Algal Research 10 (2015) 48–54

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Algal Research

journal homepage: www.elsevier.com/locate/algal

Influence of culture medium recycling on the performance of Arthrospira

platensis cultures
Orily Depraetere a,⁎, Guillaume Pierre b, Wim Noppe c, Dries Vandamme a, Imogen Foubert d,e,
Philippe Michaud b, Koenraad Muylaert a
a
KU Leuven Campus Kortrijk, Laboratory Aquatic Biology, E. Sabbelaan 53, 8500 Kortrijk, Belgium
b
Clermont Université, Université Blaise Pascal, Institut Pascal UMR CNRS 6602, 24 avenue des Landais, BP 206, 63174 Aubière cedex, France
c
KU Leuven Campus Kortrijk, IRF Life Sciences, E. Sabbelaan 53, 8500 Kortrijk, Belgium
d
KU Leuven Campus Kortrijk, Research Unit Food & Lipids, Department of Molecular and Microbial Systems Kulak, Etienne Sabbelaan 53, 8500 Kortrijk, Belgium
e
Leuven Food Science and Nutrition Research Centre (LFoRCe), KU Leuven, Kasteelpark Arenberg 20, 3001 Heverlee, Belgium

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

Article history: To reduce the water footprint of microalgae biomass production, it is essential to recycle the culture medium. The
Received 19 December 2014 influence of medium recycling on the performance of the cyanobacterium Arthrospira platensis, the most widely cul-
Received in revised form 23 February 2015 tivated microalgae, was investigated. Arthrospira was harvested with a 20 μm mesh size microstrainer, which is the
Accepted 13 April 2015
benchmark harvesting technology for Arthrospira production. Repeated recycling of the culture medium resulted in
Available online xxxx
a decline in growth rate and the maximum quantum yield of photosynthesis (Fv/Fm) when compared to a control
Keywords:
culture in fresh medium. This decline was accompanied by accumulation of organic matter in the culture medium
Medium reuse (up to 104 mg C L−1). This organic matter consists of 70% of sugars, mostly rhamnose-rich polysaccharides with
Polysaccharides uronic acids. Accumulation of polysaccharides resulted in a decrease in the filtration rate through the microstrainer
Organic matter used for harvesting. Part of the biomass escaped harvesting and was returned to the culture with the recycled me-
Harvest efficiency dium. This resulted in a change in the Arthrospira population and reduction in the harvesting efficiency, but this
Growth inhibition change in population had no effect on the growth rate. The growth rate of Arthrospira in the recycled culture medium
was primarily influenced by organic matter that accumulated in the medium.
© 2015 Elsevier B.V. All rights reserved.

1. Introduction
Microalgae have received a lot of interest in recent years as a new
source of biomass for production of biofuels or other bioproducts [1].
Like other crops, production of microalgae requires water [2,3]. The
biomass concentration of microalgae in open cultivation systems is typi-
cally only about 0.5 g dry biomass L−1. Consequently, 2000 L water is re-
quired to produce 1 kg of dry biomass. This is comparable to rice or
soybeans, two terrestrial crops with a notoriously high water demand.
This water footprint should be taken into account when considering
microalgae as a new source of biomass [4]. In contrast to terrestrial
crops, microalgae do not actively evaporate water through evapotranspi-
ration. Microalgae require water mainly as a medium to remain in sus-
pension, which implies that a large proportion of medium used for
production of microalgae can thus theoretically be re-used, provided
that essential nutrients have been repleted. Water losses are therefore
limited to evaporative losses from the water surface, biomass-bound
water and water used in biomass processing. Recycling of the culture
⁎ Corresponding author.
E-mail address: Orily.Depraetere@kuleuven-kortrijk.be (O. Depraetere).
medium after harvesting can reduce the water demand by 84% [5]. This
results in a water demand of only 320 L kg−1 dry biomass, which is low
compared to most terrestrial crops.
Today, global production of microalgae is still modest (about
10,000 ton year−1) and only a handful of species are produced on a
large scale. The bulk of the total global production can be ascribed to a sin-
gle species, the cyanobacterium Arthrospira platensis, commercially
known as ‘Spirulina’ [6]. Production of Arthrospira is relatively straightfor-
ward compared to other microalgae because contamination of the cul-
tures can easily be avoided by maintaining a high alkalinity. Under
nutrient-replete conditions, Arthrospira has a high protein content and
is attractive as animal feed. When nutrient-stressed, it can accumulate
up to 70% of carbohydrates in the form of glycogen, which makes it an at-
tractive feedstock for production of biofuels [7]. Arthrospira also contains
several high-value biochemical compounds such as the poly-
unsaturated fatty acid gamma-linolenic acid or the natural blue pigment
phycocyanin [8,9]. Because the culture medium contains high concentra-
tions of carbonate/bicarbonate (N8 g L−1), most of which is not consumed
but serves only to maintain a high alkalinity in order to avoid contamina-
tion of the culture, recycling of the medium in Arthrospira cultivation is
not only important to reduce water consumption but also to reduce the
need for chemicals.

