Journal of Biotechnology 151 (2011) 271–277

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Journal of Biotechnology
journal homepage: www.elsevier.com/locate/jbiotec

Growth of Spirulina platensis enhanced under intermittent illumination

Shengzhang Xue a,b , Zhenfeng Su a,b , Wei Cong a,∗
a
National Key Laboratory of Biochemical Engineering, Institute of Process Engineering, Chinese Academy of Sciences, P.O. Box 353, Beijing 100190, PR China
b
Graduate University of the Chinese Academy of Sciences, Beijing, PR China

a r t i c l e i n f o a b s t r a c t

Article history: The growth characteristics of microalgae under different light conditions (continuous or intermittent)
Received 8 August 2010 are essential information for photobioreactor design and operation. In this study, we constructed a thin-
Received in revised form 6 December 2010 layer (10 mm) flat plate photobioreactor device with a light/dark (L/D) alternation system to investigate
Accepted 10 December 2010
the growth of Spirulina platensis under two different light regimes: (1) continuous illumination in a
Available online 17 December 2010
wide range of light intensities (1.00–77.16 mW cm−2 ); (2) intermittent illumination in medium frequency
(0.01–20 Hz). Specific growth rate and light efficiency based on biomass production were determined for
Keywords:
each round of experiment. Four regions (light limited region, intermediate region, light saturated region
Microalgae
Spirulina platensis
and light inhibition region) were recognized according to the results under continuous illumination.
Photobioreactor Under intermittent illumination, when L/D frequency increased from 0.01 Hz to 20 Hz, specific growth
Light/dark cycles rate and light efficiency were enhanced. However, the enhancement was different, depending on the
Specific growth rate applied light intensity and light fraction. The higher the light intensity, the greater the enhancement
Light efficiency would be when L/D frequency increased from 0.01 Hz to 20 Hz; and the higher the light intensity, the lower
the light fractions is needed to maintain light efficiency as high as that under continuous illumination in
light limited region.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction
Microalgae, a photoautotrophic microorganism, use inexpen-
sive natural resources, e.g. CO2 , H2 O and inorganic salts, to
transform radiant energy into valuable products and provide pos-
sible solutions to the urgent issues facing human beings today,
such as resource and energy shortage, global warming and environ-
mental problems. The need for further upscale of their commercial
production has been recognized in recent literature (Chisti, 2007;
Pulz and Gross, 2004; Raja et al., 2008; Spolaore et al., 2006).
Spirulina platensis, has been successfully commercialized as a
nutritional supplement for humans and animals (Cohen, 1997);
and as a source of active ingredients in pharmaceuticals and
cosmetics (Belay et al., 1993). This microalga has also been suc-
cessfully employed in integrated systems for wastewater treatment
(Lalibertı̌e et al., 1997), as well as recovery and reutilization of heavy
metals as adsorbent materials (Converti et al., 2006; Solisio et al.,
2006).
According to Richmond (2004), light availability is one of the
most important factors that may limit the growth of microalgae
in any algal systems, thus, efficient use of light energy is crucial
for commercial production of microalgae. Because of self-shading
∗ Corresponding author. Tel.: +86 10 82627060; fax: +86 10 82627074.
E-mail address: weicong@home.ipe.ac.cn (W. Cong).
effect among algal cells, the light regime inside an outdoor photo-
bioreactor is characterized by a light gradient with full sunlight at
the light-exposed surface (photic zone) and total darkness in the
interior of the bioreactor (aphotic zone). Consequently, depend-
ing on the mixing characteristics of the culture suspension, algal
cells will be exposed to a series of light/dark (L/D) cycles. Most fre-
quently, these L/D cycles were in medium range (Grobbelaar, 1989;
Janssen et al., 2000b), typically 0.01–20 Hz. Growth and/or oxygen
evolution characteristics of microalgae under different L/D cycles,
which are quite important for efficient photobioreactor design and
operation, have been investigated extensively during the past few
decades for a wide range of species, e.g. Chlorella (Grobbelaar et al.,
1992; Lee and Pirt, 1981; Phillips and Myers, 1954), Scenedesmus
(Grobbelaar et al., 1996; Nedbal et al., 1996), Phaeodactylum (Terry,
1986), Porphyridium (Merchuk et al., 1998; Wu and Merchuk, 2001),
Chlamydomonas, Dunaliella (Janssen et al., 2000a). However, most
of the L/D frequencies adopted, except for the work by Grobbelaar
et al. (1996), were either too low (<0.1 Hz) or too high (>100 Hz), and
the light intensities applied were much lower than that of sunlight.
