production’’ (Sheehan et al., 1998).
Photobioreactors. The PBR systems are used to cultivate
algae in bags, tubing, or other transparent materials that are not
exposed directly to the atmosphere (Lehr and Posten, 2009; Shen
et al., 2009). Because algae are grown in a closed system, there are
fewer concerns regarding species contamination and evaporative
losses when compared with open pond systems (Posten and
Schaub, 2009; Yeang, 2008). The concentration of algal biomass
grown in PBRs typically is greater than open pond systems,
because algae are provided maximum exposure to sunlight (Chisti,
2008a). Similar to raceway ponds, unless wastewater is used, PBR
systems will require the addition of fertilizers to prepare the
nutrient broth. Tubular PBRs commonly are used to cultivate
algae, because they have large surface-to-volume ratios and can be
arranged in a number of different configurations (Shen et al.,
2009). These options include helical tubular PBRs, fence-like
PBRs, and horizontal PBRs. Each of these systems must maintain
328
a minimum flowrate to prevent sedimentation of algal biomass
within the tubing (Chisti, 2008a). However, Chisti (2007) noted
that dense algal cultures and associated photosynthetic activity
generate dissolved oxygen levels within the system that are well
beyond saturation levels. Excessive concentrations of dissolved
oxygen will inhibit photosynthesis, leading to the photooxidative
destruction of algal cells (Chisti, 2008a). Because the dissolved
oxygen cannot be removed inside the tubes, it is recommended
that degassing vessels be used to prevent excessive accumulation
of photosynthetically generated oxygen (Chisti, 2007; Tredici et
al., 1991). Furthermore, external cooling or heat exchangers are
needed to prevent excessive temperatures within the system. Flat
panel PBRs are thin rectangular containers composed of
transparent materials (Shen et al., 2009). These units often are
configured with tilt angles necessary for maximum solar exposure
(Hu et al., 1996). This allows flat panel PBRs to have their entire
surface illuminated by direct beam radiation, whereas various
portions of horizontal reactors are shaded partially throughout the
day (Hu et al., 1996). Despite the advantage of increased sunlight
exposure, Shen et al. (2009) noted that flat panel and tubular PBRs
generally yield similar algal productivity. The National Aeronau-
tics and Space Administration (Washington, D.C.) (NASA)-
developed Offshore Membrane Enclosure for Growing Algae
(OMEGA) is a novel PBR system consisting of semi-permeable
bags designed to float in the ocean. The offshore placement of the
OMEGA system eliminates competition for terrestrial resources
and may enhance marine ecosystems by simulating a reef-like
environment. The OMEGA process cultivates freshwater algae
using wastewater effluent as the growth media (Trent, 2009). This
system is advantageous because mixing, cooling, and structural
support are provided by the ocean, rather than with expensive
mechanical components that are typical of conventional PBRs
(Trent, 2009). The osmotic gradient between the wastewater and
marine environment enables the growth media to diffuse through
the semi-permeable OMEGA material, leaving concentrated algal
biomass inside the bag (Trent, 2009). The PBR material used for
this system also is gas-permeable, eliminating the need for
degassing vessels. After an incubation period of 10 to 20 days, the
algal biomass is harvested and processed into fuel (Trent, 2009).
Comparing Wastewater High-Rate Ponds, Raceway Ponds,
and Photobioreactors. Wastewater HRPs represent an inex-
pensive, low-technology approach to algal cultivation. However,
limited solar exposure resulting from increased operating depths
results in lower cell densities than are achievable with raceway
ponds and PBR systems. Flat panel and tubular PBR systems
generate higher biomass concentrations, produce less evaporative
losses, and enable improved species control when compared with
open ponds. However, Lee (2001) cited numerous publications
reporting PBR productivities within the range achievable in open
ponds. This inhibition of algal growth may result from the
accumulation of photosynthetically generated oxygen within the
PBR system, which may be remediated by periodically pumping
the culture through degassing vessels (Chisti, 2008a; Tredici et al.,
1991). The high operating and construction cost of PBR systems
may limit their application for large-scale production of low value
products, such as biofuel feedstocks (Sheehan et al., 1998; Shen et
al., 2009). While the NASA OMEGA process requires fewer
structural and mechanical components when compared with
conventional PBR systems, its technical and economical feasibil-
ity has not yet been determined. An economic analysis by
Water Environment Research, Volume 83, Number 4

