Articles

Microalgae: The Potential for

Carbon Capture
Richard Sayre

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There is growing recognition that microalgae are among the most productive biological systems for generating biomass and capturing carbon.
Further efficiencies are gained by harvesting 100% of the biomass, much more than is possible in terrestrial biomass production systems. Micro-
algae’s ability to transport bicarbonate into cells makes them well suited to capture carbon. Carbon dioxide– or bicarbonate-capturing efficiencies
as high as 90% have been reported in open ponds. The scale of microalgal production facilities necessary to capture carbon-dioxide (CO2) emis-
sions from stationary point sources such as power stations and cement kilns is also manageable; thus, microalgae can potentially be exploited
for CO2 capture and sequestration. In this article, I discuss possible strategies using microalgae to sequester CO2 with reduced environmental
consequences.

Keywords: microalgae, carbon dioxide, biofuel, biomass, global warming

T he most abundant and sustainable source of energy 

for Earth is the sun. More than 3800 zettajoules
(1 zettajoule = 1 3 1021 joules) of solar energy are absorbed
by Earth’s atmosphere and surface annually. About 0.05%
of this energy is captured in biomass each year through
the process of photosynthesis (Hill et al. 2006, Lewis and
Nocera 2006, Zhu et al. 2008). The production, decom-
position, and accumulation of biomass play a central role
in the global carbon cycle. In a well-balanced ecosystem,
carbon capture from photosynthesis, carbon deposition in
the soil and oceans, and carbon release from biological and
geological sources are in equilibrium. Since the beginning
of the industrial age (in the 1850s), however, this equilib-
rium has been perturbed by increasing carbon emissions
from the combustion of fossil fuel and biomass, and from
reduced carbon uptake as a result of global deforestation
and the loss of arable land. Currently, more than 80% of
the energy produced globally each year is generated through
the combustion of fossil fuels. Direct carbon combustion
for energy production generates more than 24 gigatons of
carbon dioxide (CO2) annually (Lewis and Nocera 2006). As
a result, atmospheric CO2 concentrations have risen from
295 parts per million (ppm) to 380 ppm over the last 100
years, and have contributed substantially to global warming,
climate change, and resultant biological extinctions (Lewis
and Nocera 2006, Battisti and Naylor 2008).
The most environmentally sustainable way to reduce
greenhouse gas emissions associated with energy
production is to generate energy from carbon-neutral
or reduced-carbon-emission sources. Wind, geothermal,
solar, hydroelectric, ocean wave, and biofuel energy are
being developed as more sustainable alternative energy
sources when compared with the combustion of fossil fuels
(Schiermeier et al. 2008). In contrast to other renewable
energy systems, biomass products can be converted into
energy-dense, liquid-storage fuels that are compatible with the
current petroleum-based energy infrastructure. Attempting
to meet all of our energy demands using biomass-derived
fuels, however, is not sustainable using current arable land.
The total energy consumed by humans accounts for more
than 26% of the global solar energy captured in biomass
each year (Long et al. 2004, Lewis and Nocera 2006). The
limitations of available arable land and the demands for
food, fiber, and environmental sustainability constrain the
use of crops or plants for biofuel production. Since biofuels
have the potential to reduce our global carbon footprint,
however, they are an attractive part of the mix of sustainable
energy solutions. In addition, oil-based biofuels are one of
the few renewable, energy-dense fuels that can replace the
petroleum-based fuels used by the aviation and long-haul
shipping sectors.
The first generation of biofuel production systems
(starch- and sugar-based ethanol production) demonstrated
the feasibility of generating liquid transportation fuels from
renewable sources, but at initially low energy-conversion effi-
ciencies and high cost. Fermentation of biomass to produce
ethanol emits substantial carbon and yields a fuel that has only
about half the energy density of oil-based fuels. Two-thirds of
the carbon in sugars fermented for ethanol production is
emitted as CO2, and additional energy inputs associated with
the use of fertilizers, pesticides, herbicides, and soil tillage,
as well as irrigation, deforestation, increased soil respiration,
erosion, and transportation of the feedstock, all reduce the
net carbon-capture efficiency of alcohol fuels derived from
biomass (Hill et al. 2006). For corn-based ethanol biofuel
systems it takes at least 10 years of production from one site

