J Appl Phycol (2008) 20:75–82
DOI 10.1007/s10811-007-9183-6
Life-history, thallus ontogeny, and the effects of temperature,
irradiance and salinity on growth of the edible green seaweed
Gayralia spp. (Chlorophyta) from Southern Brazil
Franciane Pellizzari & Eurico C. Oliveira &
Nair S. Yokoya
Received: 20 October 2006 / Revised and accepted: 1 March 2007 / Published online: 15 June 2007
# Springer Science + Business Media B.V. 2007
Abstract Gayralia K.L. Vinogr. is a monostromatic green
alga of commercial importance in the southern Brazil, and
its cultivation is being considered. This paper reports some
basic aspects of the biology of this poorly known genus.
Two populations of Gayralia spp., from outer and inner
sectors of Paranaguá Bay, showed an asexual life history
with a distinct pattern of thallus ontogeny. In one
population (Gayralia sp. 1), zooids developed an expanded
monostromatic blade directly, while in the other (Gayralia
sp. 2) zooids produced an intermediate saccate stage, before
giving rise to a monostromatic blade. Thalli of the two
species differ in size and in cell diameter. The effects of
temperature (16–30°C), irradiance (50–100 μmol
photons m−2 s−1), and salinity (5–40 psu) on the growth
of both populations were assessed. Plantlets of Gayralia sp.
1 from in vitro cultures showed a broader tolerance to all
salinity and irradiance levels tested, with the highest growth
rate (GR; mean 17% day−1) at 21.5°C and 100 μmol photons
m−2 s−1. Plantlets of Gayralia sp. 1 collected during the
winter in the field showed higher GR, ranging from 5%
day−1 to 7.5% day−1 in salinities from 20 to 40 psu, and
2.0% day−1 and 4.3% day−1 for plantlets collected during
the summer. Gayralia sp. 2 from the field showed highest
GR at salinity of 15 psu. These results suggest distinct
physiological responses of the two species, in accordance
with their distribution: Gayralia sp. 2 is limited to the inner
F. Pellizzari (*) : E. C. Oliveira
Instituto de Biociências, Universidade de São Paulo,
Cx. Postal 11461, 05422-970 São Paulo, SP, Brazil
e-mail: francianep@yahoo.com
N. S. Yokoya
Instituto de Botânica, Seção de Ficologia,
C. Postal 4005, 01061-970 São Paulo, SP, Brazil
areas of the estuary, while Gayralia sp. 1 grows in outer
areas, where salinity values are higher than 20 psu. These
data indicate that Gayralia sp. 1 has a higher potential for
aquaculture than Gayralia sp. 2 due to its larger thalli,
higher GR, and wider tolerance to environmental variations.
Keywords Aquaculture . Brazil . Edible algae . Gayralia .
Monostromataceae
Introduction
Edible monostromatic green seaweed has been gathered on
the coast of Paraná state, southern Brazil, and sold to
restaurants offering Japanese food. This activity is sporadic
and restricted by the availability of natural stocks, whose
abundance is erratic. In view of this commercial importance,
there is local interest in cultivating this seaweed as an
economic alternative for the local population, as well as to
decrease pressure on other over-exploited resources in the
region, especially the mangroves and rain forest ecosystems.
Monostromatic green algae in Brazil have been identi-
fied as Ulvaria oxysperma (Kützing) Bliding, or Mono-
stroma oxyspermum (Kützing) Doty in previous reports
(e.g., Joly 1965; Cordeiro-Marino et al. 1993; Braga 1997;
Braga et al. 1997). However, Pellizzari (2005) studied two
populations of monostromatic green algae from mangroves
and estuaries of the Paraná coast, and concluded that both
populations should be assigned to the genus Gayralia K.L.
Vinogr. based on preliminary molecular analysis of small
subunit (SSU) rDNA.
Investigation into the responses to environmental varia-
tions of seaweeds of economic interest is a common
approach to understanding their behavior in order to
improve maricultural programs. Among the abiotic factors
76
that influence the growth, reproduction and distribution of
seaweeds, temperature and salinity are paramount (e.g.,
Lüning 1981; Lobban et al. 1985; Yokoya and Oliveira
1992a, b). Some studies have reported the influence of
environmental factors on the growth of commercial species
of green seaweeds from temperate zones (Soe-Hitun et al.
1986; Guo and Mathieson 1992; Dan et al. 2002; Kalita and
Tytlianov 2003; Cohen and Fong 2004; Leskinen et al.
2004), but little is known of species from tropical regions.
Braga et al. (1997) were the first to analyze the growth rates