Benemann and Oswald (1994) concluded that raceways ponds are
likely the most feasible configuration of large-scale algal
cultivation.
Clarens et al. (2010) reported that the use of chemical fertilizers
for media preparation accounts for 50% of the energy and GHG
emissions associated with algae cultivation. Therefore, the use of
wastewater effluent as a culture medium could significantly
reduce the need for chemical fertilizers and reduce freshwater
inputs (Clarens et al., 2010; Levine et al., 2009). While research
has demonstrated that lipid accumulations of algae grown in
municipal wastewater generally are low, as a result of an
abundance of nitrate, cultivation of mixed native communities
may result in more robust operation, despite the potential decrease
in lipid content (Clarens et al., 2010). Rodolfi et al. (2008)
concluded that lipid accumulations resulting from nitrogen
deficiency do not increase net oil productivity, as higher oil
content is offset by lower biomass productivity during periods of
nutrient shortage.
Providing elevated concentrations of CO2 to algal cultures
increases growth rates (Packer, 2009). It has been suggested that
algal cultivation operations be constructed in close proximity to
coal-fired power plants, such that flue gas could be used as a CO2
source (Kadam, 2002). While this would decrease the transpor-
tation expenses associated with delivering CO2 to off-site
locations, there is uncertainty regarding how other flue gas
emissions will affect algal cultures (Clarens et al., 2010).
Furthermore, constructing algal cultivation systems next to power
plants may limit access to wastewater effluent, thereby increasing
fertilizer requirements for media preparation. A possible alterna-
tive would be to use waste CO2 generated during anaerobic
digestion processes at wastewater facilities. This would provide
access to wastewater effluent, while providing a ‘‘cleaner’’ source
of CO2 when compared with flue gas.
Algal Harvesting
In addition to cultivation challenges, the beneficial reuse of
algal biomass has been inhibited by the lack of efficient and
economical separation technologies (Craggs et al., 1997; Naghavi
and Malone, 1986; Poelman et al., 1997). The moisture content of
algal biomass, even when grown in PBR systems with high cell
density, is greater than 99% by mass. Harvesting algae is difficult
because of their small size, low specific gravity, and negative
surface charges, which produce stable algal suspensions through-
out the water column (Bare et al., 1975; Teixeira and Rosa, 2006).
The negative charges originate from the metabolic excretion of
algogenic organic matter and dissociation of functional groups
associated with cell membrane surfaces (Henderson et al., 2008a,
2008c). Thus, coagulating agents are required to neutralize the
surface charges on suspended particles, enabling the formation of
large agglomerates that facilitate solid–liquid separation (Ebeling
et al., 2003; Molina Grima et al., 2003; Packer, 2009; Rossini et
al., 1999; Stechemesser and Dobias, 2005; Tansel and Pascual,
2004).
Techniques for recovering algal biomass are classified as either
primary or secondary harvesting. Primary harvesting refers to the
separation of the algal cells from their growth media and can be
achieved by sedimentation or with flotation technology. The
product of primary harvesting is an algal slurry, with a total solids
content between 0.5 and 6%. Secondary harvesting systems
further thicken the slurry to a total solids content between 10 and
April 2011
Wiley et al.
20% using either a centrifuge or a belt filter press. The degree of
thickening required is determined by the intended end use of the
algal biomass. For example, primary harvesting is suitable for
anaerobic digestion, while algae biodiesel production requires
much drier algal paste. Therefore, careful evaluation of available
harvesting technologies is necessary when considering algal
biofuel processing (Table 2).
Sedimentation. Sedimentation is an inexpensive procedure
enabling the separation of flocculated algae cells from liquid using
gravitation forces. This process generally yields a wet, volumi-
nous sludge, as a result of poor compaction and slow settling
velocities (Koopman and Lincoln, 1983; Mulaku and Nyanchaga,
2004). Vessels designed for sedimentation require long detention
times and low weir overflow rates to prevent resuspension of the
algal cells (Koopman and Lincoln, 1983). The poor performance
and large surface area needed for gravity sedimentation has led
several authors to identify flotation technology as the most
efficient primary harvesting technology for algal biomass (Bunker
et al., 1995; French et al., 2000; Green et al., 1996; Teixeira and
Rosa, 2006).
Dissolved Air Flotation. Dissolved air flotation (DAF) units
consist of a compressor, saturator, and a flotation cell. Water
contained in the saturator is pressurized with the compressor,
increasing the solubility of gas, in accordance with Henry’s Law.
The water from the saturator then is released into the flotation cell
containing the coagulated algal aggregates. The sudden drop in
pressure causes small bubbles to precipitate from solution, which
then adhere to the flocculated particles, forcing them to the
surface as the bubbles rise through the water column (Al-
Shamrani et al., 2002; Lundh et al., 2000).
The algal mat accumulating at the surface of the flotation cell is
removed by a skimming mechanism, while a portion of the
subnatant, referred to as the recycle flow, is returned to the
saturator and re-pressurized (Chung et al., 2000). Green et al.
(1996) noted that DAF systems are capable of achieving algae
removal efficiencies of 99%. Several authors (Al-Shamrani et al.,
2002; Chung et al., 2000; Feris and Rubio, 1999; French et al.,
2000) have noted that bubble diameters between 10 and
100 microns are needed to optimize flotation efficiency. Smaller
bubbles are required for efficient separation, because they have
more surface area and a greater rising velocity than larger bubbles
(Al-Shamrani et al., 2002). Bubble size is a function of pressure in
the saturator, necessitating high operating pressures. Al-Shamrani
et al. (2002) noted that a minimum pressure of 390 kPa (56 psig) is
needed in the saturator to produce bubbles of optimal diameter.
While DAF units effectively harvest algae, they are typically
expensive to operate, as a result of the electrical requirements of
the saturation system (Haarhoff and Steinbach, 1997; Teixeira and
Rosa, 2006). Feris and Rubio (1999) estimated that DAF
saturation systems account for more than 50% of the total
operating cost. As a result, the economics of DAF systems may
prohibit their widespread application for harvesting algae.
Suspended air flotation (SAF) may be more appropriate than
DAF for algal harvesting, because it requires less energy and
provides an algal slurry of comparable quality.
Suspended Air Flotation. Suspended air flotation is a
process similar to DAF, in that it uses small bubbles that force
algal agglomerates to the surface of a flotation cell. The major
distinction from DAF is that SAF generates microbubbles with
surfactants instead of pressure, eliminating the need for a
329
Wiley et al.

