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This review paper describes systems used to cultivate microalgae for biofuel production. It addresses general
design considerations pertaining to reactors that use natural light and photosynthetic growth mechanisms,
with an emphasis on large-scale reactors. Important design aspects include lighting, mixing, water consumption,
CO2 consumption, O2 removal, nutrient supply, temperature, and pH. Though open pond reactors are the
most affordable option, they provide insufficient control of nearly all growth conditions. In contrast, a variety
of closed reactors offer substantial control, but few feature the likelihood for levels of productivity that offset
their high cost. One of the greatest challenges of closed photobioreactor design is how to increase reactor
size in order to benefit from economy of scale and produce meaningful quantities of biofuel. This paper also
highlights the concept of combining open and closed systems and concludes with a discussion regarding a
possible optimal reactor configuration.
Figure 1. Rectangular airlift reactor with separate light collection. (Reprinted with kind permission from ref 9. Copyright 2003 John Wiley and Sons.)
example, the light gradient is primarily determined by the general.16 The optimal dark period depends on the photon flux
diameter of the tube and the biomass density in the medium.9 density of the previous light period and the fluid residence time
The biomass density affects both the light intensity and the light in zones of different irradiance.17 Formulas that describe light
penetration. Optimal cell density is specific to each strain and and dark frequency values in various types of closed reactors
needs to be maintained in order for light intensity and light can be found in the literature.9 Such formulas and other
penetration to remain at optimal levels.10 Park and Lee described quantitative models for light analysis are not included in order
an important operating parameter known as the critical cell to maintain brevity. Likewise, predictive equations that model
density, which is the maximum cell concentration without mass transfer or other design parameters are excluded here but
mutual shading in algal cultures.11 can be found in referenced papers.
The wavelength of light used to cultivate algae is also a design The combination of factors such as the length of the light
factor because some experiments have shown that cultures grow and dark cycles and the light intensity result in the overall light
differently when exposed to different colors of light. However, regime in a photobioreactor. Light regime strongly influences
optimizing this aspect in systems illuminated by natural light photoacclimation, which describes the physiological responses
is much more challenging than in systems illuminated by of cells to rapid changes in light intensity. An example of a
artificial light, where the wavelength of light shone can be common response to light intensity alteration is a change in
selected. Unfortunately, more than 50% of the incident solar chlorophyll pigment content. However, a sudden surge of light
radiation from natural light cannot be used by photosynthesis.12 can be fatal for many species of algae.18 Thus, it is important
When natural light is the growth rate limiting factor, the upper to consider light regime and photoacclimation when designing
limit in the light conversion efficiency of a large-scale culture a reactor, particularly in order to maximize the photosynthetic
may result in a maximum potential yield of 30-40 g/m2 · d.13 efficiency.
Matthijs et al. found that red light matched perfectly with the Because light is the exclusive source of energy in the
requirements of the first excited state of pigments present in photosynthetic mode of cultivation, an important calculation can
the light-harvesting antenna complexes (LHC) central to pho- be performed if solar radiation per area data is available for the
tosynthesis in green algae. He also noted that the addition of location of cultivation. On the basis of this data and other values
blue light to the red LED light did not change the growth such as photosynthetic efficiency, the maximum theoretical oil
properties.14 yield per area can be determined. The maximum theoretical oil
Light and dark cycles strongly influence the growth of algae. yield per area is especially useful for comparison between actual
In both open ponds and outdoor closed reactors, natural light is and ideal reactor performance when the oil yield and area of a
subject to changes in time of day, weather, season, and reactor in operation are known. However, reactor area does not
geography.12 Unfortunately, all reactors using natural light are directly contribute to oil yield when light is collected externally
subject to the absence of light during nighttime. According to from the site of algal cultivation, which some authors have
Chisti, biomass losses might reach as high as 25% during the advocated.9 Figure 1 depicts a rectangular reactor that collects