http://dx.doi.org/10.1016/j.algal.2015.04.014
2211-9264/© 2015 Elsevier B.V. All rights reserved.
 O. Depraetere et al. / Algal Research 10 (2015) 48–54
Despite the fact that recycling of the culture medium can lead to
massive savings in water and chemical consumption, there have been
relatively few studies that have investigated the feasibility of culture
medium recycling. As far as we know, no studies have investigated
medium recycling in Arthrospira cultures. Nevertheless, it is known
that culture medium recycling causes problems in industrial production
of Arthrospira [10]. In other species of microalgae, some studies found
that recycling of the culture medium had little effect on the perfor-
mance of microalgal cultures (e.g., Chlorella vulgaris, Hadj-Romdhane
et al. [11]; Scenedesmus sp., Kim et al. [12]; Chlorella zofingiensis, Zhu
et al. [13]) while others observed a significant inhibition of growth
after repeated medium recycling (e.g. Nannochloropsis sp., Rodolfi
et al. [14]).
Medium recycling may influence the performance of the culture in
two essentially different ways. First, dissolved substances that are
excreted by the microalgae will accumulate in the medium. Microalgae
can excrete large amounts of dissolved organic matter in the culture
medium [15]. This organic matter may include growth-inhibiting sub-
stances, such as specific free fatty acids (e.g. [16]). A large part of this
organic matter consists of polysaccharides and these may influence
the rheological properties of the culture medium [17,18]. Not only dis-
solved substances but also particulate cell debris can accumulate in
the medium and this may also influence the performance of the culture,
as has been shown for Nannochloropsis [14]. Secondly, because harvest-
ing is never 100% efficient, part of the microalgal population will inevi-
table escape harvesting and will be returned to the culture when the
medium is recycled. This selective force may result in evolution of the
population and this may influence the performance of the culture [19].
One of such changes will most likely be a decrease in the harvesting
efficiency of the biomass, but other properties may change as well, in-
cluding a reduction in growth rate.
The goal of this study was to investigate the influence of repeated
recycling of the culture medium on the performance of an Arthrospira
culture. Therefore, we compared the performance of an Arthrospira
culture in which the culture medium was repeatedly recycled with
the performance of a control culture that received fresh medium. It
was also investigated to what extent changes in culture performance
were due to changes in the Arthrospira population resulting from
incomplete harvesting or due to changes in the medium. Changes in
growth rate, biomass composition as well as harvesting efficiency were
monitored.
2. Materials and methods
2.1. Arthrospira strain and culture conditions
The A. platensis strain 21.99 (SAG, Germany) was used in all experi-
ments (further referred to as Arthrospira). Arthrospira was cultured in
Zarrouk medium modified by Cogne et al. [20]. Cultures were main-
tained in 1 L bottles. The cultures were aerated with 0.2 μm filtered air
(5 L min− 1) and stirred using a magnetic stirrer. The culture bottles
were irradiated from one side with daylight fluorescent tubes, resulting
in a light intensity of 100 μmol photons m−2 s−1 at the surface of the
flasks. The photoperiod was set at 16:8 (light:dark) and room tempera-
ture was held constant at 20 ± 2 °C.
2.2. Culture medium recycling experiment
To evaluate the influence of medium recycling on the performance
of Arthrospira cultures, three replicate sequential batch cultures in
which the medium was replaced weekly with fresh culture medium
(fresh medium treatment, FM) were compared with sequential batch
cultures in which the medium was replaced with recycled culture medi-
um (recycled medium treatment, RM). In the RM treatments, the bio-
mass was harvested using a microstrainer, as this is the standard
harvesting method in commercial production of Arthrospira [21]. A
49
nylon mesh with a pore size of 20 μm was used. Preliminary tests had
shown that this mesh size had a harvesting efficiency (Eq. (1), see
below) of 88%. Nylon meshes with a wider mesh size had a substantially
lower harvesting efficiency (30 μm: 83%, 41 μm: 76%, 64 μm: 64%, and
180 μm: 5%). In the FM treatment, 80% of the culture was removed
and replaced with fresh culture medium. In the RM treatment, 90% of
the culture medium was removed and replaced with recycled culture
medium. Because harvesting was only 88% efficient in the RM treat-
ment, initial biomass concentrations were comparable in both treat-
ments despite the fact that the amount of medium that was replaced
was different. The cultures were restarted every 10 days and this cycle
was repeated 4 times. To avoid nutrient limitation, the main macronu-
trients N and P were repleted in the RM treatment based on the amount
of biomass that was harvested and the content of nutrients in the bio-
mass (10% N and 0.8% P [22]). Previous experiments had shown that
repletion of iron and trace elements was not necessary.
