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Effect of light intensity and pH condition on the growth, biomass and lipid
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DOI: 10.1080/17597269.2015.1045274

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Effect of light intensity and pH condition on the


growth, biomass and lipid content of microalgae
Scenedesmus species
a c c ab
Amrita Difusa , Jayanta Talukdar , Mohan Chandra Kalita , K. Mohanty & Vaibhav V.
ab
Goud
a
Centre for Energy, Indian Institute of Technology Guwahati, Assam-781039, India
b
Department of Chemical Engineering, Indian Institute of Technology Guwahati,
Assam-781039, India
c
Department of Biotechnology, Guwahati University, Guwahati-781014, Assam, India
Published online: 26 May 2015.
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To cite this article: Amrita Difusa, Jayanta Talukdar, Mohan Chandra Kalita, K. Mohanty & Vaibhav V. Goud (2015): Effect
of light intensity and pH condition on the growth, biomass and lipid content of microalgae Scenedesmus species, Biofuels,
DOI: 10.1080/17597269.2015.1045274

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Biofuels, 2015
http://dx.doi.org/10.1080/17597269.2015.1045274

Effect of light intensity and pH condition on the growth, biomass and lipid content of microalgae
Scenedesmus species
Amrita Difusaa, Jayanta Talukdarc, Mohan Chandra Kalitac, K. Mohantya,b and Vaibhav V. Gouda,b*
a
Centre for Energy, Indian Institute of Technology Guwahati, Assam-781039, India; bDepartment of Chemical Engineering, Indian
Institute of Technology Guwahati, Assam-781039, India; cDepartment of Biotechnology, Guwahati University, Guwahati-781014,
Assam, India

(Received 15 June 2014; accepted 18 April 2015)

The physiological factors for microalgae cultivation are the preliminary steps to overcome the potential biomass demand.
In this regard, the influence of light intensity (at six levels: 27, 40.5, 54, 67.5, 81 and 94.5 mmol m¡2s¡1) and pH
conditions (5.0, 6.0, 7.0, 8.0 and 9.0) on the growth and lipid production of Scenedesmus species ADIITEC-II and
GUBIOTJT116 was compared in the batch culture. Among the studied species, maximum specific growth (0.334 §
0.002 day¡1) and highest biomass yield (0.41 § 0.021 g L¡1) was achieved for Scenedesmus species ADIITEC-II at light
intensity 81 mmol m¡2s¡1. Similarly, Scenedesmus species GUBIOTJT116 showed increased growth and biomass
Downloaded by [New York University] at 00:56 30 May 2015

production at 81 mmol m¡2s¡1. However, illumination under 94.5 mmol m¡2s¡1 showed higher lipid production (0.533 §
0.001 g L¡1 and 0.551 § 0.001 g L¡1) for Scenedesmus sp. ADIITEC-II and GUBIOTJT116, respectively. The significant
variation in effective pH tolerance, growth and biomass production was observed in the studied pH range. The result
obtained in this study provides a potential insight of the strains and could be further explored to improve the biomass and
lipid yield.
Keywords: light intensity; pH condition; Scenedesmus species; biomass; lipid