of two different species of monostromatic green algae from
the Brazilian southeastern coast in different ranges of
temperature and salinity.
Pellizzari et al. (2007) reported a pilot experiment to
cultivate Gayralia sp. 1 in Paraná state, but many questions
have yet to be answered before this cultivation reaches a
commercial scale. Here, we present some basic information
aimed at establishing the biological basis for a rational
cultivation of this seaweed in the large estuarine system of
Paraná state.
Materials and methods
Life-history, thallus ontogeny and morphological analysis
Thalli of two populations of Gayralia were collected from
pneumatophores of Laguncularia racemosa C.F. Gaertn., a
common mangrove tree in the region, in Paranaguá Bay,
Paraná state, southern Brazil. Gayralia sp. 1 was collected
from the outer area (BP—25° 51′ S and 48° 34′W), and
Gayralia sp. 2 was collected from the inner area (GP—25°
32′ S and 48° 23′ W). Small pieces from the margins of the
thallus were isolated and cultured in seawater enriched with
25% Provasoli´s solution (PES/4, Oliveira et al. 1996), at 23±
1°C, 20±1 psu, 14:10 h light:dark cycle, and irradiance of 40±
10 μmol photons m−2 s−1. Morphology, liberation of zooids,
germination pattern and thallus ontogeny were observed under
a photomicroscope (Zeiss, Germany).
Temperature, irradiance and salinity experiments
Specimens of Gayralia sp.1 were collected from the outer
area of Paranaguá Bay, and transported to the laboratory in
an insulated container. Marginal pieces (±1 cm) were
excised from the thalli, washed in sterile seawater and put
on glass slides inside Petri dishes with sterile seawater.
After 24 h, the slides were transferred to 40 mL PES/4
medium, and cultured at 22±1°C, salinity 20 psu, and
14:10 h light:dark cycle. Germanium dioxide (1 mg L−1)
was used to suppress diatom growth when necessary. After
30 days, plantlets (ca. 1 mm) were transferred to 50 mL
glass flasks to start the experiments.
J Appl Phycol (2008) 20:75–82
Experiments on the interaction of temperature and
irradiance were carried out on a temperature and light
gradient plate (Oliveira et al. 1995, 1996), using plantlets
from in vitro cultures. Temperatures ranged from 16 to 30°C,
at intervals of approximately 1.5°C. Photon flux densities
were provided by cool–white fluorescent lamps (OSRAM) at
100, 75 and 50 μmol photons m−2 s−1, and a light:dark cycle
of 14:10 h. Plantlets (30 mg) were placed in vials with
50 mL PES/4 medium, salinity 20 psu. Each treatment was
tested in duplicate.
Effects of salinity variation were tested with plantlets
from laboratory cultures and from the field. Salinities
ranged from 5 to 40 psu, at intervals of 5 psu, and each
treatment was tested in triplicate. Seawater (32 psu) was
filtered, sterilized and concentrated by freezing. Gradual
melting and mixtures provided all the desired salinities. The
final salinity was checked with a refractometer (ATAGO
model S/MILL; Japan). In a second experiment, plants
of both populations (ca. 1 cm in diameter) were collected
from BP and GP areas, during the summer of 2003 and
the winter of 2004, respectively. Salinity experiments
were performed under the same conditions as already
described.
Medium renewal, weighing (fresh biomass), and mor-
phological observations were made weekly during 1 month
for each experiment. Before weighing, plantlets were
blotted with absorbent paper to remove excess water.
Growth rates (GR) were determined by variations in fresh
biomass (mg) following Bird et al. (1979), and expressed as
percent day−1.
Voucher specimens of Gayralia spp. were deposited in
the herbaria SP (Instituto de Botânica, São Paulo State,
Brazil) and SPF (Instituto de Biociências, Universidade de
São Paulo, Brazil).
Statistical analysis
Analysis of variance (ANOVA) using repeated measures
was conducted to evaluate the effects of temperature and
irradiance on growth of Gayralia sp.1. ANOVA of two
factors was also used to detect the influence of salinity on
growth of Gayralia spp., aiming to compare the develop-
ment of both species and during two seasons. Fisher's LSD
test was used as the post-hoc analysis.
Results
Life-history, thallus ontogeny and morphological analysis
Liberation of zooids from specimens of Gayralia sp. 1
started with disintegration of the zoosporangium wall
(Fig. 1a), and the release of four biflagellate zooids
J Appl Phycol (2008) 20:75–82 77