Table 2—Comparing the performance and energy requirements of various primary and secondary harvesting
systems.

Final slurry Energy input

Process (% total solids) (kWh/m3) Loading rates Remarks References

Sedimentation 0.5 to 3% 0.1 10 to 120 minute Low weir overflow rate Divakaran and Pillai, 2002;
mixing and necessary to prevent Henderson et al., 2008b;
sedimentation resuspension of algae Koopman and Lincoln,
timea cells. 1983; Liu et al., 1999;
Shelef et al., 1984;
Uduman et al., 2010
DAF 3 to 5% 1.5 to 20 Solids loading rate Microbubbles produced in Bare et al., 1975; Feris and
1.8 to 2.5 kg/m2?h high-pressure saturator Rubio, 1999; Feris et al.,
that requires large 2000; Shelef et al., 1984;
amounts of electrical energy. Wiley et al., 2009
High removal efficiency.
b
SAF 3 to 5% 3 3 1023 Solids loading Higher loading rates and Wiley et al., 2009
rate 5.6 kg/m2?h smaller footprint than
DAF. Requires surfactants
to generate microbubbles.
EC/EF 3 to 5% 0.3 to 2.0 Undetermined Coagulating species generated Gao et al., 2010; Mollah et
in situ. Further research is al., 2004; Poelman et al.,
needed to develop this 1997; Shelef et al., 1984;
technology Uduman et al., 2010
for harvesting algae.
Belt filter press 18% 0.5 Undetermined Less effective for bacteria- Mohn, 1980; Molina Grima
sized algae like Chlorella et al., 2003; Shelef et al.,
sp. and Scenedesmus sp. 1984
Approximately 3 to 5%
total solids content feed
required.
Centrifuge 10 to 22% 0.9 to 8.0 Solids loading Approximately 0.1 to 2% Golueke and Oswald, 1965;
rate 1.1 to total solids concentration Mohn, 1980; Molina
1.8 kg/m3 feed required. Effective Grima et al., 2003;
for concentrating Schenk et al., 2008;
bacteria-sized algae. Shelef et al., 1984
a
Data obtained from jar testing. Surface and solids loading rates unavailable.
b
Energy estimates obtained from bench-scale experiment. Scalability of this result is uncertain.

