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Microalgae are eukaryotic cellular organisms having a cell wall, plasma membrane, cytoplasm, nucleus, organelles, and

plastids. Th e size of microalgae cells ranges from a few μm to ca. 200 μm. Th ey contain chlorophyllic apparatus where
photosynthesis takes place. Together they produce close to 50% of all the photosynthetically produced O2 on Earth. The
biodiversity of microalgal species is enormous. It has been estimated that only 4–5 × 104 species out of the available 2–8
× 105 are described in the literature, belonging to Chlorophyceae (green algae), Cyanophyceae (blue-green algae),
Chrysophyceae (golden algae), and Bacillariophyceae (diatoms).3 Th e microalgae phylogeny is still being debated.

Algae generally require less land and fresh water than terrestrial plants but to harvest the maximum amount of light and
CO2 into the desired metabolite(s) is no mean task due to technological challenges that exist when growing microalgae
on a large scale. Th e potential of microalgae is strong and recent research in this fi eld has brought new insights to light,
showing their importance for the bio-based industry. Second-generation biodiesel is produced from fatty-acid waste
streams, such as cooking oil. Second-generation bioethanol is produced from lignocellulosic materials proceeding from
wood (residues), energy crops, agricultural waste, and agro-industrial residues. In these aspects microalgae are
essentially diff erent; they could be cultivated in the vicinity of already existing industrial sites as they do not depend on
(fertile) land availability.

Cultivation systems

Phototrophic algae absorb sunlight and assimilate nutrients from natural habitats (including CO2 from the atmoshpere).
Various microalgae cultivation techniques have been studied in artifi cial habitats to overcome the limitations under
natural growth conditions. Fluorescent lamps are changed for LED lamps as artifi cial light sources.5,6 Others used CO2-
containing waste streams, as the CO2 concentration in the Earth’s atmosphere is relatively very low.7 Th ree distinct
algae production systems exist: photo-autotrophic, heterotrophic, and mixotrophic.

The algae species in the former use light as energy source and mainly inorganic substrates as a carbon source.
Heterotrophic species use organic substrates both as energy and carbon sources, while the mixotrophic species use
light, CO2, and organic substrates. In contrast to microalgae, macroalgae (seaweed) have different cultivation needs that
typically require open off -shore or coastal facilities.

Considering the costs for nutrients and artificial light, photo-autotrophic production is the most viable option for the
production of algae biomass on a large scale.

Closed systems permit the cultivation of specific species for an extended duration with minimal risk of contamination.

Heterotrophic cultivation allows faster biomass production without a light source where an organic carbon substrate is
used as both energy and carbon source.

Th e main advantages of heterotrophic cultivation are a high degree of growth control, higher cell density, and lower
harvesting costs.

Dark conditions in heterotrophic cultures inhibit pigmentation and secondary metabolite production.

Mixotrophic cultivation techniques employ a combination of autotrophic and heterotrophic production. This implies that
microalgae can ingest both organic compounds and CO2 as a carbon source; the CO2 released during respiration is used
during photosynthesis. Th is means that in the dark phase less photosynthesis-derived biomass loss occurs when an
organic carbon donor is present. Typical mixotrophic metabolisms can be found in Spirulina platensis (cyanobacteria)
and in Chlamydomonas reinhardtii (green algae).8 Spirulina sp. cultures have proven to grow faster in mixotrophic
conditions, as the photo-inhibition period is shortened drastically.

Light conditioning and harvesting

Th e 400–700 nm spectral range of solar radiation is suitable for microalgae to perform photosynthesis. 63 Photons with
shorter and longer wavelengths are not favorable for the survival of microalgae, as they can damage the cells. Algae
contain pigments that absorb light in this wavelength range to make sure that not too much of this light spectrum
penetrates the cell avoiding damage.

in high cell-density cultures, cells nearer to the light source tend to absorb all the incoming light (self-shading),
preventing it from reaching distant cells.

A robust, long-lasting, effi cient and inexpensive light source is a prerequisite to commercially exploit the potential of
microalgae, such as natural sunlight. However, outdoor sunlight has some drawbacks like day-night cycle and weather
changes (fl uctuations in irradiance).

pH

In particular, the pH regulates enzymatic activity, phosphorus availability, ammonia toxicity, and the availability of
inorganic carbon. During algae growth in a closed system, the pH tends to increase to pH 10 as dissolved CO2 is
assimilated in the algae. Some studies mentioned that high pH stress inhibits the cell cycle and triggers the lipid
accumulation, which in turn enhances the lipid quality. the need for CO2 storage seems unavoidable to control the pH
but also to maintain the overall carbon balance in the sequestration.

Salinity

Th e salinity tolerance is an important aspect to reduce fresh wateruse. Media that are too saline imply osmotic stress,
which is not benefi cial to algae growth and can alter the cell metabolism, for example secretion of osmo-protective
solutes in sacrifi ce of the lipid storage.

Temperature

Higher temperatures enhance the CO2 absorption and fixation, but too high temperatures inhibit the respiration
metabolism. It is important to maintain an optimum temp for microalgal growth, mostly ranging between 18–30 °C.

Nutrient feed

The lipid composition of microalgae oils varies with the nutrient composition. According to Devi et al.(2012) mixotrophic
cultivation off ers a higher growth rate; however, higher lipid productivities were observed in nitrogen disinherited
autotrophic mode.107 Nitrogen (NH4+, NO3− and NO2−) deprivation enhances lipid production but inhibits cell growth
and propagation, as it is the main source of proteins and nucleic acids.108 Thus, in view of the production of algae
derived biodiesel, the challenge during cultivation is to combine high productivity (optimal nutrient supply) and
maximum lipid content (growth limitation). Improving the nitrogen fixation metabolism of microalgae can reduce the
need for nitrogen nutrients.4 Interestingly, in the culture of Isochrysis galbana urea was the prefered nitrogen source for
growth and lipid storage.109 Phosphorus plays a crucial role in the production of phospholipids (cell wall) and in the
intracellular energy transfer. Other micronutrients, such as vitamins, silicon and metal (complexes), are also vital for
growth.