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before the net carbon balance is positive (Searchinger et al.
2008). Further complicating matters is the fact that many
first-generation biofuel systems use food crops. Competition
for feedstocks such as corn and soybeans for food and energy
production has the potential to affect global food prices.
Cellulosics and hemicellulosics are among the most abun-
dant biopolymers in nature; converting them into sugars
using advanced enzyme catalysts promises to grow the avail-
able carbon resources for fuel production and to reduce the
land area required for biofuel production (Pauly and Keeg-
stra 2008). Hemicellulosic and cellulosic biofuel production
systems enable the conversion of nearly all sugar-based plant
polymers into fuels (ethanol, butanol, diesel). Plants that
produce high levels of cellulosic biomass, such as Miscanthus
and switchgrass, are being developed as second-generation
biofuel crops. These biofuel crops do not compete directly
with food production, require less agronomic (fertilizer,
plowing, pesticide) inputs, and have lower environmental
impacts than first-generation biofuels. However, some bio-
fuel crops (Miscanthus) have the potential to be invasive
species that may disrupt the biological integrity of local eco-
systems (Raghu et al. 2008). Overall, the energy efficiency,
environmental sustainability, and economics of various
renewable fuel production systems must be collectively eval-
uated to make informed decisions to identify the best energy
production systems for each location and market.
Microalgal biofuel systems
Recently, there has been substantial interest and invest-
ment in the development of microalgae to produce bio-
fuels. Advantages of microalgae-based biofuels are greater
Figure 1. Microalgal carbon capture and biomass production.
www.biosciencemag.org October 2010 / Vol. 60 No. 9 • BioScience 723
Articles
production yields and available land area (compared with
terrestrial crops); algae’s ability to capture CO2 as bicar-
bonate in ponds, reducing atmospheric CO2 emissions;
and reduced competition for land, particularly arable land
used for food production (figure 1). Algae are estimated to
produce two- to tenfold more biomass per unit land area
than the best terrestrial systems (Chisti 2008, Packer 2009,
Pienkos and Darzins 2009, Mata et al. 2010, Stephens et al.
2010, Weyer et al. 2010). There are several reasons for the
greater biomass yields of algae versus land plants. Generally,
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algae have higher photosynthetic efficiency than land plants
because of greater abilities to capture light and convert it to
usable chemical energy (Melis 2009, Weyer et al. 2010). Un-
der ideal growth conditions algae direct most of their energy
into cell division (6- to 12-hour cycle), allowing for rapid
biomass accumulation. Also, unlike plants, unicellular algae
do not partition large amounts of biomass into supportive
structures such as stems and roots that are energetically
expensive to produce and often difficult to harvest and pro-
cess for biofuel production. In addition, algae have carbon-
concentrating mechanisms that suppress photorespiration
(Spalding 2008, Jansson and Northen 2010). With algae,
all the biomass can be harvested at any time of the year,
rather than seasonally. In contrast, only a portion of the
total biomass of terrestrial crops (corn cob, soybean seed) is
harvested once a year. When algae are grown under stress-
ful conditions (e.g., low nitrogen) or in the presence of
supplemental reductants (sugar, glycerol), the metabolism
of some species is redirected toward the production and
accumulation of energy-dense storage compounds such as
lipids. Many unicellular algae are facultatively capable of
producing up to 60% of neutral lip-
ids (triacylglycerol [TAG]) per gram
of dry weight, making them one of
the most efficient biofuel produc-
tion systems known (Sheehan et al.
1998, Weyer et al. 2010).
Significantly, algal biofuel produc-
tion systems can be tightly controlled
and optimized. Temperature, pH, and
nutrient and CO2 concentrations can
be monitored and optimized for maxi-
mum biomass and oil yields. In addi-
tion, it may be possible to control light
quantity and quality (wavelength) by
altering pond depth or using frequency-
shifting fluorophores to increase
photosynthetically active radiation,
respectively. This level of environmen-
tal control is difficult to achieve with
land plants that have fixed plant archi-
tectures in soil open environments.
The major constraints facing bio-
fuel production from algae can be
divided into biomass production,
harvesting, and extraction systems,
Articles
and are the subject of directed research investment from the
public and private sectors (box 1). Various estimates indicate
that potential oil and biomass yields from algae ponds range
from 20,000 to 60,500 liters per hectare per year (2000 to
6000 gallons per acre per year) and 50,000 to 15,000 kilo-
grams per hectare per year (140 to 420 tons per acre per
year), respectively (Weyer et al. 2010).
Optimization of light harvesting efficiency and enhanced
metabolic flux leading to increased oil or biomass accu-
mulation promise to boost the efficiency of biomass and
oil production from algae at least two- to threefold (Wang
