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Journal of Applied Phycology (2006) 18: 259–267
DOI: 10.1007/s10811-006-9025-y C Springer 2006
Farming of the giant kelp Macrocystis pyrifera in southern Chile for
development of novel food products
Alfonso Gutierrez1
, Tom´as Correa1
, Ver´onica Mu˜noz1
, Alejandro Santiba˜nez2
,
Roberto Marcos3
, Carlos C´aceres4
& Alejandro H. Buschmann1,∗
1
i∼mar, Universidad de Los Lagos, Camino Chinquihue km 6, Puerto Montt, Chile; 2
Departamento de Gobierno
y Empresas, Universidad de Los Lagos, Casilla 557, Puerto Montt, Chile; 3
Productos del Pac´ıfico, S.A. de C.V.,
Ensenada, M´exico; 4
Consultora Los Lagos, Poblaci´on Varmontt Nr. 4, Puerto Varas, Chile
∗
Author for correspondence: e-mail: abuschma@ulagos.cl; fax: +56 65 322418
Key words: Macrocystis pyrifera, farming, food products, Chile
Abstract
This study explores the potential cultivation of the giant kelp Macrocystis pyrifera (L.) C.A. Agardh in southern
Chile, for the development of novel food products. The study demonstrates the importance of considering the
collection site of the parent sporophytes for successful cultivation. This study also shows that the ropes must be
seeded with 10,000 to 40,000 spores ml−1
, depending on the culture method used. We also demonstrated that under
environmental conditions in southern Chile, the seeded ropes must be put at sea at the latest during autumn (April) in
order to reach the harvesting season in December. However, several other management aspects must be considered
to improve the quality of the product. Our final estimation indicates that over 14.4 kg m−1
of rope (fresh weight)
can be produced and from this total production, over 70% can reach the quality to produce different food products
that are already being introduced in oriental countries. The remaining 30% can be used for abalone feeding and is
also available for the organic fertilizer industry located in Chile.
Introduction
California kelp beds started to be harvested as a source
of potash during the first decade of the 20th century
and commercial interest in the giant kelp Macrocys-
tis pyrifera expanded significantly between the 1970s
and 1980s (Neushul, 1987; Druehl et al., 1988). This
interest was primarily for the production of alginates,
but also to produce biomass as a feedstock for methane
production as a consequence of the energy crisis at
that time (North et al., 1982; Gerard, 1987; Neushul
& Harger, 1987). Nevertheless, M. pyrifera commer-
cial cultivation for methane production was never a
reality. At present, the supply of M. pyrifera biomass
for the alginate industry relies exclusively on restora-
tion practices and management of natural beds to ob-
tain a sustainable production (North, 1979; McPeak &
Barilotti, 1993; V´asquez & McPeak, 1999). After the
energy crisis and because of the low price of alginates,
farming research on Macrocystis declined sharply.
On the other hand, other brown algal species be-
gan to be commercially cultivated in Japan, China and
Korea, mainly for human consumption (Tseng, 1987;
Kaneko, 1999; Hanisak, 1998) while kelp-farming at-
tempts for this purpose have also proved technically
feasible in other regions (e.g. Druehl et al., 1988;
Kain, 1991; Merril & Gillingham, 1991). Interestingly,
the demand for brown algae is also increasing due
to the introduction of new uses such as fertilizers,
cultivation for bioremediation purposes, and abalone
as well as sea urchin feeding among others (Petrell
et al., 1993; V´asquez & Vega, 1999; Buschmann et al.,
2001c; Ugarte & Sharp, 2001; Chopin et al., 2001). In
Chile, despite the commercial importance of various
algal species, aquaculture is still limited to the red alga
Gracilaria chilensis (Buschmann et al., 2001b).
260
In Chile, Abalone and sea urchin cultures, organic
fertilizer production and novel seafoods have created
a new niche market for the giant kelp Macrocystis
pyrifera. Increased harvesting is already causing some
deterioration of different kelp populations (V´asquez &
Vega, 1999). Considerable information on Macrocys-
tis cultivation has been published in the past (North,
1979). However, some basic knowledge necessary to
run a successful commercial activity is still lacking,
especially with regard to the different environmental
conditions and complex morphological and reproduc-
tion variability between populations, that can have im-
portant commercial consequences.
