Cultivation of Chondrus canaliculatus (C. Agardh)Greville (Gigartinales, Rhodophyta) in controlledenvironments

Mario Edding Æ Erika Fonck Æ Roberto Acuna Æ Fadia Tala

Received: 31 October 2006 / Accepted: 5 October 2007 / Published online: 27 October 2007� Springer Science+Business Media B.V. 2007

Abstract Increased market demand for algal raw materials has stimulated research and

development into new cultivation technologies, particularly in countries with economically

important seaweed industries. Chondrus canaliculatus is a red alga endemic to the tem-

perate Pacific coast of South America and produces a complex carrageenan. In Chile, this

potential marine resource has been underutilized since the early 1990s, after it was

replaced mainly by imports of Kappaphycus and Eucheum from Asia. The study of the

cultivation of C. canaliculatus in outdoor tanks demonstrates the potential to produce large

quantities of algal material due to the high productivity observed. The material produced

may be important in Chile as feed for cultured algal herbivores such as Haliotis spp. Data

were obtained on the key factors affecting the growth of this alga, including light, tem-

perature, and nutrient concentrations. The results showed that, on a dry weight basis,

productivity varied seasonally between 40 g m–3 wk–1 in autumn and 200 g m–3 wk–1 in

spring, and the optimal stocking density in our experiments was 8 kg m–3. Effects of self-

shading were among the most important limiting factors in the culture. Discussion is

presented on the interactions between temperature, pH, salinity, enrichment with N and P,

and water exchange on the maximal growth of C. canaliculatus in this culture.

Keywords Algae � Chile � Chondrus � Cultivation � Productivity � Seaweed �South East Pacific

Introduction

In Chile, various species of Phaeophyceae and Rhodophyta (Norambuena 1996; Buschmann

et al. 2001; Anuario Estadıstico de Pesca 2005) have been exploited since the 1950s with

production fluctuating between 74,000 and 410,850 wet metric tonnes over the last

15 years. Most of the production has been based on the exploitation of wild stocks used as

M. Edding (&) � E. Fonck � R. Acuna � F. TalaLaboratorio de Botanica Marina, Departamento de Biologıa Marina, Facultad de Ciencias del Mar,Universidad Catolica del Norte, Casilla 117, Coquimbo, Chilee-mail: medding@ucn.cl

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raw material for the production of phycocolloids, both in Chile and abroad. In recent years,

an indeterminate quantity of seaweeds has been continuously harvested from wild stocks as

fresh food for abalone (Haliotis spp.) in mass culture. In Chile, commercial-scale seaweed

cultivation is limited to Gracilaria chilensis. However, research into potential cultivation of

other species such as the brown macroalgae Lessonia trabeculata Villouta & Santelices and

Macrocystis pyrifera (Linnaeus) Agardh and the red macroalgae Gigartina skottsbergiiSetchell & Gardner, Callophyllis variegata (Bory de Saint Vincent) Kutzing and Chondr-acanthus chamissoi (C. Agardh) Kutzing is ongoing (Bulboa et al. 2005; Buschmann et al.

2001, 2004; Edding et al. 1990; Edding and Tala 2003; Westermeier et al. 2006).

Most carrageenan is extracted from Kappaphycus alvarezii (Doty) ex P. C. Silva to

produce j carrageenan, and Eucheuma denticulatum (Burman) Collins and Hervey, to

produce i carrageenan. Both these species are cultured predominantly along the coasts of

the Philippines and Indonesia (Trono 1993; McHugh 2003). Chondrus crispus Stackhouse,

the original source of carrageenans (j and k forms), is still used to a limited extent

(McHugh 2003). Chile has imported raw Eucheuma seaweed for carrageenan processing,

80% of which is used by the food industry. This genus appears to be replacing the local

production of ‘‘lichen gomosus’’, composed of Ahnfeltiopsis furcellata (C. Agardh) P. C.

Silva and DeCew, and Chondrus canaliculatus (C. Agardh) Greville. The national demand

for carrageenophytic algae has undergone an important increase since 1995, mainly due to

the establishment of processing plants to extract the colloids, producing a total of 1586 tons

in 2004 (Anuario Estadıstico de Pesca 2005).