et al. 2009, Stephens et al. 2010). These advancements—
coupled with more energy-efficient algal harvesting and oil
extraction technologies, coproduction of income-generating
commodities including methane from the anaerobic diges-
tion of delipidated biomass, and residual biomass for animal
feeds—will collectively reduce the cost of microalgal oil
production and potentially bring algal biofuel economics to
parity with petroleum (Stephens et al. 2010).
Carbon capture by photosynthesis
One of the more attractive features of algal biomass produc-
tion is the potential to trap gaseous CO2 generated from
point sources in ponds as bicarbonate. Cyanobacteria and
eukaryotic algae transport and use bicarbonate as a source of
carbon dioxide (Spalding 2008, Jansson and Northen 2010).
At pHs between 6.4 and 10.3, the dominant (> 50%) chemi-
cal species of CO2 in water is bicarbonate, a nongaseous
form of CO2. This transiently captured carbon is pumped
into algal cells by bicarbonate transporters present in both
the plasma membrane and in the chloroplast envelope of
eukaryotic algae (reviewed in Spalding 2008). Inside the
chloroplast, bicarbonate is converted into CO2 that can be
fixed by rubisco (ribulose bisphosphate carboxylase oxy-
genase) to produce two molecules of 3-phosphoglycerate.
Through a series of reactions these three carbon organic
acids are reduced to the sugars that are substrates for starch
and oil production. However, oxygen can compete with CO2
for fixation by rubisco. The products of the oxygenase reac-
tion are 3-phosphoglycerate and 2-phosphoglycolate. The
phosphoglycolate is subsequently metabolized to glycine,
which, when condensed with another glycine molecule to
produce serine, results in the loss of CO2. This carbon loss
(one carbon per two molecules of glycine) diminishes the
ability of the Calvin cycle to regenerate the five-carbon sugar
substrate ribulose bisphosphate—required for CO2 fixation
by rubisco—further reducing the efficiency of photosyn-
thesis. This overall process is known as photorespiration
because it occurs largely in the presence of light. The process
of photorespiration reduces photosynthetic carbon fixa-
tion efficiency by 20% to 30% (Zhu et al. 2008). To reduce
the competitive inhibition of oxygen on carbon fixation by
rubisco, algae actively pump sufficient bicarbonate into cells
to elevate internal CO2 concentrations to levels above those
achievable by equilibrium with air, and competitively inhibit
photorespiration (Badger and Price 1994).
724 BioScience • October 2010 / Vol. 60 No. 9 www.biosciencemag.org
Carbon capture by algae
One advantage of aquatic carbon capture and biomass pro-
duction systems is the ability to capture CO2 in ponds in a
nongaseous form as bicarbonate to fertilize algal growth. At
moderate pHs (> pH 7) and temperatures (below 30 degrees
Celsius), the dominant form of CO2 in water is bicarbonate.
As previously discussed, algae have active bicarbonate pumps
and can concentrate bicarbonate in the cell. The bicarbo-
nate is subsequently dehydrated, either spontaneously or by
carbonic anhydrase, and the resulting CO2 is captured through
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Calvin-cycle activity, ultimately in the form of algal biomass.
Between 1.6 and 2 grams of CO2 is captured for every gram of
algal biomass produced (Herzog and Golomb 2004).
A variety of industrial sources of CO2 can be harvested
using algal ponds. However, the various anthropogenic
sources of CO2 will differ in their concentrations of CO2
and other contaminating molecules, and these attributes, as
well as gas temperature and production volume, will influ-
ence the design of CO2-delivery systems for ponds. Flue
gases from fossil-fuel power plants typically have high CO2
concentrations, ranging from 10% to 20%, but also contain
biologically significant amounts of nitrous and sulfur oxides
(NOx and SOx). The injection of power plant flue gases into
algal ponds has been shown to elevate algal biomass yields
by as much as threefold, but at a high energy cost (Jeong
et al. 2003). Fortuitously, the benefits of flue gas injection
on algal growth can be greater than the growth impacts
solely attributed to inhibition of photorespiration by high
CO2 concentrations (Douskova et al. 2009). Recent studies
have shown that direct injection of flue gasses into ponds
increases biomass productivity by 30% compared with
direct injection of an equivalent concentration of pure CO2
Box 1. Constraints limiting algal biomass yield,
biomass harvesting, and biofuel extraction.
Algal production systems (50% to 60% costs)
Identification of fastest-growing, highest-biomass-yielding
  strains
Enhancing photosynthetic efficiency
Ability to grow well across a wide range of temperature, light,
  and environments
Available genomics; transformable, stable transgene expression
Containment of genetically modified algae
Increase oil accumulation with minimal biomass penalty
Pond environmental control and biomass optimization
Crop protection; control of competing algae, bacteria,
  viruses, grazers
Removal of growth-inhibiting waste products
Recycle growth media and nutrients to reduce environmental
  impact
Harvesting and extraction systems (40% to 50%
of costs)
Low energy harvesting systems
Efficient oil extraction or secretion processes
Optimization of coproduct yields to offset oil production costs