Fig. 1a–g Thallus ontogeny of

Gayralia sp.1 from the outer
sector of Paranaguá Bay (BP).
a Zooid released. b Biflagellate
zooids. c Cell body before set-
tling (5–7 μm diameter). d Cells
with prominent eyespot. e First
cell division. f Uniseriate fila-
ment formation (7 days,
30–40 μm); multiseriate fila-
ment (14 days, 45–60 μm), rhi-
zoid formation (17 days,
100–150 μm). g Development
of a plantlet with an expanded
monostromatic blade
(30–40 days; > 200 μm)

(Fig. 1b). Zooid fusions were not observed. After a short began (Fig. 1e), giving rise to a uniseriate filament that
period of swimming, one of the flagella attached to the became multiseriate after 14 days (Fig. 1f). A small leafy
substratum while the zooid spun a bit longer before settling thallus, showing around 300 μm, was observed after 30–
(Fig. 1c). After zooid attachment (Fig. 1d), cell divisions 40 days (Fig. 1g). Cells were uninucleate, with a large
78 J Appl Phycol (2008) 20:75–82

Fig. 2a–c Thallus ontogeny of

Gayralia sp. 2 from the inner
sector of BP, showing an inter-
mediate saccate stage after
20 days of culture (a, b). After
40 and 50 days the saccate frond
opened at the tip, giving rise to a
plantlet with expanded and long
monostromatic blade (c). Bars
30 μm

central vacuole, parietal chloroplast and one pyrenoid. Responses to temperature, salinity and irradiance
Plantlets attained 0.1±0.3 cm after 50–60 days.
Zooids from plants of Gayralia sp. 2 were liberated and Gayralia sp. 1 tolerated a wide range of temperature,
germinated as described for Gayralia sp. 1, but produced an irradiance and salinity, being more sensitive to higher
intermediate tube-like stage before giving rise to a blade temperatures and lower salinities. The optimum tempera-
(Fig. 2). This stage was well-developed at around 40 days ture was between 20.0°C and 21.5°C under the highest
in culture, when the plantlets were about 100 μm long irradiance tested (post hoc 0.002 and 0.0003, respectively;
(Fig. 2a,b). The saccate stage opened up after 50 days in Fig. 3). Thallus degenerated at temperatures higher than 25°C
culture, giving rise to a flat monostromatic blade of around at 100 μmol photons m−2 s−1, and died at the end of the
200 μm (Fig. 2c). No sexual reproduction was observed in experiment. Mean GR of plants cultured at 20.0 and 21.5°C
either population of Gayralia spp. and the asexual cycle were 13.0% day −1 , under irradiances of 75 μmol
repeated successively. photons m−2 s−1 and 14.0% day−1 and 17.0% day−1 at
A morphological comparison of the two species is 100 μmol photons m−2 s−1. Two way ANOVA evaluating the
presented in Table 1. The thallus of Gayralia sp. 1 is effect of temperature and irradiance on increase of biomass in
usually larger (from 4 to 9 cm), with cell size and thallus Gayralia sp. 1 showed that GR (p=0.002) was significantly
thickness of around 8±3 μm and 25±1.8 μm, respectively. affected by temperature, but not by irradiance (P=0.542,
The thallus of Gayralia sp. 2 is smaller (maximum 4 cm), Table 2).
with cell size and thickness of around 6±1.4 μm and 15± Although a peak of GR of Gayralia sp.1 cultured in vitro
2 μm, respectively. was observed at 40 psu salinity, no significant differences
in GR were observed among the salinities tested (ANOVA
P=0.22, Fig. 4). However, plants of Gayralia sp.1 colleted
in the field showed a distinct response. Fronds collected
during winter and summer showed different responses
Table 1 Morphological characteristics of two populations of mono- when submitted to a salinity gradient, (P=0.030; Fig. 4).
stromatic green seaweeds from Paranaguá Bay (BP). Gayralia sp.1 GR of the summer plants is higher at 20 psu salinity
specimens were collected from the outer sector of the estuary and (P=0.04), and Fisher´s test showed significantly higher GR
Gayralia sp.2 from the inner sector. Values represent length and width
(mean ± standard deviation; n=6) in winter plants at salinities up to 20 psu (P=0.0009,
mainly at 40 psu). When we compared cultured plantlets
Gayralia sp. 1 Gayralia sp. 2 with plants from the field, the differences in GR were
Color Green Pale green
significant (P=0.0061).
Frond sizes (cm) 7.2×6.8±2.7×1.5 4.2×3.0±0.7×0.9 Gayralia sp. 2 had, in general, a lower GR when
Thickness (μm) 25±1.8 15±0.7 compared to Gayralia sp. 1 under salinity gradient (Fig. 5).
Cell size 7×11±1.3×1.5 5×7±0.4×0.5 Higher GR were observed at 10, 15 and 20 psu salinities,
(transverse section; μm) being 1.5, 2.5 and 1.4% day−1, respectively.
Marginal cells (μm) 8×13±1.6×1.9 6×7±0.4×0.5 The ANOVA comparing differences in GR of Gayralia
Medium cells (μm) 9×13±2×1,8 5.5×8±0.8×0.9
sp.1 and G. sp.2 collected during the summer and submitted
Basal cells (μm) 10×14±1.4×1.8 9×11±1.0×1.1
to the salinity gradient was significant (P=0.018). Fisher´s
Rhizoidal cells (μm) 35×11±1.3×0.7 25×7±2.5×0.6
test showed that GR were significantly higher for Gayralia
J Appl Phycol (2008) 20:75–82 79

