Principles for attaining maximal microalgal productivity in photobioreactors: an overview

Hydrobiologia - Efficient management of mass algal cultures requires appreciation of the most important factors governing the light regime of the average cell, i.e. the interrelationships between...     

Closed photobioreactors for production of microalgal biomasses

Microalgal biomasses have been produced industrially for a long history for application in a variety of different fields. Most recently, microalgae are established as the most promising species for biofuel production and CO(2) bio-sequestration owing to their high photosynthesis efficiency. Nevertheless, design of photobioreactors that maximize solar energy capture and conversion has been one of the major challenges in commercial microalga biomass production. In this review, we systematically survey the recent developments in this field.     

Irradiance optimization of outdoor microalgal cultures using solar tracked photobioreactors

Photosynthetic activity and temperature regulation of microalgal cultures (Chlorella vulgaris and Scenedesmus obliquus) under different irradiances controlled by a solar tracker and different cell densities were studied in outdoor flat panel photobioreactors. An automated process control unit regulated light and temperature as well as pH value and nutrient concentration in the culture medium. CO₂ was supplied using flue gas from an attached combined block heat and power station. Photosynthetic activity was determined by pulse amplitude modulation fluorometry. Compared to the horizontal irradiance of 55 mol photons m^?2 d^?1 on a clear day, the solar tracked photobioreactors enabled a decrease and increase in the overall light absorption from 19 mol photons m^?2 d^?1 (by rotation out of direct irradiance) to 79 mol photons m^?2 d^?1 (following the position of the sun). At biomass concentrations below 1.1 g cell dry weight (CDW) L^?1, photoinhibition of about 35 % occurred at irradiances of ≥1,000 μmol photons m^?2 s^?1 photosynthetic active radiation (PAR). Using solar tracked photobioreactors, photoinhibition can be reduced and at optimum biomass concentration (≥2.3 g CDW L^?1), the culture was irradiated up to 2,000 μmol photons m^?2 s^?1 to overcome light limitation with biomass yields of 0.7 g CDW mol photons^?1 and high photosynthetic activities indicated by an effective quantum yield of 0.68 and a maximum quantum yield of 0.80 (F _v/F _m). Overheating due to high irradiance was avoided by turning the PBR out of the sun or using a cooling system, which maintained the temperature close to the species-specific temperature optima.     

Batch and semi-continuous microalgal TAG production in lab-scale and outdoor photobioreactors

Microalgal triglycerides (TAGs) represent a sustainable feedstock for food, chemical and biofuel industries. The operational strategy (batch, semi-continuous, continuous cultivations) has an impact on the TAG productivity. In this study, semi-continuous (i.e. with fixed harvesting frequency) and batch cultivations were compared on TAG production both at lab-scale and in outdoor cultivations. At lab-scale, the semi-continuous TAG productivity was highest for a cycle time of 2 days (SC1; 0.21 g L^?1 day^?1) and similar to the maximum obtained with the batch (optimal harvest time; 0.23 g L^?1 day^?1). Although TAG content was lower for SC1 (22 %) than for the batch (35 %), higher biomass productivities were obtained with SC1. Outdoors, semi-continuous cultivations were subjected to a lower degree of stress (i.e. higher amount of nitrogen present in the system relative to the given irradiance) compared to lab-scale. This yielded low and similar TAG contents (10–13 %) in the different semi-continuous runs that were outdone by the batch on both TAG content (15–25 %) and productivity (batch, 0.97–2.46 g m^?2 day^?1; semi-continuous, 0.35–0.85 g m^?2 day^?1). The lab-scale experiments showed that semi-continuous strategies, besides leading to similar TAG productivities compared to the batch, could make TAG production cost effective by valorising also non-TAG compounds. However, optimization of outdoor semi-continuous cultivations is still required. For instance, the nitrogen supply and the harvest frequency should be adjusted on the total irradiance. Additionally, future research should focus on recovery metabolism upon nitrogen resupply.     