(Douskova et al. 2009). The flue gas fertilizer effect may be
due to the presence of supplemental nutrients (sulfur and
nitrate) present in flue gasses.
The efficiency of CO2 capture by algae can vary accord-
ing to the state of the algal physiology, pond chemistry, and
temperature. Carbon-dioxide capture efficiencies as high as
80% to 99% are achievable under optimal conditions and
with gas residence times as short as two seconds (Keffer and
Kleinheinz 2002). For a typical 200-megawatt-hour (MWh)
natural gas–fired power plant, it has been estimated that an
algal pond of 3600 acres would be sufficient to capture 80%
of the plant’s CO2 emissions during daylight hours, assum-
ing an algal areal biomass productivity rate of 20 grams dry
weight per square meter per day (Herzog and Golomb 2004).
Because of the greater CO2 emission levels per MWh of coal-
burning power plants, a pond approximately 7000 acres in
size would be required to capture 80% of the CO2 emissions
from a 200-MWh coal-burning power plant during the day.
Locating ponds near CO2 point sources provides several
potential cost- and energy-saving advantages. Integrated
power plant–algal pond facilities would reduce the costs of
CO2 transportation, produce limited waste heat from the
power plant for warming ponds in the winter, and could give
carbon credits to the utility. Locating ponds near CO2 sources
can be problematic, however, depending on land availability
and the climatic suitability of the site. Furthermore, CO2
capture by biomass production is not feasible in the dark.
Thus, methods to capture, concentrate, store, and transport
CO2 from source to sink (pond) during the day will need to
be developed as part of the integrated solution for capturing
as much of the CO2 emission as possible (Kadam 1997).
Carbon sequestration by algae
To have the greatest impact on greenhouse gas accumula-
tion, CO2 must not only be captured but also sequestered
over very long (geologic) time intervals. Direct injection of
CO2 into geological formations is one strategy under consid-
eration for the mitigation of CO2 from point sources (Yang
et al. 2008). However, it is difficult to ensure that CO2 stores
in geologic formations will be secure over the long term. The
escape of large amounts of CO2 to Earth’s surface could be
catastrophic (Benson and Cook 2005). A potentially lower-
risk strategy for sequestering carbon dioxide is to chemically
convert CO2 into stable liquids or solids. The production of
carbonate salts is one approach being developed (Benson
and Cook 2005); these salts could be buried or used as con-
struction material.
Another strategy for sequestering carbon is to bury it
as biomass. This strategy is particularly attractive as solar
energy is used to generate the biomass. One of the more
controversial manifestations of biomass carbon sequestra-
tion has been the proposal to fertilize the open oceans to
stimulate phytoplankton production (Stepan et al. 2002).
Fertilizing oceans with the nutrients (e.g., iron) that usually
limit algal growth can substantially enhance algal growth
and carbon capture (Buesseler et al. 2004), but the ecological
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Articles
impacts of fertilizing the oceans are largely unknown. An
alternative strategy to open-ocean fertilization of phyto-
plankton is the controlled production of algal biomass in
contained ponds or closed production systems optimized for
biomass-accumulation and capture of carbon dioxide. Two
possible algal carbon sequestration systems are considered
here: (1) permanent burial of total algal biomass in deep
geologic formations, and (2) burial of extracted or processed
carbon-rich fractions from algal biomass (table 1).
Burial of total algal biomass is the simplest technological
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approach. Direct burial of algal biomass is the most energy-
efficient way to sequester carbon, because no dewatering is
required after processing. The downside is that considerable
amounts of inorganic nutrients would also be buried with the
algal biomass. As much as 7% of algal dry weight is inorganic
elements or ash. Burying substantial amounts of nitrogen
and phosphorous would not be sustainable. An alternative
approach to burying biomass is to bury only the neutral lipid
or hydrocarbon fraction of the algal biomass. As previously
discussed, neutral lipids or TAGs account for as much as 60%
of the total dry weight of algae. More than 75% of the mass
of a typical TAG is carbon, and thus TAGs represent rich
sources of captured carbon in cells. Selective extraction of
TAGs under continuous-flow processes can be achieved using
biocompatible organic solvents that selectively extract hydro-
phobic molecules without killing the algae, allowing the algae
to more rapidly generate extractable biomass without the
costs of regenerating cellular machinery (Sayre and Periera
2008). Triacylglycerols extracted using low-energy-harvesting
and extraction processes could be injected into geologic for-
mations to “lock” the carbon in place. Significantly, burying
TAGs does not carry the risk associated with the potential
escape of gaseous CO2 from geologic formations, and because
TAGs do not contain elements other than carbon, hydrogen,
and oxygen, inorganic nutrients (e.g., nitrogen, phosphorus,
and sulfur) required for algal growth could be recycled and
not buried with the carbon.
Another strategy to sink carbon in a geologically stable
form is to convert the biomass into biochar (Hielmann et al.
2010). Biochar is also a stable form of carbon that can persist
in soils for millions of years (Amonette et al. 2007). Biochar
is more than 90% carbon and is a by-product of pyrolysis
at high temperatures in the presence of catalysts under
anaerobic conditions (Hielmann et al. 2010). Pyrolysis also
generates gaseous products including hydrogen, methane,
and CO2, which may be combusted to drive the pyrolytic
reaction. Biochar may contain inorganic elements but can
be directly added to soil as a supplement instead of being
buried, reducing energy costs and expanding the range of
possible applications. A variation of pyrolysis that generates
a liquid fuel by-product is hydrothermal liquefaction. Heat
treatment of biomass (at temperatures greater than 400
degrees Celsius) in the presence of water leads to the pro-
duction of bio-oils and other products, including biochar
(Hielmann et al. 2010). As much as 55% of the carbon in
algal biomass processed by hydrothermal liquefaction can
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Table 1. Features of various microalgal-based carbon sequestration systems.