Thus, previous studies could not fully describe the real light histo-
ries experienced by algal cells in outdoor photobioreactors and was
not able to predict the degree of benefit of productivity increase by
intensified operation. In addition, S. platensis has not been studied.
In this work, a thin-layer (10 mm) flat-plate photobioreactor
device with an L/D alternation system was constructed to give
medium frequency (0.01–20 Hz) L/D cycles. White high power LEDs

0168-1656/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.jbiotec.2010.12.012
272 S. Xue et al. / Journal of Biotechnology 151 (2011) 271–277

Fig. 1. The photobioreactor device with L/D alternation system. (1) Thin-layer flat plate photobioreactor, (2) air compressor, (3) CO2 cylinder, (4) gas flowmeters, (5)
thermal-stat water bath, (6) light source, (7) radiator, (8) timer, (9) DC power supply, (10) partly darkened rotating disc, (11) motor, (12) rotating shaft, (13) speed regulator.

(light emitting diodes) were used as the light source to provide a water jacket of the reactor. Thus, the culture conditions, except for
wide range of light intensities (0–77.16 mW cm−2 ), the highest of light regime, were kept constant for each round of experiment.
which was comparable to that of direct sunlight at summer noon.
This is important for our investigations because most commercial
2.2. Experimental set-up
productions of microalgae are conducted outdoors. Subsequently,
investigation of S. platensis growing under different light conditions
Experiments were carried out in a photobioreactor device with
(continuous and intermittent) was carried out in this device. Spe-
an L/D alternation system as shown in Fig. 1. It included the fol-
cific growth rate (SGR) and light efficiency (LE) based on biomass
lowing parts: a thin-layer flat plate photobioreactor (1) equipped
production were determined for each round of experiment to inves-
with thermal-stat and aeration systems; a light source (6) which
tigate its growth characteristics under various lighting conditions.
could give light intensities as high as 77.16 mW cm−2 , comparable
Experimental results and their significance to practical aspects of
to that of direct sunlight at summer noon; an L/D alternation sys-
photobioreactor design and operation were discussed herein.
tem which could give L/D cycles of medium frequency (0.01–20 Hz),
as well as full range of light fractions (the ratio of light period in a
2. Material and methods whole L/D cycle, 0–1); and a darkroom system which kept the reac-
tor from ambient light. A more detailed description of this device
2.1. Organism and culture condition will follow.

S. platensis (breed F3 , kindly provided by the Biochemical Engi-

2.2.1. Reactor design
neering Laboratory of Yantai University, Yantai, China) was grown
The reactor (Fig. 2) used in this device is a flat plate, bubble col-
in Zarrouk medium (1966) which included the following nutri-
umn photobioreactor with a short light path (10 mm). The effective
ents (g L−1 ): NaHCO3 , 16.601; K2 HPO4 , 0.647; NaNO3 , 2.470; NaCl,
volume is about 60 mL. Air or CO2 enriched air (10%) is bubbled
0.988; MgSO4 , 0.096; K2 SO4 , 0.988; CaCl2 , 0.040; FeSO4 , 0.005;
from a glass frit core (average aperture 80 ␮m) into the culture
EDTA–Na2 , 0.080; microelement solutions (mg L−1 ): H3 BO4 , 2.860;
chamber of the reactor, which provides well mixing of the culture
MnCl2 ·4H2 O, 1.145; CuSO4 ·5H2 O, 0.080; (NH4 )6 Mo7 , 0.019; ZnSO4 ,
suspension. The front side of the reactor is illuminated by a light
0.123. The medium was sterilized at 120 ◦ C for 15 min before use.
source (6 in Fig. 1, described below), while the back side is a water
S. platensis was cultivated in two kinds of light regimes:
jacket for thermal-stating water circulation to maintain the culture
(1) continuous illumination in a rang of light intensities
temperature.