2.3. Biomass analyses
Biomass concentration was monitored by measuring the absorbance
at 750 nm [23]. Spectrophotometric measurements were calibrated by
gravimetrical analysis, after filtration of a known volume on a pre-
weighed GF/C filter. The exponential growth rate was calculated from
changes in biomass concentration over the one-week period following
medium replacement. The maximum quantum yield of photosystem II
or Fv/Fm is a highly sensitive indicator of stress in photosynthetic organ-
isms [24]. We measured Fv/Fm using an AquaPen PAM fluorometer
(Photon Systems Instruments, Czech Republic) Samples were dark-
adapted for 30 min prior to fluorescence measurements. Fm is defined
as the maximum fluorescence, measured after supplying a high intensi-
ty light pulse. F0 is defined as the minimum fluorescence, measured
with minimal irradiance. Fv is then referred to as the variable fluores-
cence (Fm–F0) [25].
At the end of the experiment, total sugars as well as the pigments
phycocyanin, chlorophyll a and carotenoids were measured in the bio-
mass. Proteins were determined by the Bradford assay [26] and sugar
content using the phenol–sulphuric acid method according to Dubois
et al. (1956) with glucose as standard [27]. Carotenoids and total chloro-
phyll in the biomass were measured according to Lichtenthaler and
Buschman [28] and phycocyanin according to Yoshikawa and Belay
[29].
2.4. Culture medium analyses
In the culture medium, the concentration of dissolved organic
carbon (DOC) and sugars was analysed. The culture medium was first
filtered over a Whatman GF/C glass fibre filter. DOC was measured
using a Shimadzu TOC analyser after acidification to pH 2 and sparging
to remove (bi)carbonate. Total sugars in the medium were measured
according to the Dubois' phenol sulphuric acid method [27], using
glucose as the standard. To evaluate the monosaccharide composi-
tion of the sugars excreted in the recycled culture medium after 4
recycles, the organic matter was concentrated using a 3000 Da ultra-
filtration membrane and dialyzed to remove salts. The concentrate
was then freeze-dried before the polysaccharides were hydrolysed
with trifluoroacetic acid. For HPAEC–PAD, monosaccharides were
isocratically separated on a CarboPacTM PA1 column (250 mm) with
18 mM NaOH for 30 min followed by a linear gradient from 0 to 1 M
sodium acetate in 200 mM NaOH for 20 min (Dionex ICS3000 system).
The total protein content of the freeze-dried concentrate was measured
by the Lowry method [30].
2.5. Monitoring of harvesting efficiency
Because straining using the nylon mesh did not retain all Arthrospira
filaments during harvesting, changes in the efficiency of harvesting
50 O. Depraetere et al. / Algal Research 10 (2015) 48–54

were also monitored during the course of the experiment. The har-
vesting efficiency (Eq. (1)) of the microstrainer that was used was cal-
culated as follows:
Harvesting efficiency ð%Þ ¼ ðODb −ODa Þ=ODb  100
with ODb and ODa being the optical density (750 nm) of the culture
before (ODb) and after (ODa) harvesting.
The filterability of the Arthrospira medium was analysed after
removing the cells by filtration over a Whatman GF/C glass fibre filter
ð1Þ with 1.2 μm pore diameter and by measuring the filtration flux of this

Fig. 1. The OD (750 nm), growth rate (day−1) and quantum yield of Arthrospira platensis and dissolved organic carbon (DOC) (mg·L−1) and total sugars concentration (mg·L−1) in the
fresh and recycled medium.
 O. Depraetere et al. / Algal Research 10 (2015) 48–54
Table 1
Biomass composition of Arthrospira platensis harvested from recycled medium (RM) and
fresh medium (FM) treatments.