The substantial utilization of fossil fuel has contributed to optimum pH condition for the microalgae growth allows
growing concern over the search for sustainable alternate intense culture photosynthesis, which responds to an
energy sources to satisfy the existing energy demand. In increase in pH of the medium condition towards the sta-
lieu, microalgae are reckoned to be the entity with most tionary phase and which lowers the microalgae growth
potential for sustainable fuel generation and as well as rate.
able feedstock to mitigate the emitting carbon dioxide Light intensity also has a significant effect on the
which is one of the major green house gases to the atmo- growth of microalgae species. Microalgal photosynthetic
sphere.[1] In this regard, microalgae cultivation scores an kinetics utilize light condition to conduct their metabolic
extensive consideration in terms of biomass production activities.[5] Microalgae cells possess certain light satura-
for various valuable products and has been involved in tion limits (6.458 KLux) which are used as a source of
many strategic ways, leading from raceway ponds to energy for synthesizing cell protoplasm and limit the utili-
photo bioreactors. Microalgae cultivation is always main- zation of light when exposed beyond its saturation limit.
tained with a combination of basic physiological factors [6] It plays a crucial role in photoautotrophic microalgae
like light intensity, pH and temperature for its optimum cultivation, and its requirement varies depending upon the
physicochemical and biological processes. To refer, culture depth and density of the microalgae culture. The
microalgae species shows a critical influence towards the higher depth and cell concentrations of the microalgae
pH condition. Often the water bodies covered with micro- culture requires increased exposure of light intensity to
algae blooms have a lethal effect for several microalgae penetrate through the culture medium.[6] This present
species, where the increased microalgae population cre- study focused on the aspect of growth, biomass and lipid
ates high photosynthetic assimilation of carbon into bio- production of two natives isolates of Scenedesmus species
mass, which goes beyond the supply of CO2 into seawater under different light intensity and constant pH condition.
resulting an elevated pH.[2] However, microalgae are
very diverse in terms of pH tolerance and are very species
Materials and methods
specific. To compare, some species of microalgae (espe-
cially diatoms and large dinoflagellates) are very sensitive Microalgae strain and culture medium
to elevated pH (over pH 8.5) and some bloom-forming or Two microalgae strains were used in the study: Scenedes-
red-tide coastal species tolerate extremely high pH-levels mus species ADIITEC-II and Scenedesmus species
(above pH 9.0).[3] According to Goldman et al. [4] the GUBIOTJT116. Scenedesmus species ADIITEC-II was

*Corresponding author. Email: vvgoud@iitg.ernet.in

Ó 2015 Taylor & Francis


2 A. Difusa et al.

isolated from the native wetland of North East India and of biomass, known volume of algal suspension was fil-
Scenedesmus species (GUBIOTJT116) was obtained from tered through a pre-dried and pre-weighed glass fiber filter
Culture Collection of Microalgae Division, Department of (GF/C). The filter paper with biomass of algae was then
Biotechnology Gauhati University, India. The isolated dried at 80  C for 6 h and weighed until the constant dry
microalgae cultures were raised aseptically under con- weight, allowed to cool in the desiccator and then weighed
trolled laboratory conditions. The medium used during again.[8]
this study was the BG-11 medium which contains NaNO3
1.5 g, K2HPO4.3H2O 0.04 g, KH2PO4.3H2O 0.2 g, EDTA
0.0005 g, Fe ammonium citrate 0.005 g, citric acid Lipid extraction
0.005 g, Na2CO3 0.02 g and 1 ml of trace metal solution The total lipids were extracted using a modified Bligh and
per liter. The trace metal solution contains H3BO3 2.85 g, Dyer method.[9] A known volume of the culture was har-
MnCl2.4H2O 1.8 g, ZnSO4.7H2O 0.02 g, CuSO4.5H2O vested during a late exponential growth phase by centrifu-
0.08 g, CoCl2.6H2O 0.08 g and Na2MoO4.2H2O gation at 5000 rpm for 35 min. The known amount of
0.05 gl¡1. The pH of a medium was adjusted to 7.5 with dried cell biomass was finely ground with 2:1 mixture of
either 1N HCl or 1N KOH solution prior to autoclaving. chloroform (CHCl3) and methanol (CH3OH). The residue
The cultures were incubated at temperature 25 § 2 C and was extracted 23 times with chloroform or till it became
16:8 h light and dark photoperiod. colorless. The filtrate was collected, and a biphasic layer
was obtained on addition of 0.9% NaCl solution (1/3rd of
the volume). The lower organic (CHCl3) layer containing
Microalgal cultivation
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the lipid components was collected in a clean pre-weighed