C. canaliculatus is a well-known marine alga endemic to the Pacific temperate coast of

South America; it is a potentially important carrageenan-producing species occurring from

the Chinchas Islands, Peru (14�S) (Dawson et al. 1964; Acleto and Zuniga 1998) to Chiloe-

Chile (41�S) (Ramirez and Santelices 1991). The binomial taxonomy of this species is

under revision, however, since the cystocarpic morphology (Arakaki et al. 1997) and the

phylogenetic analysis based on rbcL sequencing (Hommersand et al. 1999) demonstrated

that the species does not conform to Chondrus. Until a new designation of the binomial is

agreed upon, the commonly used name, C. canaliculatus, will be used throughout this

paper. This alga grows on protected shores, forming perennial holdfasts in inter- and

subtidal areas in at least 17 known locations throughout Chile (Ramirez and Santelices

1991; Acuna 2000). Some C. canaliculatus populations diminished due to uncontrolled

human exploitation during the 1980s (Vasquez and Westermeier 1993). Carrageenan yields

from these plants are 50.5% dry wt for cystocarpic plants, and 54.4% for tetrasporic plants,

with contents of 16% and 13% of 3- and 6-anhydrogalactoses, respectively (Ayala and

Matsuhiro 1986). Alkali-modified carrageenans improves the yield of 3,6-anhydrogalac-

tose by 25% (Matsuhiro and Zalungo 1983). The presence of k, i, and j carrageenan in all

phases of its life cycle were described by Matsuhiro and Rivas (1993), while Martinez

(1998) found that the proportion of j and i carrageenan could be modified by changes in

salinity and temperature. Tapia et al. (1987) provided data on its growth in underwater

containers, with a maximum daily growth rate of 3.48%. Vega and Meneses (1999)

described the seasonal and spatial (i.e., intertidal versus subtidal stands) variations in the

productivity and reproduction of C. canaliculatus under field conditions in a population at

Puerto Aldea, in north-central Chile. Our previous experience with mass cultures of marine

algae in outdoor tanks (Edding et al. 1987; Edding 1995), as well as that of other authors

(e.g., Bidwell et al. 1985; Braud and Armat 1996) led us to examine the feasibility of

cultivating C. canaliculatus in the laboratory and in outdoor tanks as a way of insuring the

long-term stability of supply and price of carrageenan-containing raw material. The

advantages of seaweed farming would be: (a) independence from fluctuating climatic

284 Aquacult Int (2008) 16:283–295

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conditions, (b) independence from the labour-intensive seaweed collecting, (c) increased

purity of raw material, and (d) the possibility of choosing seaweed strains with high

carrageenan yields and carrageenan having desired compositions and properties. How the

environmental factors may effect the growth and productivity of C. canaliculatus is critical

for the determination of the parameters affecting its culture.

Materials and methods

Algal material

An initial stock biomass of C. canaliculatus, mainly vegetative plants, was collected from

a natural population located at Puerto Aldea (30�160S; 71�300W) at a depth of 5 m. This

location is the site of a fishing village at the southern end of Tongoy Bay, 62 km south of

Coquimbo, Chile. Here, the mountains protect the study site from the prevailing S-SW

winds except during winter when the winds blow from the north, generating heavy surf.

Algal biomass was transported to the laboratory in thermal coolers, and cleaned of her-

bivores and epiphytes. This material was maintained in a greenhouse holding pond, which

received a continuous flow of fresh seawater, and was used at our laboratory to supply algal

biomass for different experiments.

Laboratory experiments

Several experiments were designed to observe the response of Chondrus canaliculatus in a

controlled environment to environmental changes to which the algae might be exposed

during outdoor tank cultivation. The relative growth rate of C. canaliculatus was evaluated

under different light and temperature regimes, using 250 ml Erlenmeyer flasks containing

about 4 g wet weight in 50 ml of filtered (0.45 lm) seawater enriched with Provasoli

medium (Starr and Zeikus 1987). The pH was adjusted to 8.08 using TRIS buffer, and the

culture medium was replaced daily. The cultures were arranged on a light-temperature

gradient table to obtain combinations among temperatures of 8.6, 14.3, 20.7, 26.7�C and

photon flux densities of 54, 127, and 216 lmol m–2 s–1. All tests were done using three

replicates. Neutral light filters were used to provide a gradient of light from 40 W cool

white fluorescent lamps. The intensity of photosynthetic active radiation (PAR) was

determined with a Li-250 radiometer (LI-COR Inst., USA). All systems were allowed to

acclimate for a week to the test conditions, and then daily growth was estimated by

measuring biomass increments following the methods of Edding (1995).