RM FM
Ash (%) 6.2 ± 2.6 4.0 ± 0.3
Total sugars (%) 24.8 ± 3.6 7.7 ± 1.2
Protein (%) 26.7 ± 2.6 35.8 ± 1.6
Phycocyanin (%) 6.8 ± 2.3 15.4 ± 0.8
Total chlorophyll (%) 0.8 ± 0.1 1.8 ± 0.1
Carotenoids (%) 0.5 ± 0.1 0.5 ± 0.1
cell-free medium through a 25 mm GF/F glass fibre filter with 0.7 μm
pore diameter.
2.6. Comparing the influence of changes in the Arthrospira population and
changes in the medium on culture performance
A reduction in growth rate in the RM treatment compared to FM
treatment could on the one hand be due to an inhibitory effect of the
recycled culture medium, or on the other hand due to a change in the
Arthrospira population resulting from incomplete harvesting. As a result
of incomplete harvesting, part of the Arthrospira population is returned
to the culture with the recycled medium and this may result in a change
in the composition of the population. Therefore another experiment
was carried out to evaluate whether the reduction in growth rate was
due to the accumulation of organic matter in the medium or due to a
change in the Arthrospira population. This experiment was a repetition
of the medium recycle experiment except for the fact that harvesting
was done every 7 days instead of every 10 days. During the third recycle,
two additional treatments in addition to the original treatments were
added: a treatment in which the Arthrospira biomass from the RM
treatment was used as inoculum (recycled inoculum treatment, RI) in
fresh culture medium (treatment: FM + RI) and a treatment in which
Arthrospira biomass from the culture with fresh medium was used as in-
oculum (fresh inoculum, FI) in recycled culture medium (treatment:
RM + FI). Like in the first experiment, the growth rate of Arthrospira
was monitored throughout the experiment.
2.7. Influence of the polysaccharides in the culture medium on the growth of
Arthrospira
In the last experiment, it was tested whether the polysaccharides in
the culture medium inhibited the growth of Arthrospira. The growth of
Arthrospira was compared in control medium to medium to which a
polysaccharide concentrate was added. This polysaccharide concentrate
was isolated from the recycled culture medium using a 3000 Da ultrafil-
tration membrane. The concentrate was added to achieve the same
Fig. 2. Arthrospira population in the culture before harvesting (left) and in the medium after harvesting using a 20 μm microstrainer (right), the scale bar corresponds to 500 μm.
51
concentration as in the recycled medium (final concentration of
160 mg L− 1 total sugars).
p-value 2.8. Statistical analyses
N0.05
b0.001 Statistical analyses were performed using Sigma-plot 11 (Systat
b0.05 Software, Inc.). Before evaluating the results with one-way analysis of
b0.05
variance (ANOVA), normality of the data was determined with the
b0.05
N0.05 Shapiro–Wilk normality test. To analyse the pairwise differences, a
Tukey's post-hoc test was used. The significance level of statistical anal-
yses was set at p = 0.05.
3. Results and discussion
To study the effect of medium recycling on the performance of
Arthrospira cultures, three replicate semi-continuous cultures in which
the medium was replaced with fresh medium (FM) were compared
with three replicate cultures in which the medium was recycled (RM)
(Fig. 1). Both cultures performed equally well at the start of the experi-
ment. With an increasing number of medium recycles, however, the
growth rate declined in the RM treatment while it remained stable in
the FM treatment. The exponential growth rate of Arthrospira in the
FM treatment varied between 0.18 and 0.26 day−1 between different
batch cycles. In the RM treatment, the growth rate decreased from
0.24 day−1 at the start of the experiment to only 0.07 day−1 after four
recycles. The growth rate of Arthrospira in RM treatment was signifi-
cantly lower than in the FM treatment from the third recycle onwards.
As concerns the biomass composition, the protein, phycocyanin and
total chlorophyll content were lower in the biomass from the RM than
the biomass from the FM treatment while the total sugar content was
higher (Table 1). The shift from protein production to accumulation of
sugars is a typical response of Arthrospira to stress conditions, such as
nutrient stress or salt stress (e.g. [31,32]). In a recent study on medium
recycling in Scenedesmus cultures, Rocha and co-workers also observed
a similar change in the protein/carbohydrate ratio after several medium
recycles, even though growth rate was not affected by the medium
recycling [33]. The fact that the Arthrospira culture from the RM treat-
ment was under stress was confirmed by measurements of the maxi-
mum quantum yield of photosystem II or Fv/Fm, which declined
significantly in the RM treatment after the third recycle. Fv/Fm was 2
times lower in the RM than in the FM treatment after 4 recycles. Fv/Fm
is a sensitive indicator of stress in microalgae, including stress induced
by nutrient limitation, high irradiance or toxic compounds (e.g. [25,
34]). It is a parameter that is easy to measure using low-cost equipment.