Fig. 3 Growth rates (GR) of 25

Gayralia sp. 1 cultured at sev-
eral temperature and irradiances
(μmol photons m−2 s−1) after
35 days at salinity 20 psu. Each 50
20
data point is the mean of two 75
replicates. Bars Standard 100
deviation
15

GR (% day -1)
10

-5
16 18 20 21 23 24 25 26 27 30
Temperature (0C)

sp.2 at 15 psu (P=0.002) compared to the other salinities ,
and for Gayralia sp.1 at 20 psu (P=0.005).
Discussion
The occurrence of an asexual life history in both popula-
tions of monostromatic green algae from mangroves and
estuaries of the Paraná coast corroborates the conclusion
that both populations should be assigned to the genus
Gayralia based on preliminary molecular analysis of SSU
rDNA (Pellizzari 2005). However, differences in thallus
and cell sizes, and thallus ontogeny, as well as distinct
responses to environmental parameters, indicate that we are
dealing with two different species of Gayralia. Leskinen et
al. (2004), working with Ulva intestinalis and Ulva
compressa in the Baltic Sea, used halotolerance as an
additional character, besides genetic variation using internal
transcribed spacer (ITS) markers, to distinguish morpho-
logical plastic species with overlapping characters.
Based on the systematics of the genus Monostroma
described by Tatewaki (1972), Gayralia sp. 1 showed a
thallus ontogeny similar to M. latissimum and M. undu-
latum [=Protomonostroma undulatum (Wittrock) K.L.
Vinogr.], while thallus ontogeny of Gayralia sp. 2 was
similar to M. oxyspermum [=Gayralia oxysperma (Kutz.)
K.L. Vinogr. ex Scagel et al.]. However, further studies on
molecular systematics of Brazilian monostromatic green

Table 2 Results of analysis of variance (ANOVA) used to test the influence of the independent variables (physical parameters) on the growth rate
(GR) of Gayralia spp.

Variables Degrees of freedom F P

Irradiance (I) 2 0.624 0.542

Temperature (T) 9 3.885 0.002
Interaction I vs T Gayralia sp. 1 18 1.596 0.125
ANOVA (P value) Post hoc (Fisher LSD) Post hoc (Fisher LSD)
Salinity vs GR Gayralia sp. 1 (culture) 0.223
Salinity vs GR Gayralia sp. 1 0.030 P=0.04 P=0.0009
(field plants);
summer × winter
Summer; salinity 20 psu Winter; salinity 40 psu
Salinity vs GR of Gayralia sp. 1 0.018 P=0.005 Gayralia sp.1; salinity 20 psu P=0.002 Gayralia sp.2;
and sp. 2 (field plants) salinity 15 psu
80 J Appl Phycol (2008) 20:75–82

Fig. 4 Comparison of GR of 30
Gayralia sp.1 at several salin-
ities (psu) in young plants from
laboratory culture and from BP, field (summer)
collected during the winter and 25
field (winter)
summer of 2003. Each data
in vitro
point is the mean of three
replicates. Bars Standard 20
deviation