The effect of salinity on photosynthesis was evaluated in algae incubated in a walk-in

chamber, with a controlled temperature (18�C), PAR of 100 lmol m–2 s–1 and a photo-

period of 12:12 h (light:dark). Seawater salinities of 20, 25, 30, 32.7, and 42.5 psu were

established by dilution with distilled water for salinities lower than 34 psu and removing

H2O by freezing to obtain greater salinity. Plants (4 g wet weight per replicate; n = 3) were

maintained in culture for 96 h at the different salinities, and then transferred to an ambient

salinity of 34 psu for 48 h, after which productivity was measured as changes in oxygen by

using the Winkler method with light/dark bottles following the methods of Strickland and

Parsons (1972).

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Outdoor tank cultures

Culture experiments were done from austral spring 1994 to spring 1995 using 1000 l

rectangular fiberglass tanks. The tank design was the same as that which we previously

used for Gracilaria culture (Ambler et al. 1988; Edding 1995). Tanks received a contin-

uous flow of 50 lm sand-filtered seawater from La Herradura Bay (30�S), and continuous

aeration (Edding 1995).

The rates of productivity at different algal densities were observed over annual cycles.

The densities used for the algal culture were 2.1, 4.2, 8.4, and 15.5 kg m–3 (n = 2), and

the production was measured each 15 days. The increase in biomass was removed after

each measurement, determined by weighing, thus leaving the same initial amount of algae

in the tank at the beginning of each measurement period. The biomass produced was dried

in an oven at 65�C to obtain dry weights (dw), and the productivity was expressed as

g dw m–3 wk–1.

Productivity was analyzed in relation to seasonal variations in light, temperature, pH,

salinity, and nutrient concentrations in the incoming water. Water temperatures were

measured using a handheld thermometer both in the tanks and in incoming water three

times per day (09:00, 12:00, and 18:00 h). At the same time, the surfaces and underwater

PAR radiation were measured in the tanks using a Li-Cor quantum meter as cited above.

The pH was measured with a Digi-Sense meter (Cole Palmer Co., USA), and the salinity

was measured by conductivity (ECT-5 meter, Toho Dentan Co.). Seasonal variations

(n = 3) of nitrogen and phosphorus (Kjeldahl method; AOAC 2000) in the tissues of

C. canaliculatus, and nitrate, nitrite, ammonium, and phosphate were measured in triplicate

in the water tanks (Strickland and Parsons 1972).

In addition, analyses of carrageenan for each treatment (n = 3) were made by aqueous

extraction (Ayala and Matsuhiro 1986) at the end of the experiment.

The effect of N and P concentrations (n = 5, 25, 125, 625, and 3125 lM) at a ratio of

about 15:1 on productivity were recorded during three summer weeks (December 94 to

January 95), together with tissue content of these nutrients as described above. The

experiment was carried out in 1000 l tanks with an algal biomass density of 15.5 kg m–3.

The nutrient sources used were reagent-grade NaNO3 and KH2PO4. In addition, analyses of

carrageenans were made for each treatment (n = 3) at the end of each experiment. The

occurrence of epiphytism was recorded by harvesting and identifying epiphytes from the

cultured plants over a period of 30 days in each tank treatment.

The effect on productivity of changing rates of water flow through tanks was measured

in two control tanks with a flow rate of 425 l h–1 (10.2 vol d–1), two tanks with 208 l h–1

(5 vol d–l), and two tanks with 42 l h–l (1 vol d–l). Variations in temperature, pH, salinity,

and tissue N and P was measured in these tanks as described above.

Data analyses

One-way analysis of variance (ANOVA) or Kruskal–Wallis tests were performed,

depending on the normality and heterocedasticity of variances. Percentage values were

angularity transformed. The null hypothesis was rejected at a values = 0.05. A Tukey

multiple range test was used to detect significant differences within treatments. A two-way

ANOVA test was used to evaluate differences measured in the cross-gradient tests with

light and temperature in the laboratory. When the number of data points differed between

experiments, a multiple comparison procedure (Dunn’s method) was applied.