(1–77.16 mW cm−2 ); (2) intermittent illumination of four differ-
ent light intensities (3.09, 8.16, 25.87, 77.16 mW cm−2 ) and L/D
frequencies between 0.01 and 20 Hz. The cultivation process was
as follows: after acclimation to a certain light intensity for one or
two days, the algal suspension in its exponential phase was diluted
with fresh medium to a certain cell concentration to give an optical
density of 0.390 ± 0.005 at 560 nm (OD560 ). CO2 -enriched air (10%)
was bubbled into the algal suspension until its pH value reached
9.25 ± 0.03. Then the diluted algal suspension was inoculated to
a thin-layer photobioreactor (described below); afterward it was
exposed to a certain light regime for 24 h. OD560 was recorded at the
end of the growing process. Temperature was maintained at 33 ◦ C
throughout the process by circulating thermal-stated water to the Fig. 2. Experimental thin-layer flat-plate photobioreactor.
 S. Xue et al. / Journal of Biotechnology 151 (2011) 271–277
2.2.2. Light source
Illumination was provided by an array (10 × 10) of high-power
white LEDs (light emitting diodes) in conjunction with a regulated
DC power supply (9). The LEDs were mounted on a panel with an
area of 22 mm × 22 mm, and placed opposite to the front side of
the reactor to provide illumination. According to the manufacturer
(Lucky Light Electronics Co., Ltd.), its rated power was 100 W with
the relative spectral distribution shown in Fig. 3.
LED is a comparatively new light source with the features of
high efficiency, long life-expectancy and more importantly, quick
response and quick stabilization characters. Several researchers
(Grobbelaar et al., 1996; Matthijs et al., 1996; Nedbal et al., 1996,
2008; Park et al., 2000) used LED as the light source to cultivate
microalgae, though most of their LEDs failed to attain high light
intensities because of technical reasons. Our LED panel could pro-
vide ultra-high light intensity, comparable to direct sunlight at
summer noon. High-power LEDs call for efficient heat removal to
ensure the life span. In our experiment, a water cooling radiator,
usually used on computer CPU, was slightly modified to fit on the
back of the panel to remove the heat generated by the LED panel.
2.2.3. Intermittent illumination
Two modes to realize L/D alternation were integrated in this
device: for L/D cycling time greater than 1 s, the LED panel (6)
was switched ‘on’ and ‘off’ using a timer (8) in conjunction with
a high capacity regulated DC power supply (9); for L/D cycling time
less than or equal to 1 s, intermittent illumination was obtained
by rotating a partly darkened disc. In the first case, light fractions
and L/D frequencies can be easily adjusted by regulating the ‘on’
and ‘off’ time; while in the latter case, different light fractions were
obtained by adjusting the fraction of darkened area on the rotat-
ing disc, and the L/D frequency was adjusted by setting different
rotating speed of the disc as shown in Fig. 4.
2.2.4. Light intensity
Light intensity (mW cm−2 ) was measured in the PAR-range
(photosynthetic active radiation, 400–700 nm) by a PAR Sensor
(FGH-1, Photoelectric Instrument Factory of Beijing Normal Uni-
versity, Beijing, P.R. China). The photodiode of the PAR Sensor was
placed at different positions on the illuminated surface of the reac-
tor when light intensities were measured and averaged. These
values precluded the absorption of the transparent part of the rotat-
ing disc and the front wall of the reactor.
The measured values of the PAR Sensor was in mW cm−2 which
can be converted to PFD (photo flux density, ␮mol m−2 s−1 ) by a fac-
tor of 43.6 (1 mW cm−2 = 43.6 ␮mol m−2 s−1 , Biggs, 1984) for white
LED. The conversion factor depends on the spectral distribution of
the light source, and varies for different light sources.
Fig. 3. Relative spectral distribution of the LED panel.
273
Fig. 4. Front view of the partly darkened rotating disc and the reactor.