compressor and high-pressure saturator (Wiley et al., 2009).
Instead, SAF units are comprised of a mixing vessel, low-pressure
recirculation pump and flotation cell. Inside the mixing vessel,
water and surfactants combine to produce microbubbles.
The water–surfactant mixture containing the microbubbles then
is released into a flotation cell. The microbubbles adhere to the
algal conglomerates, forcing them to the surface, where they are
removed with a skimming mechanism. Wiley et al. (2009)
reported that SAF had higher loading rates when compared with
DAF for the removal of algae from wastewater. This most likely is
attributed to the faster bubble rising velocity in the water column
induced by the reduced interfacial tension between the gas and
liquid phases, and electrostatic attraction between the algae and
cationic surfactant used to produce the microbubbles (Wiley et al.,
2009). The SAF system also required a much lower ratio of
flotation water to sample when compared with DAF (120:1
compared with 2:1, respectively). The higher loading rates enable
SAF units to process more sample per unit time, with significantly
less energy consumption, as a result of the elimination of the
compression step. A potential disadvantage of SAF is that it
requires more chemical inputs than DAF, which affect the cost
and life cycle GHG emissions, as a result of chemical production.
However, the electrocoagulation and electroflotation (EC/EF)
method may be more environmentally friendly than both DAF and
SAF, because it does not require chemical inputs.
Electrocoagulation and Electroflotation. Electrocoagulation
and electroflotation is a flotation technology that generates
coagulating species for destabilizing algal suspensions in situ,
through the electrochemical oxidation of consumable metal
electrodes (Chen, 2004; Jiang et al., 2002; Mollah et al., 2001,
2004; Rodriguez et al., 2007). Electrodes, typically comprised of
aluminum or iron, release charged ions into solution when direct
current is applied to the system (Holt et al., 2005; Jiang et al.,
2002; Mollah et al., 2001, 2004). The ions immediately hydrolyze
to polymeric iron or aluminum hydroxide, both of which are
excellent coagulating agents (Emamjomeh and Sivakumar, 2009;
Mollah et al., 2004). Hydrogen and oxygen bubbles are produced
during this process at the anode and cathode, respectively, which
float the sample to the surface of the flotation cell, where it is
removed with a skimming mechanism (Mollah et al., 2004).
The only operating parameter of EC/EF is current density,
which determines the volume of bubbles released into solution.
Burns et al. (1997) noted that the bubbles generated by EC/EF
ranged in size from 17 to 40 microns, which is adequate for
efficient flotation. Current density also dictates the concentration
of coagulating species released by the electrodes during

330 Water Environment Research, Volume 83, Number 4


electrochemical oxidation. The concentration is quantified using
Faraday’s Law, which is explained in eq 1.
It
nðM Þ~ ð1Þ
zF
Where
z 5 oxidation state of oxidized electrode material,
I 5 current density (A cm22),
t 5 time (seconds),
M 5 electrode material,
n 5 moles of metal dissolved into solution, and
F 5 Faraday’s constant (96 485 C/mol).
Because EC/EF generates coagulating species in situ, costs
associated with purchasing, transporting, storing, and handling
chemicals are eliminated (Abuzaid et al., 2002; Bukhari, 2008;
Mollah et al., 2001). It has few moving parts and only requires
electricity as an input (Ofir et al., 2007). This process requires
little maintenance, with the exception of periodically replacing the
sacrificial electrodes, and can be powered with renewable sources
of energy (Mollah et al., 2004; Rodriguez et al., 2007). However,
there is very limited research regarding the application of this
technology for the purpose of harvesting algae, and questions
remain regarding the most efficient electrode design and
configuration (Holt et al., 2005).
Belt Filter Press. A belt filter press is a secondary