It has therefore been recommended as a tool to monitor lipid accumula-
tion induced by nutrient stress in large-scale cultures [35]. It may also be
used to detect declines in culture performance as a result of medium
recycling.
52 O. Depraetere et al. / Algal Research 10 (2015) 48–54
Table 2
Monosaccharide composition of the high-molecular weight fraction (N3000 Da) of
the organic matter in the recycled medium of Arthrospira platensis after 4 recycles
analysed with High Pressure Anion Exchange Chromatography with Pulsed Ampero-
metric Detection (HPAEC–PAD).
Monosaccharide composition (w/w) HPAEC–PAD
Rhamnose (%) 55.1
Glucose (%) 8.8
Xylose (%) 10.12
Galactose (%) 4.11
Arabinose (%) 2.79
Glucuronic acid (%) 7.33
Galacturonic acid (%) 5.57
Fructose (%) 6.16
In the RM treatments, Arthrospira filaments were harvested by a
20 μm mesh size microstrainer, which is the most common harvesting
method used in the industry [10]. In the RM treatment, the harvesting
efficiency declined slowly but significantly from 90 ± 1% after the first
recycle, to only 68 ± 1% after the fourth recycle. Arthrospira that were
not retained by the filter were returned to the culture when the medium
was recycled. This resulted in a change in the population of Arthrospira
in the culture. Microscopic analysis indicated that small filaments
escaped harvesting (Fig. 2). This resulted in an increase in dominance
of small filaments in the culture and a decrease in the harvesting effi-
ciency in the next round of harvesting. Belay et al. [10] noted this prob-
lem when straining is used for harvesting Arthrospira at an industrial
scale.
Repeated recycling of the culture medium resulted in accumulation
of relatively high concentrations of dissolved organic matter and
sugars in the medium. In the FM treatment, the concentrations of
DOC remained constant at around 50 mg L − 1 (Fig. 1). In the RM
treatment, on the contrary, DOC concentration increased to about
100 mg L− 1 towards the end of the experiment. Total sugars were
constant at about 50 mg L− 1 in the FM treatment and increased up
to 180 mg L− 1 total sugars in the RM treatment. Assuming a carbon
content of 40% for sugars, about 70% of the DOC at the end of the
experiment consisted of sugars. An additional 12% were proteins.
The remaining 18% may be other organic compounds, for example:
free fatty acids in culture medium of Monodus subterraneus [16]. It is
known that microalgae excrete large amounts of organic matter in the
Fig. 3. The growth rate (day−1) of the fresh and recycled medium Arthrospira platensis
culture during the recycle experiment with RM = recycled medium, FM = fresh medium,
RI = recycled inoculum and FI = fresh inoculum. a,b,cDifferent letters mean significantly
different.
Fig. 4. The OD (750 nm) of Arthrospira platensis in control medium with and without
polysaccharide concentrate.
culture medium, often 10–20% of the biomass production [15,36]. This
organic matter accumulates in the medium when the medium is
recycled. Other studies have also reported excretion of organic matter
in the culture medium of Arthrospira. Trabelsi et al. [37], for instance,
measured a concentration of 220 mg L−1 in an Arthrospira culture con-
taining only 510 mg L−1 dry weight biomass.
The organic matter excreted in the recycled culture medium was
concentrated using a 3000 Da ultrafiltration membrane for further anal-
ysis of the excreted sugars. Analysis of the sugar content of the filtrate
indicated that the membrane retained more than 99% of the total sugars
in the recycled medium. The organic matter of the recycled medium
retained by the 3000 Da ultrafiltration membrane contained 12% pro-
teins, suggesting that part of the unidentified organic matter that accu-
mulated in the recycled medium consisted of proteins. In the fresh
medium treatment, less than 1% of the sugars were retained by the
3000 Da ultrafiltration membrane. This indicates that the sugars that
accumulated in the recycled medium consisted mostly of large polysac-
charides, while the few sugars in the fresh medium treatment were
small polysaccharides or monosaccharides. The monosaccharide com-
position in the recycled medium was analysed using HPAEC–PAD anal-
ysis (Table 2 and Supplementary Fig. S1). The dominant sugar monomer
was rhamnose, which represented 55% of the total monomers. Minor
(5–10%) sugar monomers were glucose, xylose, galactose and fructose
and the uronic acids, glucuronic and galacturonic acid. Previous studies
have also reported rhamnose as the major sugar monomer in excreted
polysaccharides in the culture medium of Arthrospira [37–39].