The two isolates of Scenedesmus species were cultivated glass vial. The solvent was evaporated at 60  C in a water
at six different light intensities (27, 40.5, 54, 67.5, 81 and bath to near dryness and dried in desiccators under
94.5 mmol m¡2s¡1). The designated irradiances were reduced pressure over anhydrous Na2SO4. The dried lipid
achieved using a photometer (Lux meter: Metravi 1330, extract was measured gravimetrically and expressed in
India). Growth of cultures was initiated by the introduc- gram per liter (gl¡1) of total lipids per dry cell weight
tion of inoculums containing about 0.5 £ 106 to 2.0 £ 106 (DCW).
cells ml¡1. All of the experiments were carried out in a
250 ml culture flask (batch mode) for a culture duration of
10 days, and prior to autoclaving initial culture pH was Statistical analysis
adjusted to pH 7.5. The effects of different pH conditions,
Statistical analysis such as standard errors of means (SE),
i.e. pH 5.0, 6.0, 7.0, 8.0 and 9.0, were also studied over a
two way analysis of variance (ANOVA), etc. was done
culture duration of 10 days. The pH was calibrated to pH
using software GraphPad Prism version: 5.00.
5.0 and 6.0 using an acetate buffer. However, pH 7.0, 8.0
and 9.0 were adjusted with phosphate buffer. All experi-
ments were carried out in triplicate, and the values
Results and discussion
reported are the mean of three such experiments in which
24% variability was observed. Effect of light intensity on growth of Scenedesmus
species ADIITEC-II and GUBIOTJT116
The growth study carried out using different light intensi-
Measurement of microalgae growth ties showed a significant effect on cell growth curve pat-
Growth of microalgae was monitored by counting the cell terns of both of the isolates of Scenedesmus species. The
numbers. Cell numbers were determined every 24 hours species ADIITEC-II and GUBIOTJT116 revealed a signifi-
from a direct cell count using a Neubour Haemocytome- cant effect (p < .0001) on final cell numbers at the end of
ter. For each parameter, the average values were calcu- experiments (Figure 1). Results of the study revealed that
lated from the data generated from three replicates of under low light intensity of 27 mmol m¡2s¡1 both of the
each study. Specific growth rate (m) and doubling time isolate exhibits a slow growth rate and low biomass pro-
(T2) were calculated by using Equations (1) and (2) duction, a gradual increase in irradiance was associated
reported by Levasseur et al. [7]. with an increase in the growth and biomass, but only up to
a certain light intensity, i.e. 81 mmol m¡2s¡1. Among the
m D lnðN2 =N1 Þ=t2 ¡ t1 (1)
studied light regimes, ADIITEC-II exposed to
27 mmol m¡2s¡1 provides a continuous cell growth
T2 D 0:6931=m (2)
through an exponential phase till day 8 of cultivation with
where m is the specific growth rate, T2 is the doubling the least specific growth rate of 0.292 § 0.001 d¡1. Simi-
time, and N1 and N2 is the cell numbers at time t1 and larly, under irradiance of 40.5, 54, 67.5 mmol m¡2s¡1, oth-
time t2, respectively. erwise identical conditions, growth rates were noticed as
0.298 § 0.002 d¡1, 0.296 § 0.002 d¡1 and 0.3 § 0.001
d¡1, respectively. However, further increases in light inten-
Biomass determination sity to 94.5 mmol m¡2s¡1 showed an abrupt drop in growth
Biomass and lipid estimation was carried out by sampling rate (0.17 § 0.0004 d¡1) and the least cell number
at the end of exponential phase of growth. For the analysis (9.6£106 cells ml¡1) towards day 8 of the culture period.
Biofuels 3
Downloaded by [New York University] at 00:56 30 May 2015

Figure 1. Effect of light intensities on growth of Scenedesmus species (ADIITEC-II).