dewatering system that uses gravity and mechanical pressure to
further dewater algal biomass (Viessman and Hammer, 2004). An
algal slurry is fed onto a moving porous belt, allowing free water
to gravity drain through the pores. The sludge then enters a low-
pressure zone, where a porous upper belt converges with the lower
one, squeezing the liquid from the solids (Metcalf & Eddy, 2003;
Stechemesser and Dobias, 2005). The belts are subjected to
increasing amounts of pressure as they pass through a series of
rollers that further dewater the sludge (Metcalf & Eddy, 2003;
Stechemesser and Dobias, 2005). Belt filter presses are able to
produce an algal paste with a total solids content of 18% and
generally require less electrical energy than centrifuges (Shelef et
al., 1984). However, they are less effective at capturing solids,
because portions of biomass are squeezed through belt pores and
lost in the filtrate. Molina Grima et al. (2003) reported that belt
filter presses are satisfactory for larger microalgae, such as
Coelastrum proboscideum and Spirulina platensis, but fail to
recover smaller algaem such as Chlorella, Scenedesmus, and
Dunaliella. For these small algae, centrifugation is the most
effective secondary harvesting method.
Centrifugation. A centrifuge is a continuously operated
separation process that uses centrifugal force to separate suspended
solids from liquids (Water Environment Federation, 1996).
Recovery of algal biomass using a decanter bowl centrifuge is a
preferred method, because it is rapid and effective for removing
bacterial sized algae (Molina Grima et al., 2003; Shelef et al., 1984).
Shelef et al. (1984) obtained an algal slurry of 22% total solids
content using a decanter bowl centrifuge. Golueke and Oswald
(1965) obtained algal slurries of Chlorella sp. and Scenedesmus sp.
ranging between 10 and 18% total solids content using this type of
centrifuge. In both cases, the total solids content of the feed was
between 0.1 and 2%. This indicates that pre-concentration using
primary harvesting procedures might be necessary for cultivation
systems that generate cultures of low cell density.
April 2011
Wiley et al.
While centrifuge systems are very reliable and have a high
percentage of solids capture, the operation and maintenance costs
are substantial (Metcalf & Eddy, 2003; Shelef et al., 1984).
Schenk et al. (2008) estimated centrifuges to consume approxi-
mately 3000 kWh/ton dry algal biomass, which may limit their
application for algae biofuel processing. However, generating
biogas from algae does not require secondary harvesting
procedures, making feedstock preparation less expensive than is
necessary for biodiesel production.
Biogas from Anaerobic Digestion
Biogas generated from anaerobic digestion is an environmen-
tally friendly and versatile fuel that releases fewer GHG
emissions, nitrous oxides, and less particulate matter when
compared with kerosene, coal, diesel, and biomass solid fuels
(Akinbami et al., 2001; Börjesson and Berglund, 2006; Demirbas
and Balat, 2006; Kapdi et al., 2005).
Biogas provides indirect benefits by reducing deforestation and
devegetation commonly practiced to collect fuel sources (Akin-
bami et al., 2001). Residual solids and digestate generated during
anaerobic digestion improve soil structure and reduce fertilizer
inputs when land-applied for agricultural operations (Akinbami et
al., 2001; Börjesson and Berglund, 2006).
The mineral composition of algae meets the nutrient require-
ments of anaerobic microflora and stimulates methanogenesis
(Sialve et al., 2009). However, the relative proportions of proteins,
carbohydrates, and lipids contained in algae cells affect the
performance of algae digestion and are variable, depending on
species and growth conditions (Illman et al., 2000; Roessler,
1990). Despite this variability, theoretical biogas yields could be
calculated if the composition of the substrate is known.
Angelidaki and Sanders (2004) provided formulas adapted from
Buswell and Neave (1930) to stoichiometrically balance the
reaction (eq 2) and estimate theoretical methane yield (eq 3).
 
