Journalof Applied Phycology 8: 131-137, 1996.

131
1996 Kluwer Academic Publishers. Printed in Belgium.

Effect of blue-green light on growth rate and chemical composition of three

diatoms
Maria del Pilar Sdnchez-Saavedral & Domenico Voltolina2
1Departamentode Acuicultura, Centro de Investigaci6n Cientifica y de Educaci6n Superiorde Ensenada,
ApartadoPostal 2732, Ensenada, 22800 Baja California, Mexico
2
Present address: UniversidadAuton6ma de Sinaloa, Facultadde Ciencias del Mar, Apartado Postal 610,
Mazatlan, 82000 Sinaloa, Mexico

Received 12 March 1996; revised 25 May 1996; accepted 28 May 1996

Key words: diatom cultures, light quality, irradiance, growth rate, chemical composition, Chaetoceros sp., Skelet-
onema costatum, Thalassiosirapseudonana

Abstract

The diatoms Chaetoceros sp., Skeletonema costatum and Thalassiosira pseudonana were grown with different
irradiances of white and of blue-green light, and with a mixture of blue-green plus 6.5 mol m -2 s- of white light.
Exponential growth rates were higher in mixed blue for the first two, while T pseudonanagrew faster in white light
but, in all cases, mean cell division rates did not differ with increasing irradiances. Harvesting in stationary, rather
than in late exponential growth phase, resulted in higher protein contents for Chaetoceros sp. and S. costatum, but
for T pseudonanathe highest value was in the exponential phase. The highest protein content was in blue-green
light for the three species and it increased with irradiance. As to other fractions, the three strains showed different
responses, related to light quality and quantity, as well as to culture ages.

Introduction However, only a few authors (Flaak & Epifanio,

The culture of microalgae as food for the commercial
rearing of marine animals is of critical importance in
mariculture. Apart from the dietary requirements of
the organisms feeding on them, the nutritional value of
microalgae depends on their biochemical composition
(Brown et al., 1989), which may be manipulated by
modifying culture conditions.
Light is the driving force of photosynthesis. As
such, its quantity, its spectral composition and their
fluctuations within the culture control biomass pro-
duction rates (Dubinsky et al., 1995) and may cause
variations in the photosynthetic responses, growth rates
and cell metabolism of algae (Senger, 1987; Sukenik
et al., 1989). For instance, more amino acids and pro-
tein may be found in algal biomass produced under
blue light than under equal levels of white light, while
red light has been shown to favour glycogenesis over
proteogenesis (Voskresenskaya, 1972).
1978; Snchez-Saavedra & Voltolina, 1994, 1995)
have explored the possibility of obtaining live food
of different qualities for filter feeders using large-scale
cultures of one microalga under light with different
spectral characteristics. However, the results obtained
in these previous works differed considerably, indicat-
ing the possibility of a species-specific response to
light quality. In the present paper we compare the
growth rates and the proximate composition of the
same two diatoms (Thalassiosirapseudonana Hasle
and Heimdal and Chaetoceros sp., respectively) and
of a third one Skeletonema costatum (Greville) Cleve
grown to their late exponential and early stationary
growth phases with different photon fluence rates of
white and of blue-green light.
132

Materials and methods 1.2 l i . .

l .
l . . .
I
l l . .
l .
l . . . . . . . . . . . . . .
...
I

1.0
The marine diatoms used for this work were the
clones CH-X-1 (Chaetoceros sp.), SK-C-2 (Skelet- 0.8
onema costatum = CCMP1332), and TH-P-1 (Thalas- r
siosirapseudonana = CCMP1335), of the Centro de Er 0.6
Investigaci6n Cientffica y de Educaci6n Superior de
Ensenada (C.I.C.E.S.E.) culture collection (Trujillo- 0.4
Valle, 1993). The first is a local strain, isolated in
coastal waters and used in several Mexican commer- 0.2
.. ..... ..............
cial hatcheries, the other two were obtained from -----
. . ------
.. ..... . ....::::.
Provasoli-Guillard Center for Culture Collection for V.LU .. . . . . . . .. . . . . .
I E.
I .
F .
I .
i .
I . . . I . . . . . .i-r I