286 Aquacult Int (2008) 16:283–295

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Results

Laboratory experiments

Algae in the test cultures always presented a healthy appearance, and those in optimal

conditions grew well throughout the test period. The growth rate increased with increasing

temperature (at day 10) of culture under some of the test conditions (Table 1). No growth

was detected with the combination of lower temperature and high range of irradiance. At

the higher temperature, there was no statistically significant difference in growth attrib-

utable to an increase in light level (Table 1). In the second phase of the experiment (at day

14), there was no growth at the lowest (8.6�C) and highest (26.7�C) temperatures

(Table 1). No effect of light on growth was detected at 14.3�C and no growth was observed

at the highest light intensity. At 20.7�C, growth was observed at all light intensities with no

significant differences attributable to the different light intensities (Table 1).

For both test periods, statistical analysis (two-way ANOVA) showed significant dif-

ferences (P \ 0.05) only due to the temperature factor; there was no significant difference

(P [ 0.05) due to the light factor nor to the light/temperature interaction.

Figure 1 shows the effects of salinity on net photosynthesis by C. canaliculatus plants.

Photosynthesis differed significantly (one-way ANOVA, P \ 0.05) among the salinities.

Intermediate salinities (30 and 32.7 psu) showed higher photosynthesis, and both lower

and higher salinities depressed photosynthetic activity.

Outdoor tank cultures

Figure 2a shows that water temperatures over the culture period varied from a winter low

of 12�C to a summer high of 23�C, with only slight differences between incoming water

and tank water. In both cases, an increase in temperature was observed during the daily

measurements in each season (Fig. 2a). Irradiance values were highly variable during the

day and within each season (Kruskal–Wallis, P \ 0.05). The average irradiance reached at

noon was similar in both spring and summer (Fig. 2a). The pH in the tanks remained fairly

constant over time, with increases registered towards the end of day; in both inflow and

tank water (Fig. 2b). A significant (Kruskal–Wallis, P \ 0.05) rise in pH of the water was

detected in the water in the tanks when compared with the incoming water. There were no

significant differences (Kruskal–Wallis, P [ O.O5) in salinity between the incoming and

Table 1 C. canaliculatus growth rate at different combinations of light and temperature in laboratoryexperiments. The algae were acclimated for 1 week before the measurements began

8.6�C 14.3�C 20.7�C 26.7�C

Growth rate at day 10 (% day–1)

54 lmol m–2s–1 1.23 ± 1.23 2.07 ± 0.04 3.69 ± 0.29 3.10 ± 2.98

127 lmol m–2s–1 0.0 1.18 ± 0.48 3.59 ± 1.38 3.17 ± 1.66

216 lmol m–2s–1 0.0 0.71 ± 0.47 3.69 ± 0.25 4.16 ± 1.20

Growth rate at day 14 (% day–1)

54 lmol m–2s–1 0.0 0.14 ± 0.10 0.71 ± 0.04 0.0

127 lmol m–2s–1 0.0 0.12 ± 0.12 0.93 ± 0.65 0.0

216 lmol m–2s–1 0.0 0.0 0.72 ± 0.01 0.0

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tank water (Fig. 2b). However, differences in salinity were observed (Kruskal–Wallis,

P \ 0.05) during spring 1995, when the salinity was higher than in spring 1994 and

summer 1995 (Fig. 2b).

The PO4 level in water was significantly different (one-way ANOVA, P \ 0.05) among

seasons, with higher values in autumn and in the spring of 1995 (Fig. 2c), whereas the

tissue phosphorus levels were significantly higher (one-way ANOVA, P \ 0.05) only in

the spring of 1995 (Fig. 2c). The main N source in the tank water was nitrate, followed by

lower values for ammonium (2 lM) and nitrite (0.5 lM). Seawater nitrates remained fairly

constant over the entire study period (one-way ANOVA, P [ 0.05), with only slight

decreases in autumn (Fig. 2c). Ammonium differed among seasons (Kruskal–Wallis,

P \ 0.05), with higher values during spring 1994 and decreasing markedly during the other

seasons (Fig. 2c). The tissue N levels showed a significant (one-way ANOVA, P \ 0.05)

increase during the year of culture, from spring 1994 to spring 1995 (Fig. 2c). Ratios for

N:P were similar when comparing the water with the plant tissues.