In an independent study reported by Tang et al.[10] and under irradiance of 27, 40.5, 54 mmol m¡2s¡1 showed a
Liu et al.[11] on the growth rate of Dunaliella tertiolecta growth rate of 0.16 § 0.02 d¡1, 0.17 § 0.01 d¡1, and 0.17
and Scenedesmus sp. 11-1 under different light intensity, it § 0.01 d¡1 , respectively (Figure 2). Similarly, a study
was observed that Scenedesmus sp. 11-1 showed high toler- conducted on C. vulgaris has provided different aspects,
ance towards the light regimes, and the light intensity 400 where the low light intensity range of 37.5 to 62.5 mmol
mmol m¡2 s¡1 indicates the most suitable condition for m¡2 s¡1 showed increased growth and biomass yield, and
growth with total biomass yield (i.e. 3.88 g L¡1).[11] How- exhibited a significant decrease in response to high light
ever, in the case of GUBIOTJT116, the growth rate was intensity of 100 mmol m¡2 s¡1.[12] It has also been
continuously supported under the irradiance of 81 mmol observed that an alteration in light intensity affects the bio-
m¡2s¡1 and cell density was 4.69 £ 106 cells ml¡1 mass and biochemical compositions and also exhibits their
towards day 10 of the experiment, with maximum growth range of adaptations in different culture conditions accord-
rate 0.25 § 0.02 d¡1 and minimum doubling time 2.56 § ingly.[12] Moreover, light intensity in nature goes to about
0.01 days. The least increase in cell concentration 2,000 mmol m¡2s¡1 , which may limit the growth, as it is
(3.88 £ 106 cells ml¡1) was observed in a culture incubated beyond the light saturation constants of some algae.[13,11]
under light intensity of 94.5 mmol m¡2s¡1 with minimum Indeed, depending on species, microalgae exhibit increased
growth rate (0.1 § 0.02 d¡1). However, culture grown growth and biomass yield under certain saturation point of

Figure 2. Effect of different light intensity on growth of Scenedesmus species (GUBIOTJT116).


4 A. Difusa et al.

Table 1. Growth characteristics, biomass and total lipid content of Scenedesmus species (ADIITEC-II) under different light intensities.

Light intensity Specific growth Doubling time Biomass productivity Total lipid
(mmol m¡2s¡1) rate m, (day¡1) T2, (days) (g l¡1d¡1) content (gl¡1)

27 0.292 § 0.001 2.37 § 0.01 0.34 § 0.009 0.246 § 0.003


40.5 0.298 § 0.002 2.32 § 0.011 0.35 § 0.024 0.253 § 0.001
54 0.296 §0.002 2.34 § 0.02 0.36 § 0.02 0.241 § 0.002
67.5 0.3 § 0.001 2.31 § 0.01 0.36 § 0.015 0.285 § 0.002
81 0.334 § 0.002 2.07 § 0.011 0.41 § 0.021 0.322 § 0.001
94.5 0.17 § 0.0004 4.06 § 0.01 0.25 § 0.02 0.533 § 0.001

Data presented as § standard error of mean of triplicates (each treatment).

light. However, further increase in light intensity beyond production of 0.25 § 0.02 g l¡1 d¡1 and 0.14 §
the saturation point inhibits the growth and biomass caus- 0.02 g l¡1 d¡1 , respectively, at 94.5 mmol m¡2s¡1. On
ing photo inhibition.[14] In this phenomenon, photo oxida- the other hand, with an increased light intensity lipid yield
tion reaction takes place inside the cell, due to inability of was found to be increased (Figure 5). Compared with low
the photosynthetic apparatus to absorb the excess light. light conditions, the pattern of lipid content was almost
[1518] similar for both the species. In the case of ADIITEC-II,
the low light conditions 27, 40.5 and 54 mmol m¡2 s¡1
Downloaded by [New York University] at 00:56 30 May 2015