Carbon dioxide uptake efficiency by outdoor microalgal cultures in tubular airlift photobioreactors

The influence of solar irradiance and carbon dioxide molar fraction of injected CO(2)-air mixtures on the behavior of outdoor continuous cultures of the microalga Phaeodactylum tricornutum in tubular airlift photobioreactors was analyzed. Instantaneous solar irradiance, pH, dissolved oxygen, temperature, biomass concentration, and the mass flow rates of both the inlet and outlet oxygen and carbon with both the liquid and gas phases were measured. In addition, elemental analysis of the biomass and the cell-free culture medium was performed. The oxygen production rate and carbon dioxide consumption rate increased hyperbolically with the incident solar irradiance on the reactor surface. Carbon losses showed a negative correlation with the daily variation of the carbon dioxide consumption rate. The maximum CO(2) uptake efficiency was 63% of the CO(2) supplied when the CO(2) concentration in the gas supplied was 60% v/v. Carbon losses were >100% during the night, due to CO(2) production by respiration, and hyperbolically decreased to values of 10% to 20% in the midday hours. An increase in the carbon fixed in the biomass with the solar cycle was observed. A slight daily decrease of carbon content of the cell-free culture medium indicated the existence of carbon accumulation in the culture. A decrease in CO(2) molar fraction in the injected gas had a double benefit: first, the biomass productivity of the system was enhanced from 2.05 to 2.47 g L(-1) day(-1) by reduction of CO(2) inhibition and/or pH gradients; and second, the carbon losses during the daylight period were reduced by 60%. The fluid dynamics in the reactor also influenced the carbon losses: the higher the liquid flow rate the higher the carbon losses. By using a previous mass transfer model the experimental results were simulated and the usefulness of this method in the evaluation and scale-up of tubular photobioreactors was established.     

Physiology and nutritional aspects of actinomycetes: an overview

A review is presented of the nutritional requirements and physiology of actinomycetes, with special emphasis on: (a) selective isolation of unusual forms; (b) colonial morphogenesis and induction of sporulation; (c) biosynthesis of secondary metabolites of value, with particular reference to antibiotics and industrial enzymes. The future potential of actinomycetes as major contributors of useful bioactive metabolites is also indicated.     

Prediction of dissolved oxygen and carbon dioxide concentration profiles in tubular photobioreactors for microalgal culture

A model is developed for prediction of axial concentration profiles of dissolved oxygen and carbon dioxide in tubular photobioreactors used for culturing microalgae. Experimental data are used to verify the model for continuous outdoor culture of Porphyridium cruentum grown in a 200-L reactor with 100-m long tubular solar receiver. The culture was carried out at a dilution rate of 0.05 h-1 applied only during a 10-h daylight period. The quasi-steady state biomass concentration achieved was 3.0 g. L-1, corresponding to a biomass productivity of 1.5 g. L-1. d-1. The model could predict the dissolved oxygen level in both gas disengagement zone of the reactor and at the end of the loop, the exhaust gas composition, the amount of carbon dioxide injected, and the pH of the culture at each hour. In predicting the various parameters, the model took into account the length of the solar receiver tube, the rate of photosynthesis, the velocity of flow, the degree of mixing, and gas-liquid mass transfer. Because the model simulated the system behavior as a function of tube length and operational variables (superficial gas velocity in the riser, composition of carbon dioxide in the gas injected in the solar receiver and its injection rate), it could potentially be applied to rational design and scale-up of photobioreactors. Copyright 1999 John Wiley & Sons, Inc.     

Productivity, carbon dioxide uptake and net energy return of microalgal bubble column photobioreactors

This work examined the energy return of Chlorella vulgaris and Dunaliella tertiolecta cultivated in a gas-sparged photobioreactor design where the power input for sparging was manipulated (10, 20, and 50 W m⁻³). Dry weight, organic carbon and heating values of the biomass were measured, plus a suite of variables including Fv/Fm and dissolved oxygen. A model for predicting the higher heating value of microalgal biomass was developed and used to measure the energetic performance of batch cultivations. High power inputs enhanced maximum biomass yields, but did not improve the energy return. Cultivation in 10 W m⁻³ showed up to a 39% higher cumulative net energy return than 50 W m⁻³, and increased the cumulative net energy ratio up to fourfold. The highest net energy ratio for power input was 19.3 (D. tertiolecta, 12% CO₂, 10 W m⁻³). These systems may be a sustainable method of biomass production, but their effectiveness is sensitive to operational parameters.     

