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

BioScience 60: 722–727. ISSN 0006-3568, electronic ISSN 1525-3244. © 2010 by American Institute of Biological Sciences. All rights reserved. Request
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before the net carbon balance is positive (Searchinger et al. production yields and available land area (compared with
2008). Further complicating matters is the fact that many terrestrial crops); algae’s ability to capture CO2 as bicar-
first-generation biofuel systems use food crops. Competition bonate in ponds, reducing atmospheric CO2 emissions;
for feedstocks such as corn and soybeans for food and energy and reduced competition for land, particularly arable land
production has the potential to affect global food prices. used for food production (figure 1). Algae are estimated to
Cellulosics and hemicellulosics are among the most abun- produce two- to tenfold more biomass per unit land area
dant biopolymers in nature; converting them into sugars than the best terrestrial systems (Chisti 2008, Packer 2009,
using advanced enzyme catalysts promises to grow the avail- Pienkos and Darzins 2009, Mata et al. 2010, Stephens et al.
able carbon resources for fuel production and to reduce the 2010, Weyer et al. 2010). There are several reasons for the
land area required for biofuel production (Pauly and Keeg- greater biomass yields of algae versus land plants. Generally,

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stra 2008). Hemicellulosic and cellulosic biofuel production algae have higher photosynthetic efficiency than land plants
systems enable the conversion of nearly all sugar-based plant because of greater abilities to capture light and convert it to
polymers into fuels (ethanol, butanol, diesel). Plants that usable chemical energy (Melis 2009, Weyer et al. 2010). Un-
produce high levels of cellulosic biomass, such as Miscanthus der ideal growth conditions algae direct most of their energy
and switchgrass, are being developed as second-generation into cell division (6- to 12-hour cycle), allowing for rapid
biofuel crops. These biofuel crops do not compete directly biomass accumulation. Also, unlike plants, unicellular algae
with food production, require less agronomic (fertilizer, do not partition large amounts of biomass into supportive
plowing, pesticide) inputs, and have lower environmental structures such as stems and roots that are energetically
impacts than first-generation biofuels. However, some bio- expensive to produce and often difficult to harvest and pro-
fuel crops (Miscanthus) have the potential to be invasive cess for biofuel production. In addition, algae have carbon-
species that may disrupt the biological integrity of local eco- concentrating mechanisms that suppress photorespiration
systems (Raghu et al. 2008). Overall, the energy efficiency, (Spalding 2008, Jansson and Northen 2010). With algae,
environmental sustainability, and economics of various all the biomass can be harvested at any time of the year,
renewable fuel production systems must be collectively eval- rather than seasonally. In contrast, only a portion of the
uated to make informed decisions to identify the best energy total biomass of terrestrial crops (corn cob, soybean seed) is
production systems for each location and market. harvested once a year. When algae are grown under stress-
ful conditions (e.g., low nitrogen) or in the presence of
Microalgal biofuel systems supplemental reductants (sugar, glycerol), the metabolism
Recently, there has been substantial interest and invest- of some species is redirected toward the production and
ment in the development of microalgae to produce bio- accumulation of energy-dense storage compounds such as
fuels. Advantages of microalgae-based biofuels are greater 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,
Figure 1. Microalgal carbon capture and biomass production. harvesting, and extraction systems,

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Articles

and are the subject of directed research investment from the Carbon capture by algae
public and private sectors (box 1). Various estimates indicate One advantage of aquatic carbon capture and biomass pro-
that potential oil and biomass yields from algae ponds range duction systems is the ability to capture CO2 in ponds in a
from 20,000 to 60,500 liters per hectare per year (2000 to nongaseous form as bicarbonate to fertilize algal growth. At
6000 gallons per acre per year) and 50,000 to 15,000 kilo- moderate pHs (> pH 7) and temperatures (below 30 degrees
grams per hectare per year (140 to 420 tons per acre per Celsius), the dominant form of CO2 in water is bicarbonate.
year), respectively (Weyer et al. 2010). As previously discussed, algae have active bicarbonate pumps
Optimization of light harvesting efficiency and enhanced and can concentrate bicarbonate in the cell. The bicarbo-
metabolic flux leading to increased oil or biomass accu- nate is subsequently dehydrated, either spontaneously or by
mulation promise to boost the efficiency of biomass and carbonic anhydrase, and the resulting CO2 is captured through