showed minimum lipid content, i.e. 0.24 § 0.003, 0.25 §


Effect of light intensity on biomass and lipid content of 0.001, and 0.24 § 0.002 gl¡1, respectively. Likewise,
Scenedesmus species ADIITEC-II and GUBIOTJT116 minimum lipid accumulation was also observed in the
For the growth of microalgae cells, light is the major case of species GUBIOTJT116 under the same light inten-
source of energy. Therefore, light intensity played a sig- sities and thereafter increased beyond the light intensity of
nificant effect on biomass production of the studied spe- 67.5 mmol m¡2 s¡1. Thus, in the studied species, light
cies towards the exponential growth phase. As shown in intensities ranging from 67.5 to 94.5 mmol m¡2 s¡1
Figure 5, each microalga has a unique requirement of light tended to promote the lipid yield. Total lipid content of
intensity for biomass and lipid production. At light inten- the studied microalgae species was determined towards
sity of 81 mmol m¡2s¡1, ADIITEC-II exhibits maximum the late exponential phase during the culture period. As
growth and biomass production (0.41 § 0.021 g l¡1d¡1) noted from the data obtained, significant variation was
compared to GUBIOTJT116 (0.32 § 0.02 g l¡1 d¡1) observed in respond to different light intensities. Light
(Tables 1 and 2). An increase in light intensity to regime carries an essential part in the production of tria-
94.5 mmol m¡2s¡1 lowered the biomass production. For cylglycerides and its composition varies from species to
the same light regime, an effective increase in lipid con- species in respond to different light intensities.[6] There
tent was observed for ADIITEC-II and GUBIOTJT116 are many contradictory reports on accumulation of lipid
(0.53 § 0.001 gl¡1) (0.55 § 0.001) gl¡1, respectively. in relation to irradiance. Many have reported that
Both of the species at series of light regimes (27, 40.5, 54, increased light intensity supports more enhancements of
67.5 mmol m¡2s¡1) follow the steady increase in biomass lipid and hydrocarbon contents rather than biomass pro-
productivity. Therefore, in the present study, it was duction. [19,20] However, high irradiance has been
observed that gradual increases in light intensities from reported to be responsible for promoting increased growth
27 mmol m¡2s¡1 to 81 mmol m¡2s¡1 favored the cell den- but reduced lipid content.[21] It was reported that triacyl-
sity increase which results in maximum biomass produc- glycerol is the main component of neutral lipid and, dur-
tion (Figures 3 and 4). However, the biomass productivity ing synthesis, it requires a large amount of ATP and
of ADIITEC-II and GUBIOTJT116 was limited beyond NADPH via photosynthesis. Therefore, on exposure to
the light regime 81 mmol m¡2s¡1 with minimum biomass large quantities of light energy, excess light energy may

Table 2. Growth characteristics, biomass and total lipid content of Scenedesmus species (GUBIOTJT116) under different light
intensities.

Light intensity Specific growth Doubling time Biomass productivity Total lipid
(mmol m¡2s¡1) rate m, (day¡1) T2, (days) (gl¡1d¡1) content (gl¡1)

27 0.16 § 0.01 3.8 § 0.18 0.2 § 0.01 0.278 § 0.001


40.5 0.17 § 0.002 3.8 § 0.04 0.22 § 0.02 0.274 § 0.002
54 0.17 § 0.002 3.8 § 0.04 0.22 § 0.02 0.285 § 0.001
67.5 0.2 § 0.02 3.0 § 0.37 0.27 § 0.02 0.333 § 0.002
81 0.25 § 0.01 2.56 § 0.13 0.32 § 0.02 0.362 § 0.001
94.5 0.1 § 0.003 5.77 § 0.13 0.14 § 0.02 0.551 § 0.001

Data presented as § standard error of mean of triplicates (each treatment).