2.3. Analytical methods
Previous investigation showed that S. platensis was growing in
its exponential phase, although with some small errors, between
CDW 0.15–2.0 g L−1 under continuous light. For each round of
experiment, specific growth rate (SGR) and light efficiency (LE)
based on biomass production were determined to investigate
growth characteristics of S. platensis under different lighting con-
ditions:
ln CDW1 − ln CDW0
SGR = (1)
t1 − t0
(CDW1 − CDW0 ) × HG × V
LE = × 100% (2)
(I0 − I1 ) × S × (t1 − t0 ) × 10−3
where CDW0 and CDW1 (g L−1 ) are cell dry weight of S. platen-
sis at the starting time t0 and the ending time t1 (s), respectively;
HG (kJ gCDW −1 ) is the enthalpy of dry biomass (for S. platensis,
21 kJ gCDW −1 ; Watanabe and Saiki, 1996); V is the culture volume
(L); S is the illuminated area of the bioreactor (cm2 ); I0 and I1
(mW cm−2 ) are the incident and outlet light intensity of the biore-
actor, respectively. Above all, the numerator of Eq. (2) means the
chemical energy immobilized by the photosynthetic microorgan-
ism, where the denominator means the light energy irradiated into
the culture.
According to our experimental results, CDW showed good
linearity with OD560 of S. platensis suspension in the range of
0.10–0.90, as described by Eq. (3) with a correlation coefficient (R2 )
of 0.997:
CDW = 0.672 × OD560 + 0.028 (3)
where OD560 was measured by a visible spectrophotometer (723N,
Shanghai Precision & Scientific Instrument Co., LTD, P.R. China)
against a reference of fresh culture media; and CDW was deter-
mined by filtration of a sample (10 mL) through Millipore filters
(0.45 ␮m pore diameter) after washing with deionized water to
eliminate salt precipitates; the filters were then dried in oven at
120 ◦ C for 6 h and weighed for determination of cell dry weight. For
samples of OD560 being larger than 0.9, the algal suspension was
firstly diluted by fresh media to a certain multiple to give OD560
value below 0.9 before CDW was determined.
3. Results and discussion
In order to reflect the true responses of microalgae to differ-
ent light conditions it is desirable that the light history perceived
by each algal cell in the reactor is exactly the same. In this study,
because of three reasons: (1) the reactor used was a flat plate thin-
layer one, (2) cell concentration during growth was relatively low,
274 S. Xue et al. / Journal of Biotechnology 151 (2011) 271–277
Fig. 5. Specific growth rate (SGR, ) and light efficiency (LE, ) of S. platensis under
continuous illumination of different light intensities. The solid line connecting the
solid squares represents a B-spline fit of SGR.
and (3) adequate mixing conditions in the bubble column reactor
minimized the self-shading effects among algal cells, each algal cell
inside the reactor was assumed to have the same opportunity to
receive light, thus implying that each experiment truly represents
the physical responses of algal cells to various light conditions, irre-
spective of the flowing conditions in the reactor. This assumption
holds for all the experimental runs in this study.
3.1. Continuous light of various intensities
The knowledge of quantitative dependence of SGR of S. platensis
on light intensity, though not much reported heretofore, is quite
important for photobioreactor design and operation. Qiang (Qiang
and Richmond, 1996) and Ravelonandro et al. (2008) investigated
the growth of S. platensis under different light intensities, how-
ever, they failed to preclude self-shading effects among algal cells
by employing dense culture or long light-path reactor, thus fail-
ing to reflect the true responses of S. platensis to various light
intensities. Moreover, the light intensity ranges they covered were
limited.
Fig. 5 plots SGR and LE of S. platensis under low optical den-
sity (OD560 = 0.390) in the thin-layer (10 mm) photobioreactor as a
function of light intensity under continuous illumination. Appar-
ently, light intensity can be separated into four regions according
to its relationship with SGR:
(1) light limited region (0–3.5 mW cm−2 ): SGR increases linearly
with increasing light intensities, indicating first-order kinetics.