An integrated wavelength-shifting strategy for enhancement of microalgal growth rate in PMMA- and polycarbonate-based photobioreactors

This study investigates the spectral shifting of UV-A radiation, using fluorescent material, as a tool for enhancing Chlorella sp. growth rate in simulated photobioreactors made of UV-stabilized polycarbonate (PC) and poly (methyl methacrylate) (PMMA). The feasibility of using a fluorescent coating as a wavelength shifter layer, to shift UV-A radiation to the PAR range, was explored. For this purpose, a variety of concentrations of fluorescent dye dissolved in thermoplastic acrylic resin were prepared and used to coat PMMA and PC sheets which then were placed between the radiation source and the culture flask. Compared with the uncoated sheets, the panels coated with the wavelength shifter layer exhibited 74% and 45% (for PC and PMMA substrates, respectively) increase in biomass productivity during the same culture period. It was also found that the elimination of UV-A radiation increased chlorophyll a content of the cells.     

Growth and biochemical characterization of microalgal biomass produced in bubble column and airlift photobioreactors: studies in fed-batch culture

Relatively large (0.19 m column diameter, 2 m tall, 0.06 m³ working volume) outdoor bubble column and airlift bioreactors (a split-cylinder and a draft-tube airlift device) were compared for monoseptic fed-batch culture of the microalga Phaeodactylum tricornutum. The three photobioreactors produced similar biomass versus time profiles and final biomass concentration (4 kg m⁻³). The maximum specific growth rate observed within a daily illuminated period in the exponential growth phase, had a value of 0.08 h⁻¹ on the third day of culture. Because of night-time losses of biomass, the specific growth rate averaged over the 4-days of exponential phase was 0.021 h⁻¹ for the three reactors.

The biomass in the vertical column reactors did not experience photoinhibition under conditions (photosynthetically active daily averaged irradiance value of 1150±52 μE m⁻² s⁻¹) that are known to cause photoinhibition in conventional thin-tube horizontal loop reactors. Because of good gas-liquid mass transfer, the dissolved oxygen concentration in the reactors at peak photosynthesis remained <120% of air saturation; thus, oxygen inhibition of photosynthesis and photo-oxidation of the biomass did not occur. Carbohydrate accumulation (up to 13% w/w) by the biomass was favored during light-limited linear growth. A declining light intensity caused a more than five-fold increase in cellular carotenoids but the chlorophylls increased only by about 2.5-fold during the course of the culture. In the stationary phase, up to 2% of the biomass was chlorophylls and carotenoids constituted up to 0.5% of the biomass dry weight.     

Cannabidiol: an overview of some chemical and pharmacological aspects. Part I: chemical aspects

Over the last few years considerable attention has focused on cannabidiol (CBD), a major non-psychotropic constituent of Cannabis. In Part I of this review we present a condensed survey of the chemistry of CBD; in Part II, to be published later, we shall discuss the anti-convulsive, anti-anxiety, anti-psychotic, anti-nausea and anti-rheumatoid arthritic properties of CBD. CBD does not bind to the known cannabinoid receptors and its mechanism of action is yet unknown. In Part II we shall also present evidence that it is conceivable that, in part at least, its effects are due to its recently discovered inhibition of anandamide uptake and hydrolysis and to its anti-oxidative effect.     