Biofuels 5

Figure 3. Relationship between biomass productivity and lipid content of Scenedesmus species (ADIITEC-II) under the light intensity
of 27, 40.5, 54, 67.5, 81, and 94.5 mmol m¡2s¡1.

be directed toward the lipid accumulation on a unit bio- condition, cell numbers of ADIITEC-II and
Downloaded by [New York University] at 00:56 30 May 2015

mass basis through the process of photosynthesis.[11] GUBIOTJT116 reached the maximum value of 7.77 £
Thus, the study describes that modifications of light inten- 106 cells ml¡1 and 4.05 £ 106 cells ml¡1 with maximum
sity in accordance to microalgae species may serve the specific growth rate of 0.3 § 0.02 d¡1 and 0.2 § 0.04 d¡1,
required yield of biomass and lipid. respectively, towards the final day of the experiment. As
illustrated in Table 3, the maximum biomass productivity
of 0.38 § 0.02 gl¡1d¡1 for ADIITEC-II and 0.23 § 0.02 g
Effect of pH on growth and biomass production of l¡1 d¡1 for GUBIOTJT116 was observed at pH 7.0. In
Scenedesmus species ADIITEC-II and GUBIOTJT116 comparison, acidic conditions (pH 5.0 and pH 6.0) do not
Growth of both Scenedesmus species was studied under enhance the cell density and show minimum biomass pro-
controlled pH condition and was controlled by using buf- ductivity. Therefore, a linear decline in the cell density of
fers. Microalgae growth is also sensitive to pH change, both of the isolates was observed towards the acidic con-
and shows an increase trend of pH towards the stationary ditions. However, a steady cell growth with maximum
stage of the growth phase due to their ability to metabolize biomass productivity was observed in ADIITEC-II at pH
the inorganic carbon (CO2).[22] The effects of pH on 8.0 and pH 9.0 with yield of 0.27 § 0.02 gl¡1d¡1 and 0.26
growth of ADIITEC-II and GUBIOTJT116 are shown in § 0.01 gl¡1d¡1, respectively. For GUBIOTJT116 at pH
Figures 6 and 7. From the data obtained, it was observed 8.0 specific growth observed was 0.15 § 0.02 d¡1, but an
that the pH of the media has a significant effect (p < increase in pH to 9.0 slowed the growth rates. Subse-
.0001) on final cell numbers and growth rates of both of quently, the biomass productivity was decreased slightly
the isolates. Among the studied pH range, growth of both (i.e. 0.21 § 0.02 and 0.2 § 0.02 g l¡1 d¡1) during pH 8.0
of the species was enhanced at pH 7.0. Under this and 9.0. Similar response was also observed when the

Figure 4. Relationship between biomass productivity and lipid content of Scenedesmus species (GUBIOTJT116) under the light inten-
sity of 27, 40.5, 54, 67.5, 81, and 94.5 mmol m¡2s¡1.
6 A. Difusa et al.

Figure 5. Total lipid content of Scenedesmus species ADIITEC-II and GUBIOTJT116.


Downloaded by [New York University] at 00:56 30 May 2015

species was grown at lower pH resulting in less cell den- entire nitrate taken up was released as ammonia during
sity and least biomass productivity among the studied pH this condition in the presence of CO2, which might be the
conditions. At pH 5.0 and pH 6.0, the biomass productiv- reason for an abrupt rise in the alkalinity at the later stage
ity was recorded as 0.08 § 0.02 g l¡1d¡1 and 0.1 § of growth.[24] Hence, an intermediate key is required to
0.01 g l¡1d¡1, respectively. According to reported litera- control the fluctuating pH in order to support the continu-
ture, Scenedesmus species WC-1 showed highest cell den- ous growth of the strain.
sity in the buffered cultures pH 7.4 and 9.3 followed by
pH 10.3 unbuffered media condition.[23] However, A. fal-
catus could grow in a wide range of culture pH 5.010.0, Conclusion
but favored maximum growth between the pH range 7.0 The present work demonstrates significant increases in the
to 9.0.[8] The results agree well with the reported litera- growth, biomass and lipid content of Scenedesmus species
ture,[8,23] where a similar pH trend ranging from 7.0 to ADIITEC-II compared to GUBIOTJT116 by using appro-
9.0 provided favorable condition for increased cell num- priate light intensities and pH conditions. Under higher
ber and high growth rate. Usually, the microalgae cultures light intensity, increased lipid content was observed in
provide gradual elevation of the pH conditions towards both the species, but a further increase of the light regime
the final stage of the growth. According to a study con- beyond 81 mmol m¡2 s¡1 resulted in slower rate of growth
ducted by Eisele and Ullrich, pH 8.2 was found to be the and biomass; even though response of both of the isolates
most efficient for rapid nitrate uptake and almost the towards the light regime was similar. Among the studied