Filali Mouhim et al. [38] noted that extracellular polysaccharides
from Arthrospira significantly increase the viscosity of the medium at
concentrations of 50–100 mg L−1, which is similar to the concentrations
measured in our experiment. During harvesting of Arthrospira in the
recycled medium treatment, a reduction in the filtration rate during
straining was noted after repeated recycling of the culture medium. To
evaluate whether the filterability of the culture medium had decreased,
the filtration flux of fresh medium was compared with recycled medi-
um (collected after 4 recycles) by filtration over a standard GF/F filter.
The filtration flux of fresh culture medium was 1.15 mL s− 1 but this
decreased to 0.29 mL s−1 for recycled medium with a concentration of
137 mg L−1 total sugars. This suggests that polysaccharides that accu-
mulate in the culture medium result in a reduced filterability of the
medium and thus complicate harvesting by means of straining.
Repeated recycling of the culture medium clearly caused a decrease
in the performance of the Arthrospira culture. This may be due to a
change in the composition of the medium caused by accumulation
of excreted organic matter or due to a change in the population of
 O. Depraetere et al. / Algal Research 10 (2015) 48–54
Arthrospira due to incomplete harvesting by the straining method. To
discriminate between these two causes, an additional experiment was
carried out in which in an Arthrospira population from the RM treat-
ment was transferred to fresh medium and, vice versa, a population
from the FM treatment was transferred to cell-free medium from the
RM treatment (Fig. 3). When an Arthrospira population that had gone
through repeated cycles of harvesting was transferred into fresh medi-
um (FM + RI treatment), it grew just as well as an Arthrospira popula-
tion that had never been subjected to medium recycling (FM + FI
treatment). When Arthrospira filaments that had been raised in fresh
culture medium were transferred to the recycled medium (treatment
RM + FI), however, the growth was significantly reduced, although
not as much as in the RM + RI treatment. This suggests that the reduc-
tion in culture performance was due to changes in the medium compo-
sition and not to changes in the Arthrospira population resulting from
incomplete harvesting.
Because the recycled culture medium contained high concentrations
of excreted polysaccharides, it was tested whether these polysaccha-
rides inhibited the growth of Arthrospira (Fig. 4). The addition of this
polysaccharide concentrate to the medium resulted in a lower growth
rate, but this effect was not significant and was smaller than the
reduction in growth observed in a culture after the medium had been
recycled several times. This suggests that polysaccharides may play a
role in the reduction in growth rate in a culture with recycled culture
medium, but other factors also play a role. Inhibition may be caused
by smaller molecules such as free fatty acids, as has been observed in
Monodus cultures [16].
Our experiment demonstrates that repeated recycling of the culture
medium causes two important problems in Arthrospira cultures. On the
one hand, incomplete harvesting results in a change in the Arthrospira
population, with a larger proportion of the population that escapes
harvesting by straining. This causes a decrease in the harvesting effi-
ciency but has no effect on the growth rate of the Arthrospira population.
On the other hand, Arthrospira excretes large amounts of organic matter
and this accumulates in the culture medium when it is recycled. A large
part of this organic matter consists of polysaccharides. These cause an
increase in the viscosity of the medium and this in turn reduces the
filtration rate when the biomass is harvested by straining. Some organic
compounds that accumulate in the culture medium appear to influence
the performance of the Arthrospira culture, as is apparent from the
lower growth rate, a reduction in the quantum yield of photosystem II
and a change in the biomass composition.
Supplementary data to this article can be found online at http://dx.
doi.org/10.1016/j.algal.2015.04.014.
Acknowledgements
The research presented in this paper was financially supported
by the Agency for Innovation by Science and Technology (IWT strategic
research grant O. Depraetere number 111275) and the Research Foun-
dation Flanders (FWO travel grant O. Depraetere number V431813N).
D. Vandamme is a postdoctoral researcher funded by the Research
Foundation Flanders (FWO grant D. Vandamme number 12D8914N).
Thanks to Laura Van Maele and Sanjaya Lama for their help in the lab
and to Prof. Christophe Vial, Dr. Cédric Delattre, Dr. Alina-Violeta Ursu,
Fatou Ba, Christine Gardarin and Jo Tixeuil for their assistance at the
Blaise Pascal University.
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