300 400 500 600 700

Marine Phytoplankton (Andersen et al., 1991).
Wavelength
Non-axenic batch cultures of each strain were run
in triplicate, in 250 ml Erlenmeyer flasks with Guillard
2 . l . ll.. . *,I . . .. . . .rll l. ,,l ,, ,,,l
& Ryther's (1962) 'f' medium, prepared with sterile
I.
natural seawater (salinity = 33 1 g kg-'). Cultures
conditions were 23 1 C and pH between 7.5 and
1:3 o b
8.5. Stirring was manual, twice daily, and lighting was ' 0.1
continuous, provided by commercial fluorescent lamps
Sylvania Cool White F40CW or General Electric Blue- 0.
green F40B (peak of emission = 440 nm) with known 4
spectral characteristics (Figure 1) (Sdnchez-Saavedra TO. 4
& Voltolina, 1994). In the case of blue-green, one set of
cultures was isolated from ambient light and a second 0.;
one was allowed to receive 6.5 umol m- 2 s - I of the
1O _. . ., . .I . ,.. . I.........., . I
laboratory lighting, with the same spectral characterist- v
300 400 500 600 700
ics as the experimental white light. In this case, the dis-
Wavelength
tance of the cultures from the light sources was adjus-
ted so that the total irradiance, measured at the centre Figure 1. Spectral distribution (in relative units) of the three fluor-
of the culture vessel with a Biospherical Instrument escent light sources: --- blue light, - mixed blue light; ......
ambient light; b: white light.
QSL-100 quanta meter, was the same as those of white
and blue-green light. Growth experiments were run in
light gradients, from 100 to 547 /mol m - 2 s-l for both
types of blue-green, and from 100 to 718/ mol m-2 s- Proximate cell compositions were determined for
for white light, and were repeated at least three times each culture in late exponential and in stationary phase
for each value irradiance. of growth in three or four triplicate sets of samples for
The growth caracteristics obtained with these each type of analysis and determined independently
experiments for each strain and irradiance were used in each case: soluble protein content, using bovine
in separate sets of experiments, to obtain samples for albumin (98%) as standard, was determined according
proximate analysis during the late exponential and the to Lowry et al. (1951) and Malara & Charra (1972a)
stationary phases of growth, 24 h before and 24 h after after pretreatment for 10 min with 0.1 N NaOH at
reaching the maximum value of cell concentrations at 100 C for Chaetoceros sp. and at 80 C for S. cost-
100 and 400 iumol m- 2 s -1 of irradiance. atum and with 0.5 N NaOH at 80 C for 10 min for
Each set of cultures was started with inocula T pseudonana, adjusting the final pH value to <8.0
acclimated to the light condition tested, for at least (Firber-Lorda, 1986). Carbohydrates were measured
eight to ten generations. Increases in cell concentra- with the phenol-sulphuric method by Dubois et al.
tions were checked by daily measurements of optical (1956), using glucose (99%) as standard, as described
density at 550 nm and by occasional hemacytometer by Malara & Charra (1972b), pretreatment was as in
count. Mean cell division rates were calculated as in Whyte (1987). Lipids were extracted following Bligh
Fogg & Thake (1987), using log2 transformation. & Dyer (1959) and Chiaverini (1972) and quantified