The light intensities applied by Ravelonandro et al. (2008) for
S. platensis growth was mostly in this range.
(2) intermediate region (3.5–10 mW cm−2 ): SGR increases slowly
with increasing light intensities, illustrating the reaction to fol-
low mixed-order (0–1) kinetics;
(3) light saturated region (10–30 mW cm−2 ): SGR almost keeps
constant with increasing light intensities, indicating zero-order
kinetics. The highest value of SGR was about 1.9 day−1 . This is
highly in accordance with the result of Olaizola (Olaizola and
Duerr, 1990) who found the grow rate of S. platensis was light-
saturated at irradiance around 450 ␮mol m−2 s−1 .
(4) light inhibition region (>30 mW cm−2 ), SGR decreases rapidly
with increasing light intensities, indicating complete inhibi-
tion by light. For instance, when light intensity increased to
50.34 mW cm−2 , the SGR was only 0.539 day−1 , even smaller
than that under 1 mW cm−2 . We have also tested the growth
of S. platensis under light intensities larger than 60 mW cm−2 ,
showing even negative SGR, possible due to cell degradation
under prolonged light inhibition (Wu and Merchuk, 2001).
The highest LE was found in light limited region, e.g. around 8%
with some fluctuations. For the other three regions, LE decreased
rapidly with increasing light intensities, to approach zero when
light intensity increased to 50.34 mW cm−2 . Apparently, under light
limited condition, where SGR increased linearly with increasing
light intensities, all of the light energy was utilized to produce
biomass, while in the intermediate region, only part of the light
input was utilized to produce biomass, the remainder being lost as
heat and fluorescence (Janssen et al., 2000b). Hence, LE began to
decrease as compared to that in light limited condition. This effect
became more obvious in light saturated and light inhibition region,
where increased light input did not lead to any biomass increase or
even decrease it. As a consequence, light efficiencies continued to
decrease with increasing light intensities. For light intensity larger
than 60 mW cm−2 , LE was negative (the data were not included in
Fig. 5) because of negative biomass increase. The results could be
explained by biological adaptation mechanisms (Rier et al., 2006):
in weak light, algal cells are prone to synthesize more pigments
to capture light, and thus could use photons in a more economic
mode; while in strong light, algal cells do not need too much light
and are apt to waste photons. This feature could be utilized by algal
production by diluting solar light through partly overlapped photo-
bioreactors (Carlozzi, 2003) or using light redistribution materials
in photobioreactors, such as optical fibers (Janssen et al., 2003).
3.2. Growth characteristics of S. platensis under intermittent
illumination
As stated above, for reliability of interpreting the final results it
is desirable that the light flashes seen by each algal cell in the reac-
tor be exactly the same. For the first mode to realize L/D alternation
(using a timer), it is correct, while for the second mode (rotating a
partly darkened disc), some small and unmeasurable error is intro-
duced since the algal cells are themselves moving erratically in
the suspension during exposure to the transparent portion of the
disc. However, this error is insignificant and can thus be neglected
because the rotating disc moves much faster than the algal cells
under the operation conditions.
3.2.1. Specific growth rate
Four light intensities, 3.09, 8.16, 25.87 and 77.16 mW cm−2 , cor-
responding to the four different regions respectively, were utilized
to investigate the influence of different-frequency L/D cycles on S.
platensis growth, the results of which were shown in Fig. 6.
Fig. 6 showed that SGR increased with increasing L/D frequency
for all four light intensities but with different degrees. Under
light limited condition (Fig. 6(a)), a minor increase was detected.
The higher the light intensity, the greater the SGR increase was
observed. The largest SGR increase under our experimental con-
ditions was found under 77.16 mW cm−2 (light inhibited region)
when L/D frequency increased from 0.01 Hz to 20 Hz. In general,
for all the four light intensities, high frequency L/D cycles led to
high SGR; low frequency L/D cycles (<0.1 Hz) led to low or even
negative SGR. It implied that long stay of algal cells in dark zone
was uneconomical for algal culture because some of the biomass
might be wasted for maintaining in dark.