Algal carbon capture and
sequestration systems
Permanent burial of total fresh biomass
Permanent burial of algal lipids
Soil amendment with algal biochar
Advantages
Captures the most carbon
No biomass processing
No loss of inorganic nutrients
Long-term energy reserve
Easily handled as liquid
Potential soil supplement
Permanent carbon sequestration
  without burial
Liabilities
Burial of inorganic nutrients and water in biomass
Sequesters < 50% of carbon present in biomass
Energy costs associated with biomass processing
Sequesters less than 55% of carbon present in biomass
Energy costs associated with biomass processing
Potential dispersal of some fraction of inorganic nutrients

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be recovered as biochar, a carbon-capture efficiency greater
than that achieved by extraction of oils from algae (Hiel-
mann et al. 2010). Hydrothermal processing also has the
advantage of reduced energy consumption compared with
direct pyrolysis (Hielmann et al. 2010).
Summary
At present, there are few examples of continuous long-term
biomass production or carbon-capture systems using algae
(Jeong et al. 2003, Doucha et al. 2005). However, laboratory
and pilot plant studies suggest that capturing CO2 by algae
is a potentially viable strategy for mitigating CO2 emissions
from anthropogenic sources. The long-term prognosis for
large-scale carbon capture using algae will depend on the
passage of legislation promoting carbon-capture technolo-
gies and the economics of algal pond systems. Significantly,
life-cycle analyses suggest the costs for capturing carbon and
producing liquid biofuels from algae may approach the costs
of producing petroleum-based fuels in the next 5 to 10 years
(Schenk et al. 2008, Lardon et al. 2009, Stephens et al. 2010).
Acknowledgments
Funding was provided for this review by the US Air Force
Office of Scientific Research and the Department of Energy’s
Energy Frontier Research Center program to Richard Sayre.
Additional thanks to Kevin Berner at Phycal LLC for infor-
mation on algal biofuels life-cycle analyses.
The Danforth Center is a nonprofit research center. Sayre
is also chief technology officer of Phycal LLC in Cleveland,
Ohio. Phycal is a for-profit algal biofuels company.
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Richard Sayre (rsayre@danforthcenter.org) is director of the Enterprise Insti-
tute for Renewable Fuels, at the Donald Danforth Plant Science Center, in St.
Louis, Missouri.