night, depending on the light intensity during the day, the light externally. Experiments conducted by Feuermann et al.19
temperature during the day, and the temperature at night.5 and Zijffers et al.20 suggested that natural light can efficiently
Janssen et al. noted that the length of the light/dark cycles be collected at a separate location and delivered to a photo-
experienced by algae influenced photosynthetic efficiency. bioreactor using fiber optic cables. Zijffers et al. employed the
Cycles on the order of milliseconds increased the photosynthetic light guide technology in a flat-plate reactor known as the Green
efficiency (PE) of Dunaliella tertiolecta, but cycles on the order Solar Collector.20 The schematic of this reactor is shown in
of seconds lowered the PE compared to continuous lighting.15 Figure 2. In such a design, light must enter the light guides and
The time length of the dark reactions in photosynthesis may experience total internal reflection. It must also refract out of
serve as the rate-limiting step for photosynthesis and growth in the guide when surrounded by the algal suspension. In order to
3518 Ind. Eng. Chem. Res., Vol. 49, No. 8, 2010
The time taken by the fluid to travel the length of the degasser reactor. The pH affects the liquid chemistry of polar compounds
must at least equal the time required by the oxygen bubbles to and the availability of nutrients such as iron, organic acids, and
rise out.36 If practical, the capture and sale of this oxygen even CO2.39,40 Because pH is so influential, Suh and Lee stated
stripped from the reactors may be an opportunity to reduce the that commercial pH controllers must be used in reactors to
cost of biofuel production. optimize growth.21
2.6. Nutrient Supply. In order to grow, algae require more 2.9. Other Considerations. Tredici and Zittelli asserted that
than the reactants in the photosynthesis reaction. Two major a sustainable production process, which relies on a homogeneous
nutrients are nitrogen and phosphorus, which both play a role and stable environment for microalgae cells, is more important
in controlling growth rates and lipid production. Other essential in industrial applications than high yields.7 When evaluating a
nutrients are carbon, hydrogen, oxygen, sulfur, calcium, mag- proposed reactor configuration, it is important to consider the
nesium, sodium, potassium, and chlorine. Nutrients needed in production process as a whole. For instance, a major advantage
minute quantities include iron, boron, manganese, copper, mo- of high cellular lipid content is the improved efficiency of oil
lybdenum, vanadium, cobalt, nickel, silicon, and selenium.21 extraction and other downstream processing.38 The ease of
An analysis of a common medium known as N-8 revealed integration of the reactor design with downstream processes is
the deficiency of iron, magnesium, sulfur, and nitrogen at high another key comparison parameter to reflect upon.
cell concentrations. Additional experiments showed that the Techniques rooted in biology, rather than reactor design, can
separate addition of each of the four elements did not improve have a dramatic impact on the economics of algae production.
culture performance, but that balanced supplementation resulted Chisti suggested that genetic engineering may have the greatest
in improved performance. The experimenters therefore asserted likelihood of improving the economics of biofuel production
that balancing the nutrients based on the elemental composition from microalgae.5 Genetic engineering could enhance fuel
of the biomass should be the basis for effective medium production in a variety of ways, including improving photo-
design.37 However, Chisti noted that some nutrients need to be synthetic efficiency, increasing biomass productivity, increasing
present in excess. For example, phosphorus must be supplied cellular lipid content, and improving temperature tolerance of
in excess because the phosphates react with metal ions.5 algae to reduce cooling expenses.41 In addition, genetic engi-
Applying stress in the form of limited nutrients (especially neering could increase algal cells’ tolerance to light saturation,
N or P) can increase lipid percentages within the biomass. photoinhibition, and photooxidation.5
However, this stress application also curtails the growth rate Rodolfi et al. mentioned that a strain should be highly
and thus may lower overall lipid production. The trade-off productive in outdoor culture and adaptable to the varying
between productivity and lipid content stems from the high conditions of an outdoor environment. The authors asserted that
metabolic cost of lipid biosynthesis. Rodolfi et al. described there may be many suitable strains of microalgae among the
three different situations of nutrient supply: nutrient-sufficient, thousands of natural strains available, and that for immediate
nutrient-limited, and nutrient-deficient. The first case should be purposes, there is no need to genetically modify microalgae.