with the colorimetric method by Pande et al. (1963)
using tripalmitin (99%) as standard. All calibrations
curves were routinely checked once weekly with new
standards.
A two-way analysis of variance and the aposteriori
Student Newman Keuls (SNK) test (Sokal & Rohlf,
1979), all for a = 0.05, were used to compare the
effect of each variable.
Results
In all cases, mean cell division rates, from inoculum to
the end of exponential growth, did not differ signific-
antly with increasing irradiance. However, the three
species showed different responses to light quality.
Chaetoceros sp. and S. costatum grew faster in mixed
blue-green than in blue-green light only, and the slow-
est growth was in white light. This was, on the other
end, the best growth-promoting type of light, followed
by mixed blue, for T pseudonana.As to cell numbers,
Chaetocerossp. and T pseudonanahad the highest cell
densities in white light, followed by mixed blue-green,
while S. costatum showed an inverse trend, with the
highest and lowest cell concentrations in blue-green
and in white light, respectively (Tables 1 and 2).
As to proximate composition, responses to light
quality and quantity, as well as to culture age, differed
for each species so that, for the sake of simplicity, the
effect of each variable, summarized in Tables 3 and 4,
will be described separately.
The effects of culture age on Chaetoceros sp. and
S. costatum was a consistent tendency, independent
from light quality and quantity, to higher protein and
lipid contents during the early stationary than in the
exponential growth phase. Carbohydrates, as well as
the ash content had an inverse trend.
On the other hand, upon reaching the stationary
phase, T: pseudonana showed a sharp decrease in
proteins, paralleled by equally sharp increases in all
other fractions in blue and mixed blue-green light. In
white light, only the ash content increased while car-
bohydrates and lipids remained practically unchanged.
As a general response to irradiance, protein con-
tents increased with increasing photon flux density, but
differences were significant only during exponential
growth and simply indicative of this trend in the sta-
tionary phase. As to other organic fractions, the three
strains showed different responses, in some cases also
related to light quality: Chaetocerossp., for instance,
showed an irradiance-related decrease in lipids while
133
carbohydrates remained practically unchanged. The
same tendency was evident for S. costatum in the
case of lipids, although differences never reached the
level of significance. On the other hand, carbohydrates
decreased with irradiance in the case of cultures grow-
ing exponentially in blue-green and mixed blue-green
light, but had an inverse tendency in white light.
In the case of T pseudonana,lipids increased with
irradiance in white and mixed blue-green light for cul-
tures in exponential growth, but not in blue-green light
only. In the stationary phase there was a tendency to
lower carbohydrates with high light, although the dif-
ference was not significant in most of the cases.
The response to light quality is complicated by the
responses to age and irradiance already described. In
general terms, short-wave radiation causes an increase
in proteins, but the presence of a small percentage of
white light seems to have an effect which is not appar-
ently commensurate to such a small amount of con-
tamination by other wavelengths. In all cases, cultures
grown in mixed blue-green have a higher protein con-
tent than those grown in other types of light of equal
photon fluence rate, those in blue-green are second
and white light-grown cultures have the lowest protein
content.
An even more striking effect of light type is the low
amount of carbohydrates of mixed blue-green cultures
of Chaetoceros, which was between one third and 50%
of the levels found with the other types of light.
Discussion
Our results seem to disagree on a number of points
from some of the common generalizations on microal-
gae. For instance, the protein content is expected to
be higher during exponential, rather than in station-
ary phase of growth. Two out of the three species we
tested did not fulfil this expectation, possibly due ot the
late conversion of their nitrogen products into soluble
protein (Fernandez-Refriz et al., 1989).
Another common assumption is that, if the growth
rate of one microalga is not dependent from the exper-
imental variable, the biomass harvested will have a
similar composition. Even if the yields and the growth
rates of our strains were found to be independent from
irradiance for each type of light, we found signific-
ant irradiance-related differences in chemical compos-
ition, during both phases of growth.
Evidently, this is due to changes in biochemical
pathways induced by quantitative changes of the light
134
Table 1. Mean cell division rate () and standard deviation (in brackets) during exponential growth of three
diatoms cultured with different photon fluence rates of blue-green (A), blue-green plus 6.5 timol m- 2 s-l
of white (B) and white light (C). Equal letters indicate lack of significant differences (Two-way ANOVA and
Student-Newman-Keuls a posteriori test, a = 0.05): a<b<c.

Species Light Photon fluence rates (mol s-l m- 2 )

100 210 362 547 718

Chaetoceros sp. A 1.39(0.08)b 1.39(0.03)b 1.38(0.07)b 1.40(0.03)b -

B 2.17(0.02)c 2.15(0.04)c 1.96(0.05)c 2.63(0.07)c -
C 0.73(0.04)a 0.75(0.07)a 0.70(0.05)a 0.71(0.07)a 0.74(0.52)a

S. costatum A 1.02(0.01)b 1.03(0.01)b 1.03(0.01)b 1.03(0.01)b -

B 2.59(0.04)c 2.65(0.02)c 2.59(0.02)c 2.61(0.01)c -
C 0.95(0.03)a 1.15(0.06)b 0.97(0.06)b 0.97(0.04)a 1.07(0.04)b

T pseudonana A 1.95(0.04)b 1.83(0.03)a 1.94(0.03)b 1.96(0.03)b -

B 2.53(0.02)d 2.35(0.04)c 2.03(0.04)ab 2.27(0.04)c -
C 2.74(0.06)e 2.97(0.05)f 2.76(0.03)e 3.23(0.09)9 2.88(0.06)f

Table 2. Mean cell concentrations (cell 103 ml- 1) and standard deviation (in brackets) during exponential
growth of three diatoms cultured with different photon fluence rates of blue-green (A), blue-green plus
6.5 mol m- 2 s- of white (B) and white light (C). Equal letters indicate lack of significant differences
(Two-way ANOVA and Student-Newman-Keuls a posteriori test, a = 0.05): a<b<c.