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oil production from algae at least two- to threefold (Wang Calvin-cycle activity, ultimately in the form of algal biomass.
et al. 2009, Stephens et al. 2010). These advancements— Between 1.6 and 2 grams of CO2 is captured for every gram of
coupled with more energy-efficient algal harvesting and oil algal biomass produced (Herzog and Golomb 2004).
extraction technologies, coproduction of income-generating A variety of industrial sources of CO2 can be harvested
commodities including methane from the anaerobic diges- using algal ponds. However, the various anthropogenic
tion of delipidated biomass, and residual biomass for animal sources of CO2 will differ in their concentrations of CO2
feeds—will collectively reduce the cost of microalgal oil and other contaminating molecules, and these attributes, as
production and potentially bring algal biofuel economics to well as gas temperature and production volume, will influ-
parity with petroleum (Stephens et al. 2010). ence the design of CO2-delivery systems for ponds. Flue
gases from fossil-fuel power plants typically have high CO2
Carbon capture by photosynthesis concentrations, ranging from 10% to 20%, but also contain
One of the more attractive features of algal biomass produc- biologically significant amounts of nitrous and sulfur oxides
tion is the potential to trap gaseous CO2 generated from (NOx and SOx). The injection of power plant flue gases into
point sources in ponds as bicarbonate. Cyanobacteria and algal ponds has been shown to elevate algal biomass yields
eukaryotic algae transport and use bicarbonate as a source of by as much as threefold, but at a high energy cost (Jeong
carbon dioxide (Spalding 2008, Jansson and Northen 2010). et al. 2003). Fortuitously, the benefits of flue gas injection
At pHs between 6.4 and 10.3, the dominant (> 50%) chemi- on algal growth can be greater than the growth impacts
cal species of CO2 in water is bicarbonate, a nongaseous solely attributed to inhibition of photorespiration by high
form of CO2. This transiently captured carbon is pumped CO2 concentrations (Douskova et al. 2009). Recent studies
into algal cells by bicarbonate transporters present in both have shown that direct injection of flue gasses into ponds
the plasma membrane and in the chloroplast envelope of increases biomass productivity by 30% compared with
eukaryotic algae (reviewed in Spalding 2008). Inside the direct injection of an equivalent concentration of pure CO2
chloroplast, bicarbonate is converted into CO2 that can be
fixed by rubisco (ribulose bisphosphate carboxylase oxy- Box 1. Constraints limiting algal biomass yield,
genase) to produce two molecules of 3-phosphoglycerate. biomass harvesting, and biofuel extraction.
Through a series of reactions these three carbon organic
acids are reduced to the sugars that are substrates for starch Algal production systems (50% to 60% costs)
and oil production. However, oxygen can compete with CO2 Identification of fastest-growing, highest-biomass-yielding
for fixation by rubisco. The products of the oxygenase reac-   strains
tion are 3-phosphoglycerate and 2-phosphoglycolate. The Enhancing photosynthetic efficiency
phosphoglycolate is subsequently metabolized to glycine, Ability to grow well across a wide range of temperature, light,
  and environments
which, when condensed with another glycine molecule to
Available genomics; transformable, stable transgene expression
produce serine, results in the loss of CO2. This carbon loss
Containment of genetically modified algae
(one carbon per two molecules of glycine) diminishes the
Increase oil accumulation with minimal biomass penalty
ability of the Calvin cycle to regenerate the five-carbon sugar
Pond environmental control and biomass optimization
substrate ribulose bisphosphate—required for CO2 fixation
Crop protection; control of competing algae, bacteria,
by rubisco—further reducing the efficiency of photosyn-   viruses, grazers
thesis. This overall process is known as photorespiration Removal of growth-inhibiting waste products
because it occurs largely in the presence of light. The process Recycle growth media and nutrients to reduce environmental
of photorespiration reduces photosynthetic carbon fixa-   impact
tion efficiency by 20% to 30% (Zhu et al. 2008). To reduce Harvesting and extraction systems (40% to 50%
the competitive inhibition of oxygen on carbon fixation by of costs)
rubisco, algae actively pump sufficient bicarbonate into cells Low energy harvesting systems
to elevate internal CO2 concentrations to levels above those Efficient oil extraction or secretion processes
achievable by equilibrium with air, and competitively inhibit Optimization of coproduct yields to offset oil production costs
photorespiration (Badger and Price 1994).

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(Douskova et al. 2009). The flue gas fertilizer effect may be impacts of fertilizing the oceans are largely unknown. An
due to the presence of supplemental nutrients (sulfur and alternative strategy to open-ocean fertilization of phyto-
nitrate) present in flue gasses. plankton is the controlled production of algal biomass in
The efficiency of CO2 capture by algae can vary accord- contained ponds or closed production systems optimized for
ing to the state of the algal physiology, pond chemistry, and biomass-accumulation and capture of carbon dioxide. Two
temperature. Carbon-dioxide capture efficiencies as high as possible algal carbon sequestration systems are considered
80% to 99% are achievable under optimal conditions and here: (1) permanent burial of total algal biomass in deep
with gas residence times as short as two seconds (Keffer and geologic formations, and (2) burial of extracted or processed
Kleinheinz 2002). For a typical 200-megawatt-hour (MWh) carbon-rich fractions from algal biomass (table 1).
natural gas–fired power plant, it has been estimated that an Burial of total algal biomass is the simplest technological