The effects of medium frequency L/D cycles to microalgae
growth have been widely investigated, whereas conflicting results
were found. On one hand it was suggested that L/D cycles of
medium frequency enhanced productivity greatly (Lee and Pirt,
1981; Merchuk et al., 1998; Terry, 1986); whereas several others
found that medium frequency L/D cycles exerted little influence on
productivity (Grobbelaar, 1989, 1991). In addition, Janssen (Janssen
 S. Xue et al. / Journal of Biotechnology 151 (2011) 271–277 275

Fig. 6. Specific growth rate of S. platensis under L/D cycles of four different light intensities as affected by L/D frequency. Four plots take the same scale for both the
X (logarithmic) and Y coordinates. The two dashed lines, the lower one and the upper one, in each diagram represent no light integration (SGR = k × f (I)) and full light
integration (SGR = f (k × I)), respectively, under that specific light intensity (I) and light fraction (k). The spacing between the two lines represented the enhancement potential
by applying L/D cycles to algal culture as compared to continuous lighting.

et al., 2000a,b) concluded that whether L/D cycles of medium fre-
quency could enhance productivity depended on the light fraction
of the L/D cycle. However, our results clearly showed that whether
medium frequency L/D cycles could increase SGR largely depended
on the applied light intensity. This result is meaningful for outdoor
microalgae production. Because, however, a high L/D frequency in
photobioreactor requires a high degree of turbulence and thus a
large stirring or pumping energy input, whether it pays largely
depends on the incident light intensity. At noon when the light
intensity is high, algal culture is much favored by intensified tur-
bulence, which gives an overall high L/D frequency, than that at
morning or afternoon when the light is weak. The higher the light
intensity, the more turbulence will be desirable.
3.2.2. Light integration effect
The effect of L/D cycles on algae is often discussed with the help
of the term “light integration effect” (Terry, 1986). In continuous
light, the specific growth rate or oxygen production rate (denoted
as SGR here) can be written as a saturating function of light inten-
sity (I): SGR = f (I). When an L/D cycle with a light fraction of “k” is
applied to algae, with full light integration, SGR = f (k × I); whereas
with no light integration, SGR = k × f (I). Since the SGR is a saturating
function of light intensity, it is clear that: f (k × I) > k × f (I). Full light
integration means the algae would be able to use intermittent light
as efficiently as continuous lighting of the same time-averaged PFD,
while no light integration means that the SGR under L/D cycles is
proportional to that under continuous illumination of same light
intensity by a factor of light fraction, thus no LE enhancement.
Fig. 6 showed that the SGR of most experimental runs under
intermittent illumination was between no light integration and full
light integration, and the degree of light integration had a positive
correlation with L/D frequency as a whole. This result was in accor-
dance with previous works (Phillips and Myers, 1954; Terry, 1986).
As can be seen in Fig. 6(a), SGR under L/D frequencies less than
1 Hz was smaller than the lower line, which indicated that light
integration effect was totally absent under these conditions. The
reason was possibly that under long L/D cycles of low light intensity
(light limited condition) respiration effect during the dark period
of L/D cycles began to exert larger influence than the light inte-
gration effect, caused the SGR to be smaller than that of zero light
integration. Therefore, it is not economical for S. platensis to grow
under long L/D cycles of low light intensity. This finding was con-
sistent with Ohira et al. (2000) who concluded that S. platensis had
no light integration effect. However, this was not true when it came
to high frequency (>1 Hz) L/D cycles as was clearly shown in all four
plots in Fig. 6. According to previous study (Matthijs et al., 1996;
Nedbal et al., 1996; Phillips and Myers, 1954), full light integration
was found under high frequency (>100 Hz) L/D cycles. In our study,
however, full light integration has not been achieved but is more
closely approached, especially under high frequency L/D cycles of
ultra-high irradiance.
Fig. 6(d) also indicates that SGR of S. platensis under ultra-high
irradiance will show negative if the light period is too long (>10 s)
in the L/D cycle. This corresponded to the results under contin-
uous illumination where SGR showed negative as light intensity
surpassed 60 mW cm−2 . It indicated that S. platensis would not be
sustainable if it was exposed to a high irradiance (77.16 mW cm−2 )
for more than 10 s.
3.2.3. Light efficiency
Light efficiency is another important parameter to evaluate
microalgae growth under different conditions. Fig. 7 showed that
276 S. Xue et al. / Journal of Biotechnology 151 (2011) 271–277
Fig. 7. Light efficiency of S. platensis under intermittent illumination of four different
light intensities as affected by L/D frequency.