Outdoor helical tubular photobioreactors for microalgal production: modeling of fluid-dynamics and mass transfer and assessment of biomass productivity

The production of the microalga Phaeodactylum tricornutum in an outdoor helical reactor was analyzed. First, fluid dynamics, mass-transfer capability, and mixing of the reactor was evaluated at different superficial gas velocities. Performance of the reactor was controlled by power input per culture volume. A maximum liquid velocity of 0.32 m s(-1) and mass transfer coefficient of 0.006 s(-1) were measured at 3200 W m(-3). A model of the influence of superficial gas velocity on the following reactor parameters was proposed: gas hold-up, induced liquid velocity, and mass transfer coefficient, with the accuracy of the model being demonstrated. Second, the influence of superficial gas velocity on the yield of the culture was evaluated in discontinuous and continuous cultures. Mean daily values of culture parameters, including dissolved oxygen, biomass concentration, chlorophyll fluorescence (F(v)/F(m) ratio), growth rate, biomass productivity, and photosynthetic efficiency, were determined. Different growth curves were measured when the superficial gas velocity was modified-the higher the superficial gas velocity, the higher the yield of the system. In continuous mode, biomass productivity increased by 35%, from 1.02 to 1.38 g L(-1) d(-1), when the superficial gas velocity increased from 0.27 to 0.41 m s(-1). Maximal growth rates of 0.068 h(-1), biomass productivities up to 1.4 g L(-1) d(-1), and photosynthetic efficiency of up to 15% were obtained at the higher superficial gas velocity of 0.41 m s(-1). The fluorescence parameter, F(v)/F(m), which reflects the maximal efficiency of PSII photochemistry, showed that the cultures were stressed at average irradiances within the culture higher than 280 microE m(-2) s(-1) at every superficial gas velocity. For nonstressed cultures, the yield of the system was a function of average irradiance inside the culture, with the superficial gas velocity determining this relationship. When superficial gas velocity was increased, higher growth rates, biomass productivities, and photosynthetic efficiencies were obtained for similar average irradiance values. The higher the superficial gas velocity, the higher the liquid velocity, with this increase enhancing the movement of the cells inside the culture. In this way the efficiency of the cells increased and higher biomass concentrations and productivities were reached for the same solar irradiance.     

Scale-up aspects of photobioreactors: effects of mixing-induced light/dark cycles

The green micro-algae Chlamydomonas reinhardtiiand Dunaliella tertiolecta were cultivated undermedium-duration square-wave light/dark cycles with acycle time of 15 s. These cycles were used to simulatethe light regime experienced by micro-algae inexternally-illuminated (sunlight) air-lift loopbioreactors with internal draft tube. Biomass yieldin relation to light energy was determined as gprotein per mol of photons (400–700 nm). Between 600and 1200 mol m^-2 s^-1 the yield at a10/5 s light/dark cycle was equal to the yield atcontinuous illumination. Consequently, provided thatthe liquid circulation time is 15 s, a considerabledark zone seems to be allowed in the interior ofair-lift loop photobioreactors (33% v/v) without lossof light utilization efficiency. However, at a 5/10 slight/dark cycle, corresponding to a 67% v/v darkzone, biomass yield decreased. Furthermore, bothalgae, C. reinhardtii and D. tertiolecta,responded similarly to these cycles with respect tobiomass yield. This was interesting because they werereported to exhibit a different photoacclimationstrategy. Finally, it was demonstrated that D.tertiolecta was much more efficient at low (average)photon flux densities (57–370 mol m^-2s^-1) than at high PFDs (> 600 mol m^-2s^-1) and it was shown that D. tertiolectawas cultivated at a sub-optimal temperature (20 ^°C).     