Productivity of C. canaliculatus varied significantly (two-way ANOVA, P \ 0.001)

with season and culture density (Fig. 3). In overall terms, the productivity showed a

seasonal trend, with maximum values in spring 1994 and 1995, decreasing during summer

and autumn 1995, with an increase in biomass beginning again toward winter. During the

summer, it was impossible to quantify the productivity of Chondrus in the smaller density

tanks due to the high degree of epiphytism occurring on the plants. During spring 1994, a

significant (one-way ANOVA, P \ 0.05) depression in productivity was found when

compared with the other densities (Fig. 3). Almost equal productivities (one-way ANOVA,

P \ 0.05) were obtained in autumn with tank loadings of 4.2 and 8.4 kg m–3, which were

both higher than when the initial loading was 15.5 kg m–3 (Fig. 3). Productivity was not

significantly different among the four tank loading levels during the winter (one-way

ANOVA, P = 0.059) (Fig. 3). During spring 1995, high productivity was apparent at a

density of 8.4 kg m–3, although a statistically significant difference could not be detected

among the different algal densities (one-way ANOVA, P = 0.166). Over a 112-day period

during winter–spring 1995, the biomass of C. canaliculatus doubled after the first 12 weeks

(tank loading 8.4 kg m–3). The culture productivity fluctuated between a maximum dry

weight of 195 g m–3 wk–1 reached in the fifth week, and a minimum of 54 g m–3 wk–1 in the

14th week, with an average dry weight value of 128 g m–3 wk–1.

300

400

500

600

20 25 30 32.7 45

Salinity (psu)

Pho

tosy

nthe

sis

(um

ol O

2 g-

1 h-

1 )

Fig. 1 Net photosynthesis of C. canaliculatus at different salinities. Values represent means ± 1 standarderror

288 Aquacult Int (2008) 16:283–295

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Over the test period, the carrageenan yield was greater in the culture system containing

8.4 kg m–3, with an average carrageenan content of 26%, followed by the culture at

15.5 kg m–3 with a carrageenan yield of 21%, and at 4.2 kg m–3 with an average of 15%

carrageenan. In the system containing 2.1 kg m–3, the yield of carrageenan could not be

determined due to the high degree of contamination of the culture by epiphytes. The

variability in carrageenan yield can be explained by the quality of the plants, since

0

5

10

15

20

25

Wat

er t

empe

ratu

re (

°C)

0

1000

2000

3000 PF

D (µm

ol m-2 s -1)

a

6.5

7.5

8.5

9.5

pH

34

35

36

37

Salinity (psu)

Inflow water pH Tank water pH

Inflow water salinity Tank water salinityb o o

0

2

4

6

8

10

12

Spring1994

Summer Fall Winter Spring1995

Wat

er n

utri

ents

(µM

)

0

10

20

30

40

50

Tissue N

and P (m

g g-1)

NO3 NO2 NH4PO4 Tissue N Tissue P

c

T° inflow water T° tank water PFD

Fig. 2 Seasonal variations in (a) photon flux density and water temperature in the tanks, (b) salinity and pHof inflow and tank water, (c) nutrient in the incoming water and in C. canaliculatus tissues. For photon flux,water temperature, pH, and salinity, the points within each season represent the averages of the dailymeasurements taken at 9:00, 12:00 and 18:00 h. Values represent means ± 1 standard error

Aquacult Int (2008) 16:283–295 289

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populations are exposed to different environmental conditions and seasonality also affect

carrageenan production.

The total productivity was not statistically significant (one-way ANOVA, P = 0.167) at

different nutrient concentrations. Nevertheless, a marked increase in the productivity was

observed during the 3rd week of culture, both at N/P ratios of 125:8.3 and 625:41.7

(Fig. 4a), while for 625:41.7 treatment the tissue N increased (Fig. 4b). In the other

treatments, a final decrease in the levels of tissue N was recorded (Fig. 4b). In all nutrient

treatments, the P concentration in the tissue increased slightly from initiation of the cul-

tures to their termination (Fig. 4b). The initial yield of carrageenan was 20%, and a final

average yield of 31% was recorded in all the cultures. Epiphytes growing on C. canali-culatus during the study period included benthic diatoms, Ectocarpus acutus Setchell and

N.L. Gardner, E. confervoides Le Jolis, Hincksia granulosa (J.E. Smith) P.C. Silva, En-teromorpha intestinalis (Linnaeus) Link, and Ulva costata (M.A. Howe) Hollenberg. The

last named species was persistent and abundant (Fig. 4c). Epiphytes were more frequent in

the fertilized tanks than in controls, although it was observed that the higher levels of

nutrients negatively affected their growth (Fig. 4c), and decreased their biomass and cover.