a b
Cn Ha Ob z n{ { H2 O?
4 2
   
n a b n a b
z { CH4 z { z CO2 ð2Þ
2 8 4 2 8 4
 
n a b :
z { 22:4
2 8 4
B0 ~ ð3Þ
12nzaz16b
Where
B0 5 theoretical methane yield (L CH4/g VS), and
22.4 5 molar volume of methane, STP.
The theoretical CH4 yields from substrates that are typical of
anaerobic digesters can be found in Table 3 (Angelidaki and
Sanders, 2004; Sialve et al., 2009). However, observed biogas
production from algal biomass may be less than the theoretical
predictions. This is because a fraction of the substrate is used for
cell maintenance and anabolism and because resistant algae cell
walls may limit accessibility to intracellular components (Angel-
idaki and Sanders, 2004; Sialve et al., 2009).
Golueke et al. (1957) compared the digestibility of algal sludge
consisting of Chlorella sp. and Scenedesmus sp. to raw wastewater
sludge. The results from this experiment indicated that biogas
production per kilogram of volatile solids (VS) destroyed was
331
Wiley et al.
Table 3—Theoretical methane yields for substrates
comprising the majority of algal cellular constituents
(adapted from Sialve et al., 2009; Angelidaki and
Sanders, 2004).
Substrate Composition CH4 yield (L/g VS)
Lipids C57H104O6 1.014
Proteins C6H13.1O1N0.6 0.496
Carbohydrates (C6H10O5)n 0.415
comparable for both treatments (1.02 m3/kg VS for wastewater
sludge versus 0.986 m3/kg VS algal sludge), but volatile solids
destruction rates were higher for wastewater sludge (60% versus
44%) when operated at mesophilic conditions. The volatile solids
destruction rate of the algal sludge improved to 54% when the
temperature was increased from 35 to 50uC, indicating that
thermophilic operation reduces the ability of algal cells to resist
bacterial attack (Golueke at al., 1957). The composition of gas
produced during these experiments was 61 to 63% CH4, 31 to 32%
CO2, and 5 to 8% other (Golueke et al., 1957).
In addition to resistance to biodegradation, algal biomass has a
low carbon-to-nitrogen ratio (C:N), which can reduce methane
production (Yen and Brune, 2007). Thus, co-digestion of algal
biomass with high-carbon content materials may improve digester
performance. Yen and Brune (2007) doubled the methane
production of algal sludge comprised mostly of Chlorella sp.
and Scenedesmus sp. through co-digestion with waste paper. This
study estimated algal biomass alone to have a C:N of 6:1, which
was adjusted to a range of 20 to 25:1 with waste paper addition
(Yen and Brune, 2007).
Biogas generated from anaerobic digestion is a cleaner burning
alternative to wood and other biomass fuels commonly used for
indoor cooking in rural areas (Figure 1) (Demirbas and Balat,
2006; Eaton, 2009). Many biomass-cooking systems lack working
chimneys, resulting in poor indoor air quality (Fullerton et al.,
2008; Masera et al., 2005; Smith, 2006). These poorly ventilated
Figure 1—(a) Star-burner burning biogas and (b) smoke from a woodfuel cooking system in Chiapas, Mexico.
Prolonged exposure to indoor air pollution from cooking can result in negative health effects (photos courtesy of
Alexander Eaton).
332
indoor environments lead to increased incidents of respiratory
infections and eye diseases, particularly in women and children, as
these groups most often are present during cooking (Kapdi et al.,
2005; Smith, 2006). While biogas can provide improved indoor
environmental quality and reduce disease, technical issues, such as
gas storage and distribution, may be a potential barrier to adoption
(Akinbami et al., 2001). A possible solution would be to compress
biogas into cylinders, thereby making it transportable without a
distribution network. However, Kapdi et al. (2005) noted that
incombustible biogas constituents (CO2, H2S, and water vapor)
reduce the heating value and must be removed using scrubber
systems to make biogas compression economical. After scrubbing,
the biogas becomes enriched in methane and can be a direct
substitute for natural gas, making it suitable as vehicle fuel or for
electricity generation.
Sialve et al. (2009) determined that direct anaerobic digestion
of algal biomass might be more energetically favorable than
anaerobically digesting cellular residues following lipid extrac-
tion. Other recent work suggests that bioenergy pathways resulting
in biogas may have greater GHG and energy security advantages
than pathways resulting in liquid fuels (Campbell et al., 2009).
These results indicate that biogas production may be the most
practical process for converting algae into energy. However,
Börjesson and Berglund (2006) noted that energy inputs and
environmental effects vary greatly among biogas production
systems. Therefore, a life-cycle assessment (LCA) of the entire
algal biogas production chain is needed to fully assess the
environmental effects.
Biodiesel from Algae
Biodiesel is a non-toxic and biodegradable alternative fuel
derived from renewable sources (Hossain et al., 2008). Biodiesel
provides comparable engine performance to petroleum diesel fuel,
while reducing sulfur and particulate matter emissions (Miao and
Wu, 2006; Scragg et al., 2002). During the manufacturing process,
TAGs are transesterified with an acid or alkali catalyst to produce
biodiesel and glycerol (Chisti, 2007; Johnson and Wen, 2009; Van
Water Environment Research, Volume 83, Number 4