Modeling of biomass productivity in tubular photobioreactors for microalgal cultures: effects of dilution rate, tube diameter, and solar irradiance

A macromodel is developed for estimating the year-long biomass productivity of outdoor cultures of microalga in tubular photobioreactors. The model evaluates the solar irradiance on the culture surface as a function of day of the year and the geographic location. In a second step, the geometry of the system is taken into account in estimating the average irradiance to which the cells are exposed. Finally, the growth rate is estimated as a function of irradiance, taking into account photoinhibition and photolimitation. The model interconnects solar irradiance (an environmental variable), tube diameter (a design variable), and dilution rate (an operating variable). Continuous cultures in two different tubular photobioreactors were analyzed using the macromodel. The biomass productivity ranged from 0.50 to 2.04 g L-1 d-1, and from 1.08 to 2. 76 g L-1 d-1, for the larger and the smaller tube diameter photobioreactors, respectively. The quantum yield ranged from 1.1 to 2.2 g E-1; the higher the incident solar radiation, the lower the quantum yield. Simultaneous photolimitation and photoinhibition of outdoor cultures was observed. The model reproduced the experimental results with less than 20% error. If photoinhibition was neglected, and a growth model that considered only photolimitation was used to fit the data, the error increased to 45%, thus reflecting the inadequacy of previous outdoor growth models that disregard photoinhibition. Copyright 1998 John Wiley & Sons, Inc.     

Hydrodynamic aspects and Arthrospira growth in two outdoor tubular undulating row photobioreactors

Two tubular undulating row photobioreactors (TURPs) with a very high illuminated surface/volume ratio (400 m⁻¹) were designed and constructed for the growth of photosynthetic micro-organisms. Experiments were conducted under outdoor conditions; and Arthrospira recycling was performed with airlifts (one for each row). The rows in each reactor faced east-west and consisted of a flexible polyvinyl chloride pipe (22 m long, 0.01 m bore) arranged in a sinusoidal shape. We studied the hydraulic performance of the sine-shaped photobioreactor rows during culture recycling in the TURPs at a very high Reynolds number (4200), when Arthrospira showed Newtonian fluid behavior. The sinusoidal pipe arrangement imposed a sine waveform on the culture, which led to better light utilization. During summer, a volumetric productivity of 2.2 g l⁻¹ day⁻¹ was reached in the TURP-5r (5 rows m⁻²), whereas an area productivity of 35 g m⁻² day⁻¹ was obtained in the TURP-10r (10 rows m⁻²). This was due to more light being available in the TURP-5r, because its rows were more spaced out and the photic ratio (R _f) was low (3.0). In the TURP-10r, the closer rows caused a dilution of the sunlight, but gave a better light distribution inside the Arthrospira culture and improved the light utilization. This was attributed to the high R _f (6.0) of this reactor. Received: 8 October 1999 / Received revision: 20 January 2000 / Accepted: 23 January 2000     

Ontogenetical aspects of nutritional requirements in fish

Changes in morphology i.e. "metamorphosis", mode of respiration, allometric growth of organs, mode of swimming and efficiency of biochemical pathways are briefly reviewed. It is suggested that these processes form the basis for progressive changes in nutrient requirements involving formation and development of several organs, systems and function. Digestive tract morphology changes during ontogenesis and aspects of fish metabolism, protein synthesis rate and body growth rate are interconnected and an attempt is made to explain these processes so as to understand the specificity of larval and juvenile fish nutrient requirements as compared to subadults. Protein and amino acid requirements given the body mass perspective and the generalization of the protein maintenance requirement in protein requirements for maximum growth was estimated to amount to 5-20%. Several cases of amino acid deficiency symptoms showed strong dependence on fish weight (age), but even most numerous studies on salmonids are lacking complete research throughout the life history of one species in defined nutritional and environmental conditions. Larval and juvenile fish have reduced capacity of catabolic adaptability and this fact links them to strictly carnivorous mammals. An attempt is made, for the first time, to relate amino acid needs of fish to young and/or carnivorous mammals. Vitamin requirements of fish are reviewed, taking into account the relationship between body size and time of the first appearance of deficiency symptoms. These are virtually no studies on vitamin requirements in larval warm-water fishes and very few on first feeding salmonids. The same applies to the vitamin need in reproductory fish. Fatty acid deficiencies manifest themselves faster in juvenile fish, but larval fish might require separate classes of lipids, phosphatidylcholine, in the diet to develop and grow at all. It seems that the studies on nutrient requirements have so far not used an ontogenetical perspective, but evidence given throughout this work argues that it would be worthwhile.