The epiphytes were abundant in spring and summer, but were absent from the cultures

during fall and winter.

The water temperature (Fig. 5a), pH, and salinity (Fig. 5b) increased in tanks with the

lower seawater flow rates (one-way ANOVA, P \ 0.05). The productivity among tanks

having different flow rates showed no significant differences (one-way ANOVA,

P = 0.773) (Fig. 5a). A reduction in the tissue levels of nitrogen and phosphorus was

observed with decreasing water flow rate (Fig. 5c).

Discussion

The present study is part of pioneering efforts in Chile to culture economically valuable

marine algae in land-based seawater tanks; C. canaliculatus was of interest based on its

high content of carrageenan and other food additives needed by the food industry in the

temperate SE Pacific region (McHugh 2003). The culture of algae in land-based tanks

(Braud and Amat 1996) rather than at sea (Norambuena 1996) is feasible on the Chilean

0

50

100

150

200

250

Spring1994

Summer Winter Spring1995

Pro

duct

ivit

y (g

dw

m-3

wk-

1 )

2.1 km-3 4.2 km-3 8.4 km-3 15.5 km-3

Fall

Fig. 3 Productivity of C. canaliculatus cultured in outdoors 1000 l tanks at different culture densities inCoquimbo, Chile. Values represent means ± 1 standard error

290 Aquacult Int (2008) 16:283–295

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coast since there are no major climatic variations between the seasons that would interrupt

the culture process. Also, tank cultures lend themselves to controlled regimes of light and

nutrients, and proper management of the water quality control epiphytes (Bouarab et al.

2001). These considerations can collectively lead to higher productivity rates in tank

cultures in different seasons.

Our study clearly demonstrated the dependence of C. canaliculatus on environmental

factors. Growth of the vegetative plants in response to light and temperature variation was

0:05:0

.3

25:1.

7

125:8

.3

625:4

1.7

3125

:208.3

1º week

3º week0

50

100

150

200

250

Pro

duct

ivit

y(g

dw

m-2

wk-

1 )

0

4

8

12

16

20

0:0 5:0.3

25:1.

7

125:8

.3

625:4

1.7

3125

:208.3

Tis

sue

leve

l (m

g g

dw-1

)

Ni Nf Pi Pf

0

20

40

60

80

0:0 5:0.3

25:1.

7

125:8

.3

625:4

1.7

3125

:208.3

Nutrients (N:P µM)

Dis

lodg

ed e

piph

ytes

(%

)

0

1

2

3

4

5

Biom

ass (%dw

)

Ectocarpales Ulvales Epiphyte biomass

a

b

c

Fig. 4 C. canaliculatus (a)productivity, (b) tissue nitrogen(Ni = initial N; Nf = final N) andphosphorous (Pi = initial P;Pf = final P), and (c) epiphytechanges during cultivation inoutdoor tanks with differentnutrient concentrations. Theexcess production of C.canaliculatus was removedweekly. Values representmeans ± 1 standard error

Aquacult Int (2008) 16:283–295 291

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similar to that of carposporic plants of this species cultured under similar conditions

(Acuna 2000). The results have shown good growth of the plants at 14�C and 20�C, at

lower and middle irradiance levels. Generally, the higher (environmental) temperature

ranges proved lethal for some seaweed (Breeman 1988; Martinez 1999), and the lowest

ranges produced decreases in growth and productivity (Martinez 1998; Pang et al. 2005).

Changes in pH in the culture water have been related to the inorganic carbon availability

in the seawater, and are active in controlling the balance among inorganic ionic forms of

carbon in the water (Zou et al. 2004). Thus, productivity should be limited both by the

ionic species of carbon available and the amounts of inorganic carbon in the water. Craigie

100

200

300

400

Pro

duct

ivit

y(g

dw m

-3w

k-1 )

16

17

18

19

20

Tem

perature (°C)

Productivity Temperaturea

8.0

8.5

9.0

9.5

10.0

pH

34.2

34.4

34.6

34.8

35.0

Salinity (psu)

pH Salinityb

0

5

10

15

20

25

Inflow 10.2 1

Water exchange (volumes per day)

Tis

sue

N (

mg

g dw

-1)

3

4

5

Tissue P

(mg g dw

-1)

N Pc

5

Fig. 5 C. canaliculatus (a)weekly productivity and tankwater temperature in cultivationwith different rates of waterexchange, (b) tank water salinity,and pH variations, and (c) tissuenitrogen and phosphorus. Valuesrepresent means ± 1 standarderror

292 Aquacult Int (2008) 16:283–295

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and Shacklock (1995) reported that adding CO2 or H2CO3 to cultures attenuated the pH

change and maintained the inorganic C availability in C. crispus culture. Salinity is also a

factor that must to be taken into account in open-air tank cultures of C. canaliculatus, as

the present results showed that both low and high salinity values depressed its photo-

synthesis. Continuous replacement of water in the tanks helps prevents changes in salinity,

where there may be dilution of the water by rain, or increases in salinity due to

evaporation.