LE under L/D cycles was largely dependent on the light intensities
applied. High LE was derived under low light intensity; the higher
the light intensity, the lower the LE. However, when it came to LE
increase due to increased L/D frequency, the largest increase was
detected under the largest intensity, corresponding to the results
under continuous light.
Light efficiency enhancement due to increased L/D frequency is
also affected by the light fractions of the L/D cycles, as was clearly
shown in Fig. 8. Light efficiency under light fraction of 0.05 is higher
than that of 0.1 and 0.2, especially when it comes to high frequency
L/D cycles. It is an interesting finding for S. platensis that LE increases
when light fraction decreases under the same L/D frequency. The
possible reasons were that under L/D cycles of large k values, the
photosynthetic units of S. platensis were light-saturated before light
period ended for each cycle, and the following incident light was
wasted for most part. While under small-k-L/D-cycles, where the
light period were shorter compared to that of large-k-L/D-cycles,
algal cells could timely leave light period and enter dark period
to stabilize carbon, thus making high light efficiency. There was
another way to explain this phenomenon. As was stated above,
full light integration, which means the algal cells would be able
to use intermittent light as efficiently as continuous light of the
same time-averaged light intensity, k × I, could be more closely
approached under high frequency L/D cycles. Under continuous
Fig. 8. Light efficiency of S. platensis under intermittent illumination of different
light fractions of ultra-high irradiance.
illumination, light efficiency had a negative correlation with light
intensity, thus leading to the light efficiency increasing when k was
reduced under high frequency L/D cycles of the same light intensity.
This finding provided basis and expectations for high/ultra-high cell
density culture which could obtain an overall high light efficiency
if it was operated under high light intensity and high L/D frequency
conditions.
High efficiency microalgae culture does not just happen; it
involves a comprehensive consideration of photobioreactor design
and operation. Based on our results, the overall light efficiency of
a photobioreactor is mainly determined by two factors: diluted
light intensity (k × I) and L/D frequency. High efficiency microal-
gae culture could be expected only if the diluted light intensity
enters the light limited region (0–3.5 mW cm−2 ), and the over-
all L/D frequency in reactor is high enough to make sure all the
light energy absorbed by algal cells could be utilized efficiently
to produce biomass. In other words, no light energy is wasted to
generate heat and/or fluorescence. Efficient photobioreactor design
and operation should take good advantage of these two factors: (1)
reactor thickness along light-path and algal concentration should
be appropriate to give a certain light fraction, making sure that
k × I enters the light limited region, and (2) photobioreactor con-
figuration should facilitate the operation conditions to generate
controllable flowing conditions which could give desired L/D cycles.
For example, under low light intensities, high L/D frequency was
not needed because the SGR and LE enhancement was limited by
increased L/D frequency. While under high light intensities, inten-
sified turbulence should be imposed to algal suspension to speed up
the shift of algal cells between light region and dark region, which
helps generate high frequency L/D cycles.
4. Conclusions
A thin-layer photobioreactor device with an L/D alternation
system was constructed to investigate S. platensis growth char-
acteristics under continuous and intermittent illumination of
various light intensities. Four regions, namely light limited region
(0–3.5 mW cm−2 ), intermediate region (3.5–10 mW cm−2 ), light
saturated region (10–30 mW cm−2 ) and light inhibition region
(>30 mW cm−2 ), were recognized according to their relationship
to SGR under continuous illumination. The highest light efficiency
was obtained under light limited region; while in the other three
regions, LE decreased rapidly with increasing light intensity. Light
efficiency in light saturated and light inhibited region was compar-
atively low, which can be significantly improved by applying low
light fraction, high frequency L/D cycles to algal culture.
The practical expectation of the present work is the contribu-
tion of intensified turbulence and high-density algal suspension
to increase growth rate and light efficiency in algal cultures. The
feasibility of increasing turbulence depends on the gain in light effi-
ciency as compared to increased power requirement for stirring or
pumping the suspension. In general, the higher the light intensity,
the more turbulence will be desirable, taking into account of the
pumping cost.
Acknowledgements
The authors wish to thank the support of National Key Technol-
ogy R&D Program of China (2006BAD09A12).
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