Gerpen, 2005). While biodiesel is an acceptable substitute for
petroleum diesel, it is unlikely that its production using terrestrial
biomass feedstocks will displace annual demand for petroleum
diesel. Chisti (2007) noted that 326% of available cropping area in
the United States would be required for soy-based biodiesel to
meet 50% of the annual demand for transportation fuel. In
contrast, it has been estimated that algal biodiesel could satisfy
this demand with reduced land use effects (Chisti, 2007). Despite
this potential, there presently are no large-scale operations
producing algal-derived biodiesel (Lardon et al., 2009).
Biodiesel can be produced through direct transesterification of
algal biomass or by a two-step process by which lipids are
extracted, collected, and transesterified (Johnson and Wen, 2009).
Either process requires lipid extraction using combinations of
solvents and alcohols, such as chloroform/methanol, hexane/
ispropanol, or petroleum ethers and methanol (Johnson and Wen,
2009; Mulbry et al., 2009). The direct method is advantageous,
because it combines lipid extraction and transesterification into
one process, making it less time-consuming than extraction-
transesterification processes (Johnson and Wen, 2009). However,
Johnson and Wen (2009) discovered that biodiesel yield from wet
biomass processed using direct transesterification was significant-
ly less that that obtained from dry biomass, indicating that
biomass drying is required for this process. In contrast, the two-
step extraction transesterification process yielded similar results
for both wet and dry biomass (Johnson and Wen, 2009).
Lardon et al. (2009) conducted an LCA to estimate which algal
biodiesel process options would yield the most favorable energy
balance. This study compared algae grown in normal conditions
(sufficient nitrogen) and in low nitrogen conditions, assuming a
lipid content of 12 and 25%, respectively. Extraction procedures
using both wet (20% total solids) and dry (,90% total solids)
biomass also were compared. Results from this assessment
concluded that the wet lipid extraction of algae grown under
normal conditions provided the most favorable total energy
balance, followed by the wet extraction of lipid from the low
nitrogen media. Scenarios requiring 90% total solids content were
less encouraging, as a result of the additional energy required for
drying the biomass (Lardon et al., 2009). Biodiesel production from
dry biomass may become more feasible if solar drying processes are
developed. However, the practical feasibility and lipid stability
during solar drying processes are uncertain (Lardon et al., 2009).
Cultivation systems producing large quantities of lipid-rich
algal biomass and the development of technologies that reduce the
energy inputs required for harvesting and biodiesel production are
needed to improve the economic viability of algal biodiesel
(Lardon et al., 2009; Levine et al., 2009; Sialve et al., 2009). A
promising strategy to improve net lipid production is to initially
cultivate algae in nutrient-rich conditions to develop biomass, then
‘‘starve’’ the cells of nitrate to stimulate lipid productivity. Hu et
al. (2008) also noted the promise of genetic engineering to
enhance the lipid productivity of algal cells. However, previous
attempts to identify and isolate mechanisms initiating the ‘‘lipid
trigger’’ have been unsuccessful (Sheehan et al., 1998). Thus,
more research focusing on the biosynthesis of algal lipids is
needed for genetic manipulation to improve to feasibility of
biodiesel production (Hu et al., 2008).
Alternatives to centrifugation still are needed to produce an
algal paste of sufficient dryness to enable lipid extraction and
transesterification. Chisti (2008b) suggested that sedimentation
April 2011
Wiley et al.
followed by vacuum belt filters might be an adequate substitute
for centrifugation. However, previous research has cited the poor
reliability of sedimentation, and further investigation is needed to
assess the performance of vacuum filtration for concentrating
bacterial-sized algae (Koopman and Lincoln, 1983; Mulaku and
Nyanchaga, 2004; Shelef et al., 1984).
Energy-efficient and inexpensive lipid extraction and transes-
terification processes also are needed for algal biodiesel to be
cost-competitive with petroleum fuels. Several studies have
suggested that biogas generated from anaerobically digested algal
cells following lipid extraction can be used to offset energy
demands from algal biodiesel production processes (Chisti, 2007,
2008a, 2008b; Sialve et al., 2009). However, the effects that
extraction solvents will have on the digester microbial community
are unknown. Toxicity issues might be avoided using environ-
mentally friendly, solvent-free algal biodiesel processes suggested
by Amin et al. (2009) and Xiong et al. (2009). These techniques
use osmotic shock to rupture algal cells, enabling oil fractioniza-
tion, while enzymes from Candida sp. are used for transesterifi-
cation.
Process Train Combinations
Wastewater treatment ponds generally have decreased algal
culture densities compared with raceway ponds and PBR systems.
This makes wastewater ponds an unlikely algal biodiesel
feedstock, as both primary and secondary harvesting procedures
would be necessary. However, coupling wastewater treatment
ponds with anaerobic digestion could provide adequate wastewa-
ter treatment, while simultaneously producing biogas for cooking,
heating, lighting, and power generation. These low-technology
systems are capable of meeting effluent standards established in
the United States, making them an inexpensive alternative to
electromechanical processes. These systems also are appropriate
for less developed areas, where an estimated 2 billion people lack
access to adequate sanitation, and indoor air quality is poor
because of biomass cooking systems (Mara, 2001; Muga and
Mihelcic, 2008). Additionally, stabilized algal sludge resulting
from anaerobic digestion represents a potential source of
recovered nitrogen and phosphorus fertilizer (Mulbry et al.,
2005; Yen and Brune, 2007).
The missing link connecting these processes is a primary
harvesting system matching the resource availability of rural
areas. Sedimentation settling rates are too slow and yield a sludge
that is too dilute for anaerobic digestion. The DAF systems are
highly mechanized, require chemical coagulants, and consume
large amounts of electricity. The SAF units are less mechanized
and require fewer electrical inputs, but are more chemically
intensive. The EC/EF process could bridge wastewater treatment
ponds and anaerobic digestion within an appropriate technology
framework. It generates coagulating species in situ through the
electrochemical oxidation of metallic electrodes and requires only
electricity as an input, which could be provided by photovoltaic
modules. There are no moving parts, and the only routine
maintenance would be periodically replacing the electrodes.
Anaerobic digestion of algal biomass from raceway ponds and
PBR systems would have similar process train components as
wastewater systems (Figure 2). Industrialized areas could ‘‘clean’’
biogas with scrubbing technologies and generate a source of
bioelectricity to power battery electric vehicles. Campbell et al.
(2009) suggested that bioelectricity pathways deliver more
333
Wiley et al.