Nutrient dynamics play a key role in obtaining a high-quality product, both in its

individual appearance, and in the management of epiphytes. Addition of high concentra-

tions of nutrients produced a decline in productivity in the first weeks, but also reduced

contamination by epiphytes, which appeared to be more affected than C. canaliculatus.

Manipulation of nutrient concentrations may be a useful method for controlling epiphytism

in these tank cultures in the future, although both photosynthesis and carrageenan content

could decline under these conditions (Chopin and Floc’h 1992). This supposition was not

apparent in the present study, however, as the carrageenan yield from C. canaliculatusremained unchanged in spite of the different nutrient treatments. Also, it is important to

mention that the carrageenan yield obtained by Ayala and Matsuhiro (1986) was based on

plants collected in April and December in the same locality as this work. The carrageenan

yield obtained in this work can be explained by the life history of the algae and the

environmental conditions plants have in the cultivation tanks.

The seasonal pattern of productivity of C. canaliculatus followed that reported for other

algae from temperate coasts, with maxima in spring and minima toward winter (Edding

et al. 1987). The continuous production observed for C. canaliculatus cultivated in tanks

could reach a mean of 6.5 wet Kg year–1 m–3 (128 g dw m–3 wk–1) at the most favorable

density. Similar productivity values were found for Gracilaria chilensis (125 g dw m–3 wk–1,

Edding 1995), Chondracanthus chamissoi (160 g dw m–3 wk–1, M. Edding, unpublished)

and Ulva sp. (92 g dw m–3 wk–1) (J. Macchiavello, personal communication) cultivated in

the same tank system. This observation was carried out at the experimental level, the next

step being to consider economic evaluation of the model at the pilot scale. The productivity

of C. canaliculatus was in the same range as the values observed in tank culture of

C. crispus in Canada (Craigie and Shacklock 1995). The climatic conditions in Canada,

however, included low water temperatures between December and April, resulting in very

low rates of production, during which time the culture had to be suspended (Craigie and

Shacklock 1995). In Chile, this type of culture could be carried out all year, since the

normal summer–winter temperature variation in the seawater seldom exceeds 10�C, not

going below a minimum of 11�C. Algal production in tanks may be severely altered,

however, by climatic and atmospheric changes that accompany the El Nino southern

oscillation (ENSO) phenomenon, which occurs periodically on the Chilean coast, and often

produces mass mortalities of naturally occurring algal populations (Glantz 1998; Edding 1999).

Different culture objectives can be obtained by varying the nutrient inputs to the cul-

tures. Concentrations of 125 N:8.3 P stimulate productivity, whereas higher concentrations

could be used in spring and summer to control epiphytism. Future research should include

scaling up of the culture and verification of the optimal conditions, as well as the selection

of strains and clones providing optimum production of biomass and carrageenans. A good

measure of success was achieved in this area by Craigie and Schaklock (1995) in the

culture of C. crispus in Canada; these authors selected clones having optimal growth,

desirable types and yields of carrageenans, and favorable colour mutants. Factors such as

carbon supply, pest control, and disease prevention also need to be considered in futures

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studies on C. canaliculatus in tank culture. Chondrus canaliculatus biomass needs to be

evaluated in practical terms for its potential use in the carrageenan industry, for the culture

of marine herbivores such as Haliotis spp., and even for human consumption.

Acknowledgements This study was made possible by support from Multiexport Algas SA (Chile). Wethank Mr. Danilo Peralta for expert technical help and acknowledge technical assistance from the studentsRoberto Silva, Carmen Rivera, Susana Espana, and Fungyi Chow. Finally, we would like to thank JulioNazar, Lautaro Moreno, and Dr. Louis DiSalvo for helpful reviews of this manuscript, as well as theanonymous reviewers for their comments.

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