evident, but the difference between the latter two cases may be Genetic engineering can improve productivity and economics,
subtle. Nutrient limitation occurs when cells are grown in an but it will require long-term research and funding, as well as
environment of a constant, but insufficient, supply of a limiting overcoming regulations against the release of genetically
nutrient, to which the cells generally adapt. Nutrient deficiency modified organisms.38
is characterized by the culture’s reliance on endogenous reserves In addition, Pulz and Gross presented several reasons to be
because there are no nutrients in the environment. Rodolfi et wary of genetic engineering. First, the authors claimed that
al. compared the growth of a few robust strains under all three increases in lipid content and other valuable cellular components
conditions, with the nutrient-deficient scenario applied to are inherently constrained by cellular metabolism. Second,
microalgae previously grown in a nutrient-sufficient environ- genetically modified algae may have a variety of detrimental
ment. The authors found that the genus Nannochloropsis was effects on the environment. Finally, Pulz and Gross argued that
an exception to the rule and had both enhanced lipid content genetically modified algae are not as fit as natural strains and
and lipid productivity in an N-deficient environment.38 thus unlikely to overcome competition without the aid of other
2.7. Temperature. Temperatures experienced by algae grown agents.42
outdoors can vary as much as the extreme outdoor temperatures Nevertheless, genetic engineering has tremendous potential
characteristic to the geographic region of cultivation. Although and has already achieved successes in the laboratory. For
algae may be able to grow at a variety of temperatures, optimal example, Mussgnug et al.6 described experiments that altered
growth is limited to a narrow range specific to each strain. For the light harvesting complexes (LHCs), which were mentioned
example, Abu-Rezq et al. found that the optimum temperature earlier in the Lighting section. The purpose of the LHCs is to
range for Nannochloropsis, Tetraselmis, and Isochrysis was capture solar energy and control the flow of the excitation energy
19-21, 19-21, and 24-26 °C, respectively.31 to the photosynthetic reaction centers. They also facilitate the
Seasonal and even daily fluctuations in temperature can dissipation of light energy as heat or fluorescence when
interfere with algae production. Temperatures can reach as high irradiation exceeds photosynthetic capacity. This second trait
as 30 °C higher than ambient temperature in a closed photo- is especially undesirable in algal bioreactors because it reduces
bioreactor without temperature control equipment.21 Evaporate efficiency. To resolve this issue, the authors used RNAi
cooling or shading techniques are employed frequently to inhibit technology to create a mutant of C. reinhardtii (referred to as
temperatures of that magnitude. In addition, a lower temperature Stm3LR3) that significantly down regulated the amount of LHCI
appears to reduce the loss of biomass due to respiration during and LHCII complexes. Their experiments, which were success-
the night.5 ful, also showed that the reduction was permanent, something
2.8. pH. Each strain of algae also has a narrow optimal range that had not previously been reported in literature. The strain
of pH. The pH of the medium is linked to the concentration of of Stm3LR3 resulted in a decrease in dissipation of captured
CO2. Suh and Lee21 mentioned that pH increases steadily in light energy, an increase in photosynthetic quantum yield, and
the medium as CO2 is consumed during flow downstream in a reduced sensitivity of the system to photoinhibition. Further-
Ind. Eng. Chem. Res., Vol. 49, No. 8, 2010 3521
Figure 11. “Inclined-tube ALR configuration: (A) schematic presentation of one airlift ’triangle’. Solid arrows indicate the direction of the gas flow, and
open arrows indicate the direction of the liquid flow (B) An array of 30 ALRs, each with a volume of 30 L, with an algal culture grown on a flue gas. Inset:
installation of the array of ALRs on the roof of the MIT Cogeneration Power Plant”.54 (Reprinted with kind permission from ref 54. Copyright 2005
American Chemical Society.)
Figure 13. “A schematic drawing of 2 units, 200 L each, connected together. (a) point of connection of two reactor units, (b) inner supports, (c) braces for
keeping together the front and back plates, (d) distance between the bottom of the reactor and the inner supports, (e) passage made between the two units
to create a ‘common volume’ between units”.55 (Reprinted with kind permission from ref 55. Copyright 2001 Elsevier.)
and Redalje,56 as well as Olaizola,57 described a two-stage best reactor type, based on photosynthetic efficiency and areal
commercial-scale production system that was in continuous productivity, appears to be column reactors, at least on the small
operation from December 1997 to September 2001. The scale used in experiments from the literature. However, technical
Aquasearch facility, located in Hawaii, was designed to constraints prevent the size of this reactor type from being
maximize the production of astaxanthin from Haematococcus increased to commercial scale without the use of multiple small
pluVialis, but the strain also produces oil under the same units, which are unlikely to be economical. Combinations of
conditions. The facility featured 25 000 L closed photobiore- open and closed reactors seem promising from a productivity
actors and 50 000 L open ponds, with a total capacity of over perspective. However, there is not enough economic information
600 000 L equally divided between photobioreactors and ponds. available to assess whether the increased productivity can offset
During the final year of operation, the average areal productivity the extra capital investment required, particularly with regard
was 10.2 g/m2 · d, which corresponded to a photosynthetic to biofuel applications. Thus, at this time, no specific reactor
efficiency of 3.0%.56 type is optimal for the commercial cultivation of microalgae
The two-stage process began with growth in an industrial- for biofuel production. This agrees with the general conclusion
scale closed reactor. The highly controlled environment in this arrived at by other authors.12,35
step maximized cell growth. Next, the algae were exposed to
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