Figure 2—Process train options for anaerobically digesting algal biomass cultivated in wastewater systems,
raceways ponds, and PBRs. Coupling anaerobic digestion processes and wastewater treatment is an appropriate
technology approach to algal biofuels that can address public health issues in developing areas. Industrialized areas
could generate bioelectricity from algal-derived biogas for transportation and other needs.

transportation and GHG offsets than liquid biofuels. This pathway
may be more energetically favorable than algal biodiesel, as a
result of the decreased processing necessary to produce biogas
(Sialve et al., 2009).
Cultivating algae in raceway ponds using wastewater effluent is
less expensive and produces fewer GHG emissions than PBR
systems (Benemann and Oswald, 1994; Clarens et al., 2010;
Lardon et al., 2009; Sheehan et al., 1998). Wastewater effluent
typically is nutrient-rich and does not present conditions of nitrate
limitation that facilitate lipid accumulation. However, it has been
observed that higher lipid concentrations are offset by lower
biomass productivity during periods of nutrient shortage,
indicating that net oil production may be greater in nutrient-rich
conditions (Clarens et al., 2010; Rodolfi et al., 2008). The total
solids content of algal biomass achievable in raceway ponds and
PBR systems approaches that which is acceptable for direct feed
into a centrifuge, eliminating the need for primary harvesting.
Centrifugation appears to be the most effective concentration
method, because it has a higher percent solids capture than a belt
filter press (Molina Grima et al., 2003).
Producing biodiesel from algal lipids can occur via direct
transesterification or by a two-step extraction-transesterification
procedure. Direct transesterification is less time-consuming, but
necessitates biomass drying, which increases energy needs and
GHG emissions. In contrast, the two-step process can be
performed using wet biomass, thereby eliminating the need to
dry the algal slurry (Johnson and Wen, 2009). The use of solvent-
free extraction and transesterification processes might reduce
costs and GHG emissions associated with algal biodiesel
production. Furthermore, energy demands associated with biodie-
sel production may be offset by anaerobically digesting the algal
cell following extraction and transesterification, provided the
residuals are non-toxic (Figure 3).
Conclusion
The lower density of algal biomass present in wastewater ponds
systems suggests that anaerobic digestion is the most appropriate
energy pathway. As these systems produce less biomass than
raceway ponds and PBR systems, it is unlikely that wastewater
ponds will be used for large-scale algae cultivation. Instead, these

Figure 3—Pathways for producing biodiesel from algal biomass. The increased cell densities achievable in raceway
ponds and PBRs may eliminate the need for primary harvesting. Following transesterification processes, cellular
residues can be digested anaerobically to generate electrical power.

334 Water Environment Research, Volume 83, Number 4


systems represent a low-technology, community-scale approach to
algal biofuels. Raceway ponds appear more cost-effective than PBR
systems for industrial-scale algae cultivation. However, the NASA
OMEGA process may be less expensive than traditional PBR
systems, as a result of fewer infrastructure needs for mixing,
cooling, and degassing. The use of wastewater effluent as a
cultivation broth in mass cultivation operations is advantageous,
because it releases fewer GHG emissions than synthetically
prepared media. Anaerobic digestion of mass-produced algae
requires less processing than algal biodiesel and could be a more
energetically favorable pathway. Large-scale production of algal
biodiesel remains energy-intensive and expensive, as a result of
harvesting procedures and lipid extraction/transesterification pro-
cedures. However, these energy demands may be partially offset, if
electricity is produced by anaerobic digestion of algal waste
residuals following biodiesel processing. The CO2 removed during
biogas scrubbing can be injected to the cultivation broth to
stimulate algal growth. Both industrial-scale and appropriate
technology approaches to algal biofuels face technical obstacles
throughout the process train. Based on the assessment, the
following knowledge gaps and research needs have been identified:
(1) Presently, lipid accumulation and biomass productivity appear
to be mutually exclusive. Development of cultivation systems
capable of generating large quantities of lipid-rich algal
biomass is necessary to improve the economic viability of
algal biodiesel.
(2) If genetic engineering is used to enhance lipid productivity,
concentrated research on the biosynthesis of algal lipids is
needed to effectively manipulate these mechanisms.
(3) Anaerobic digestion of algal biomass generated during
wastewater treatment processes represents an appropriate
technology approach to algal biofuels that can improve global
health issues stemming from poor sanitation and indoor
cooking systems. An appropriate technology algal harvesting
must be developed to bridge these technologies.
(4) Both primary and secondary harvesting procedures are
energy-intensive and expensive. Improving this aspect of
algal processing will reduce costs associated with biogas and
biodiesel production.
(5) Detailed LCAs for algal biogas and biodiesel production are
needed to determine which biofuel pathway is most favorable,
with respect to GHG emissions and net energy yields.
(6) Development of more efficient and environmentally friendly
lipid extraction/transesterification processes are needed to
improve the sustainability of algal biodiesel.
Credits
The authors thank Alexander Eaton (International Renewable
Resources Institute, Mexico City, Mexico) for permitting the use
of his photographs in this manuscript.
Submitted for publication June 21, 2010; revised manuscript
submitted September 9, 2010; accepted for publication September
13, 2010.
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