GR (% day -1)
15

10

-5
5 10 15 20 25 30 35 40
Salinity (psu)

algae are necessary to clarify the species identities. The
physiological responses and thallus ontogeny of Gayralia
spp. are partially in accordance with observations reported
for monostromatic green species identified as Monostroma
sp. and Ulvaria oxysperma in Brazil (Cordeiro-Marino et
al. 1993; Braga et al. 1997), which were considered as
eurythermic and euryhaline species. These physiological
responses are typical of monostromatic green seaweeds that
are distributed mainly in estuaries of subtropical and
temperate zones and are subjected to a broad fluctuation
of salinity (Biebl 1937).
Exponential growth occurred from 16 to 24°C in
Gayralia sp. 1 (data not shown) and the upper limit of
survival was around 26°C, which was lower than observed

Fig. 5 Comparison of GR of 6
Gayralia sp.1 and Gayralia sp.2
collected from the outer and
inner sectors of BP, respectively, 5
during the summer of 2004. Gayralia sp.2
Each data point is the mean of Gayralia sp.1
three replicates. Bars Standard 4
deviation

3
GR (% day-1)

-1

-2
5 10 15 20 25 30 35 40
Salinity (psu)
J Appl Phycol (2008) 20:75–82
by Braga et al. (1997) for a monostromatic green species
collected from the southern coast of São Paulo State. These
responses suggest the existence of ecotypic differentiation
among populations from São Paulo and Paraná states.
Although statistical analysis indicates that GR are
affected predominantly by temperature, Gayralia sp. 1
showed the highest growth rates from 20°C to 22°C, at
100 μmol photons m−2 s−1, and from 20 to 40 psu salinity,
ranges that should be utilized to select seasons and areas for
cultivation. These values are in accordance with observa-
tions on the distribution of the species in the area made by
Pellizzari (2005), who found larger fronds during the winter
with temperature ranging from 19 to 21°C. Specimens kept
at higher temperatures (ca. 25°C) showed low tolerance to
higher irradiances, a fact already observed by Cordeiro-
Marino et al. (1993) for Monostroma sp. collected at
Espírito Santo state in southeastern Brazil. Although we
utilized a small range of irradiances, it was quite clear that
this could be an important factor interacting with temper-
ature in controlling the GR and distribution of Gayralia
spp. in the estuary, since the species did not grow at low
irradiance (50 μmol photons m−2 s−1).
According to Druehl (1981), the adverse effects of high
temperatures could be minimized at higher salinities or
lower irradiances. However, in general, intertidal mono-
stromatic chlorophytes have higher GR at higher irradi-
ances. Guo and Mathieson (1992) did not observe
photoinhibition until 570 μmol photons m−2 s−1 in four
species of monostromatic chlorophytes [Gayralia oxy-
sperma, Ulvaria obscura (Kützing) P. Gayral ex C. Bliding,
Monostroma grevillei (Thuret) Wittrock and Protomonos-
troma undulatum f. pulchrum (Farlow) M.J. Wynne].
Growth rates of Gayralia sp.1 do not differ significantly
over the salinity range tested. However, they differ when
comparing GR of G. sp.1 between summer and winter, and
also comparing GR of G. sp. 1 and G. sp.2.
Gayralia sp.1 grew in all salinities tested, but showed
higher GR at salinities of 20 to 40 psu. Monostroma
grevillei and M. pulchrum, from New England (USA),
showed salinity tolerance from 0 to100 psu, although
optimum growth occurred from 10 to 40 psu (Guo and
Mathieson 1992), which is similar to our results. The
differences between GR of Gayralia sp.1 collected during
the winter (ca. 6% day−1) and plants cultured in vitro (ca.
10% day−1) could be attributed to the optimum conditions
established in culture and to acclimation.
Gayralia sp.2 had a lower GR and tolerance to a
narrower salinity variation than Gayralia sp.1. The growth
of the former diminished abruptly in salinities higher than
30 psu, suggesting a distinct physiological response. This
behavior could explain the restricted distribution of Gayralia
sp. 2 in the inner sectors of the estuary and tidal channels of
Paranaguá Bay, while Gayralia sp.1 occurs preferentially in
81
the outer sectors of the estuary, where salinity is higher than
20 psu. Based on these results, Gayralia sp. 1 was selected
for a mariculture program in Paraná state (Pellizzari et al.
2007). Another advantage of this species is that cultivation in
the outer sector of the estuary could avoid conflict with the
utilization of the inner areas for oyster and shrimp
cultivation.
Acknowledgments The authors are grateful to the Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES, Brazil) for
providing a scholarship to F.P., and to the Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq, Brazil) for
supporting E.C.O. and N.S.Y. We thank Mauricio Camargo for
comments on statistics.
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