Species Light Photon fluence rates (mol s-l m-2 )

100 210 362 547 718

Chaetocerossp. A 988(45)a 970(36)a 943(35)a 980(52)a -

B 1027(43)b 1129(26)b 1140(90)b 1040(52)b -
C 1340(29)c 1293(48)c 1322(53)c 1293(96)c 1323(90)c

S. costatum A 940(26)c 923(18)c 946(30)c 960(21)c -

B 790(45)b 810(25)b 797(36)b 820(42)b -
C 520(17)a 536(19)a 517(23)a 508(29)a 523(25)a

T pseudonana A 1248(96)a 1198(87)a 1240(83)a 1190(73)a -

B 1325(81)b 1420(76)b 1420(76)b 1340(36)b -
C 1536(99)c 1620(80)c 1551(67)c 1730(91)c 1630(86)c

environment, not unlike the one described by Helle-
bust (1970) for two marine coccolithophorids, whose
growth was saturated at light intensities far below those
necessary for maximum photosynthesis.
As to light quality, several reports in the literat-
ure suggest that the response of photosynthetic organ-
isms to blue light are by no means homogeneous and
that the expected higher growth rates, higher protein
and chlorophyll contents and increase of tricarboxilic
acid synthesis independently from the photosynthetic
activity is a composite picture, from which individual
species may deviate in one or more aspects (Wallen &
Geen, 1971; Gostan et al., 1986).
In this work we found that the differences measured
in growth rate and cell numbers for the three species
are due only to light quality, and that none was limited
in these aspects by the quantity of light. In two spe-
cies (Chaetoceros sp. and S. costatum) the response
was close to the expected since, as described by oth-
er authors (Hauschild et al., 1991; Ojala, 1993), blue
light caused an increase in growth rate but, in both
cases, only if this was enriched by a small amount of
white. We found a different response to blue light for
T pseudonana, similar to that described by Flaak &
Epifanio (1978) for the same species, since its growth
rate was higher in white than in blue light.
As to biochemical composition, blue light had the
expected effect of higher protein contents for the three
species, again made more evident by the mixture with
white light and by an increase with increased irradiance
 135

Table 3. Mean proximate composition (in percentage of the total dry weight) and standard deviation (in brackets)
during exponential growth of three diatoms cultured with different photon fluence rates of blue-green (A), blue-green
-2 -
plus 6.5 mol m s of white (B) and white light (C). Equal letters indicate lack of significant differences (Two-way
ANOVA and Student-Newman-Keuls aposterioHtest, a = 0.05): a<b<c.

Species Light /mol s- m- 2 Protein Lipids Carbohydrates Ash content

Chaetocerossp. A 100 33.26(1.49)b 20.12(2.16)d 15.26(0.56)c 26.13(2.01)a

400 42.26(1.16)c 10.15(0.41)a 16.26(0.76)c 23.16(2.01)a
B 100 38.12(1.29)b 24.12(0.87)f 7.23(0.89)a 30.41(3.06)a
400 51.26(2.15)d 16.44(0.37)c 6.43(0.39)a 26.33(1.22)a
C 100 30.15(1.46)a 22.41(0.49)e 12.23(0.76)b 26.84(1.36)a
400 36.18(1.37)b 12.44(0.62)b 13.29(0.47)b 23.22(1.44)a

S. costatum A 100 39.16(0.97)a 6.49(0.83)ab 19.21(0.77)d 39.36(1.67)a

400 43.16(0.87)c 5.43(0.46)a 17.21(0.69)c 38.44(1.39)a
B 100 42.17(0.94)c 7.43(0.92)ab 15.23(0.33)b 37.43(1.76)a
400 47.15(1.32)d 5.20(0.72)a 13.26(0.23)a 34.17(0.47)a
C 100 37.23(1.46)a 6.62(0.46)ab 21.73(0.49)e 38.43(1.69)a
400 39.26(0.87)b 4.17(0.37)a 24.33(0.71)f 36.11(1.72)a