Figure 6. Growth curve of Scenedesmus species (ADIITEC-II) in different pH conditions.


Biofuels 7

Figure 7. Growth curve of Scenedesmus species (GUBIOTJT116) in different pH conditions.


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Table 3. Growth characteristics and biomass production of Scenedesmus species (ADIITEC-II) and (GUBIOTJT116) under different
pH conditions.

Scenedesmus species (ADIITEC-II) Scenedesmus species (GUBIOTJT116)

pH Specific growth Doubling time Biomass productivity Specific growth Doubling time Biomass productivity
conditions rate m, (day¡1) T2, (days) (gl¡1d¡1) rate m, (day¡1) T2, (days) (gl¡1d¡1)

5 0.08 § 0.004 8.55 § 0.64 0.1 § 0.02 0.04 § 0.003 13.8 § 0.64 0.08 § 0.02
6 0.06 § 0.004 11.35 § 0.25 0.08 § 0.01 0.05 § 0.004 7.7 § 0.25 0.1 § 0.01
7 0.3 § 0.002 2.16 § 0.005 0.38 § 0.02 0.2 § 0.0004 2.8 § 0.005 0.23 § 0.02
8 0.2 § 0.002 2.88 § 0.03 0.27 § 0.02 0.15 § 0.001 4.07 § 0.03 0.21 § 0.02
9 0.2 § 0.004 3 § 0.04 0.26 § 0.02 0.13 § 0.001 4.6 § 0.04 0.2 § 0.02

Data presented as § standard error of mean of triplicates (each treatment).

species, ADIITEC-II showed the highest biomass produc- [3] Hansen PJ, Lundholm N, Rost B. Growth limitation in
tivity, and GUBIOTJT116 was better in terms of lipid marine red-tide dinoflagellates: effects of pH versus inor-
yield. Therefore, the most suitable range of light intensity ganic carbon availability. Mar Ecol-Prog. Ser. 2007;
334:6371.
must be selected for optimum growth and lipid yield. Also [4] Goldman JC, Azov Y, Riley CB, Dennett MR. The effect
the pH conditions showed significant effects on growth of pH in intensive microalgal cultures. I. Biomass regula-
and biomass of both species, and pH 7.0 to 9.0 was found tion. J Exp. Mar. Biol. Ecol. 1982;57:113.
to be the most optimum condition. Finally, results of the [5] Latasa M. Pigment composition of Heterocapsa sp. and
study indicate that both of the species can be enhanced for Thalassiosira weissflogii growing in batch cultures under
different irradiances. Sci. Mar. 1995;59:2537.
biomass and lipid by triggering the optimum range of light [6] Wahidin S, Idris A, Shaleh Muhamad RS. The influence of
regime and pH conditions. light intensity and photoperiod on the growth and lipid
content of microalgae Nannochloropsis sp, Bioresource
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[7] Levasseur M. Thompson PA, Harrison PJ. Physiological
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No potential conflict of interest was reported by the authors. sources. J. Phycol. 1993;29:587595.
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