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algal pond of 3600 acres would be sufficient to capture 80% approach. Direct burial of algal biomass is the most energy-
of the plant’s CO2 emissions during daylight hours, assum- efficient way to sequester carbon, because no dewatering is
ing an algal areal biomass productivity rate of 20 grams dry required after processing. The downside is that considerable
weight per square meter per day (Herzog and Golomb 2004). amounts of inorganic nutrients would also be buried with the
Because of the greater CO2 emission levels per MWh of coal- algal biomass. As much as 7% of algal dry weight is inorganic
burning power plants, a pond approximately 7000 acres in elements or ash. Burying substantial amounts of nitrogen
size would be required to capture 80% of the CO2 emissions and phosphorous would not be sustainable. An alternative
from a 200-MWh coal-burning power plant during the day. approach to burying biomass is to bury only the neutral lipid
Locating ponds near CO2 point sources provides several or hydrocarbon fraction of the algal biomass. As previously
potential cost- and energy-saving advantages. Integrated discussed, neutral lipids or TAGs account for as much as 60%
power plant–algal pond facilities would reduce the costs of of the total dry weight of algae. More than 75% of the mass
CO2 transportation, produce limited waste heat from the of a typical TAG is carbon, and thus TAGs represent rich
power plant for warming ponds in the winter, and could give sources of captured carbon in cells. Selective extraction of
carbon credits to the utility. Locating ponds near CO2 sources TAGs under continuous-flow processes can be achieved using
can be problematic, however, depending on land availability biocompatible organic solvents that selectively extract hydro-
and the climatic suitability of the site. Furthermore, CO2 phobic molecules without killing the algae, allowing the algae
capture by biomass production is not feasible in the dark. to more rapidly generate extractable biomass without the
Thus, methods to capture, concentrate, store, and transport costs of regenerating cellular machinery (Sayre and Periera
CO2 from source to sink (pond) during the day will need to 2008). Triacylglycerols extracted using low-energy-harvesting
be developed as part of the integrated solution for capturing and extraction processes could be injected into geologic for-
as much of the CO2 emission as possible (Kadam 1997). mations to “lock” the carbon in place. Significantly, burying
TAGs does not carry the risk associated with the potential
Carbon sequestration by algae escape of gaseous CO2 from geologic formations, and because
To have the greatest impact on greenhouse gas accumula- TAGs do not contain elements other than carbon, hydrogen,
tion, CO2 must not only be captured but also sequestered and oxygen, inorganic nutrients (e.g., nitrogen, phosphorus,
over very long (geologic) time intervals. Direct injection of and sulfur) required for algal growth could be recycled and
CO2 into geological formations is one strategy under consid- not buried with the carbon.
eration for the mitigation of CO2 from point sources (Yang Another strategy to sink carbon in a geologically stable
et al. 2008). However, it is difficult to ensure that CO2 stores form is to convert the biomass into biochar (Hielmann et al.
in geologic formations will be secure over the long term. The 2010). Biochar is also a stable form of carbon that can persist
escape of large amounts of CO2 to Earth’s surface could be in soils for millions of years (Amonette et al. 2007). Biochar
catastrophic (Benson and Cook 2005). A potentially lower- is more than 90% carbon and is a by-product of pyrolysis
risk strategy for sequestering carbon dioxide is to chemically at high temperatures in the presence of catalysts under
convert CO2 into stable liquids or solids. The production of anaerobic conditions (Hielmann et al. 2010). Pyrolysis also
carbonate salts is one approach being developed (Benson generates gaseous products including hydrogen, methane,
and Cook 2005); these salts could be buried or used as con- and CO2, which may be combusted to drive the pyrolytic
struction material. reaction. Biochar may contain inorganic elements but can
Another strategy for sequestering carbon is to bury it be directly added to soil as a supplement instead of being
as biomass. This strategy is particularly attractive as solar buried, reducing energy costs and expanding the range of
energy is used to generate the biomass. One of the more possible applications. A variation of pyrolysis that generates
controversial manifestations of biomass carbon sequestra- a liquid fuel by-product is hydrothermal liquefaction. Heat
tion has been the proposal to fertilize the open oceans to treatment of biomass (at temperatures greater than 400
stimulate phytoplankton production (Stepan et al. 2002). degrees Celsius) in the presence of water leads to the pro-
Fertilizing oceans with the nutrients (e.g., iron) that usually duction of bio-oils and other products, including biochar
limit algal growth can substantially enhance algal growth (Hielmann et al. 2010). As much as 55% of the carbon in
and carbon capture (Buesseler et al. 2004), but the ecological algal biomass processed by hydrothermal liquefaction can

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Articles

Table 1. Features of various microalgal-based carbon sequestration systems.

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

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Funding was provided for this review by the US Air Force version and Management 38: S505–S510.
Office of Scientific Research and the Department of Energy’s Keffer JE, Kleinheinz GT. 2002. Use of Chlorella vulgaris for CO2 mitigation
Energy Frontier Research Center program to Richard Sayre. in a photobioreactor. Journal of Industrial Microbiology and Biotech-
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