T pseudonana A 100 44.12(0.69)c 15.33(0.72)b 12.33(0.43)c 22.47(1.99)a

400 48.27(0.33)e 17.27(0.69)d 11.28(0.23)b 24.72(2.06)a
B 100 46.47(0.47)d 13.23(0.47)a 10.11(0.62)a 26.41(2.31)a
400 52.21(1.69)f 16.44(0.33)c 12.44(0.39)c 23.12(1.76)a
C 100 31.27(0.42)a 20.12(0.12)e 13.62(0.42)d 21.33(1.49)a
400 36.47(1.33)b 20.41(0.36)e 15.23(0.26)e 20.88(1.20)a

Table 4. Mean proximate composition (in percentage of the total dry weight) and standard deviation (in brackets)
during stationary phase of growth of three diatoms cultured with different photon fluence rates of blue-green (A),
-2
blue-green plus 6.5 /mol m s- of white (B) and white light (C). Equal letters indicate lack of significant differences
(Two-way ANOVA and Student-Newman-Keuls aposterioritest, a = 0.05): a<b<c.

Species Light pmol s- m-2 Protein Lipids Carbohydrates Ash content

Chaetocerossp. A 100 37.13(1.22)b 21.46(1.16)d 12.29(0.55)c 21.12(1.41)a

400 45.26(0.96)c 12.13(1.11)a 13.33(0.67)c 19.47(2.47)a
B 100 41.33(0.99)b 25.37(0.96)f 5.33(0.96)a 26.11(1.13)a
400 53.46(1.21)d 17.61(0.68)c 4.37(0.93)a 22.17(1.72)a
C 100 33.29(0.97)a 23.17(0.99)e 10.29(0.62)b 24.29(1.62)a
400 39.18(0.86)b 14.14(0.84)b 11.43(0.72)b 20.17(1.17)a

S. costatum A 100 42.17(1.12)b 8.45(0.62)a 14.33(0.77)b 36.21(0.89)b

400 45.23(1.43)b 7.36(0.72)a 12.11(0.89)b 35.43(0.72)b
B 100 44.27(1.29)b 9.36(0.61)a 12.21(0.92)a 35.21(0.99)b
400 49.86(0.43)c 7.17(0.66)a 10.40(0.96)a 32.11(0.69)a
C 100 39.21(0.26)a 8.23(1.43)a 17.11(0.87)c 35.11(0.77)b
400 41.53(0.23)b 6.14(1.17)a 18.24(0.67)c 32.11(0.74)a

T pseudonana A 100 20.16(0.44)a 22.33(0.79)b 26.18(0.74)c 36.14(0.72)a

400 24.45(0.47)b 21.46(0.87)b 24.12(0.89)b 35.26(0.72)a
B 100 37.93(0.72)c 19.36(0.63)a 17.33(0.99)a 39.12(0.74)b
400 38.43(0.86)c 18.73(0.49)a 16.44(0.68)a 38.41 (0.82)b
C 100 18.61(1.12)a 20.12(0.72)b 16.93(0.77)a 38.10(0.96)b
400 19.61(1.17)a 22.21(0.86)b 15.76(0.74)a 38.44(0.94)b
136
levels. From these discrepancies between our results
and those of the general picture, it is clear that the blue
light effect is not homogeneous, since different species
may use different carbon sinks in dependence from the
particular light environment. As mentioned by other
authors, this may be used to obtain microalgae bio-
mass of different composition by simply changing light
quality and intensity or the time of harvesting. For this
reason, it is necessary to obtain information about the
particular response of each species to different culture
conditions, since the results may be modified consid-
erably by a slight change of the experimental envir-
onment. The potential of these light-induced changes
has not been suficiently explored for aquaculture, espe-
cially in the light of the possibility of obtaining several
types of diets, suitable for feeding different organisms
with different dietary requirements, or to use in mixed
diets in substitution of those presently used, which are
obtained by culturing different microalgae species.
We have shown that, besides harvesting at differ-
ent phases of growth, the chemical composition of
microalgae and hence their nutritional value for filter
feeders, may be varied by modifying the quality and
quantity of light. We have also shown that the results
may change with slight modifications of the experi-
mental conditions since, for equal photon flux densities
of predominantly blue-green light, we obtained signi-
ficant biochemical differences by enriching the light
source with a fraction of white which seemed initially
unimportant.
Acknowledgements
To Centro de Investigaci6n Cientifica y de Educaci6n
Superior de Ensenada (C.I.C.E.S.E.), and Consejo
Nacional de Ciencia y Tecnologfa (Co.Na.C. y T.) for
financial support for this investigation.
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