Considering this new market scenario, the potential
impact on natural populations and the lack of biological
knowledge necessary to produce a high quality prod-
uct, this paper deals with the cultivation of M. pyrifera
in southern Chile. Specifically, the effect of the ori-
gin of the parental plants on the survival and growth
of young sporophytes cultivated on ropes was tested,
in both hatchery and field conditions. Finally, a pilot
cultivation was established to determine the potential
yields of M. pyrifera in southern Chile and we describe
some of the food products developed.
Materials and methods
Study sites
Fertile sporophylls of Macrocystis pyrifera were col-
lected at six localities in southern Chile: Metri
(41◦
35 S; 72◦
42 W), Pargua (41◦
47 S; 73◦
25 W),
Calbuco (41◦
46 S; 73◦
08 W), Pucatrihue (40◦
33 S;
73◦
43 W), Bah´ıa Mansa (40◦
34 S; 73◦
44 W) and Cu-
raco de Velez (42◦
26 S; 73◦
35 W) (Figure 1). Site se-
lectionwasbasedonthepresenceofabundantkelppop-
ulations and different water movement conditions. The
plants were collected by scuba divers and transported,
within 6 h, on ice to the seaweed culture laboratory
in Metri. All field cultivation experiments were carried
out in Metri and the pilot culture in Calbuco (Figure 1).
Cultivation of different populations
The sporophylls collected in Metri, Pargua, Calbuco,
Pucatrihue and Bah´ıa Mansa, were washed under tap
water and UV treated filtered seawater (0.2 μm) con-
taining commercial iodine (0.5% for 10 s), packed in
filter paper, covered with aluminum foil and stored at
15 ◦
C (Figure 2A). After 12 h, 10 to 15 sporophylls
were placed in 20 L sterile plastic containers filled with
filtered (0.2 μm) and autoclaved seawater (Figure 2B).
Sporulation started in all cases after 25 to 35 min and,
after 1 h the sporophylls were removed and the water
was filtered with a 100 μm mesh. Eight PVC cylin-
ders covered with a 1.5 mm nylon rope were introduced
into each of the 20 L containers to allow for spore set-
tlement (Figure 2B). After 12 h, the eight PVC cylin-
ders were removed and placed in a 30 L glass tank
filled with autoclaved, filtered, and Provasoli enriched
seawater (McLachlan, 1973; Figure 2B). Culture was
carried out at a photon flux density of 30–40 μmol
m−2
s−1
; a temperature of 9–10◦
C; a salinity of 30%
and a pH of 7.8–7.9. Photoperiod was 16:8 (L:D) dur-
ing the first week; 14:10 (L:D) during the second week;
12:12 (L:D) during the third week and 10:14 (L:D)
thereafter (following a previously determined proto-
col; Buschmann unpublished results). After 44 days,
3 cm pieces of rope were randomly cut off and plant
density (number of sporophytes fronds per cm of rope
fragment) estimated under a stereomicroscope. Fur-
thermore, the maximum lengths of the juvenile sporo-
phytic fronds were determined using an ocular microm-
eter. The data were statistically analyzed by a one-way
ANOVA after logarithmic transformation to ensure the
normality and homocedasticity of the data. If signifi-
cant differences were detected between treatments, a
Tukey a posteriori-test (according to Steel and Torrie,
1985) was performed. As the data were obtained from
independent tanks, seeded from independent plants and
comparisons between times were not considered, no
pseudoreplication exists (sensu Hurlbert, 1984).
After 60 days in the hatchery (September), seeded
ropes were attached to a 3 m long horizontal supporting
rope (18 mm diameter) in groups of three placed at 2 m
depth (Figure 3) in Metri (Figure 1). Three seeded ropes
were used for each one of the five original populations.
Plant density and length of the different Macrocystis
pyrifera populations were estimated, after one and two
months in the field, by random sampling of 5 cm seeded
rope sections under a stereomicroscope. All data were
analyzed as above.
Pilot study
Parent sporophytes were collected in Curaco de Velez
(Figure 1) and seeded on ropes following the same
methods and culture conditions described above. The
ropes were seeded in mid January and were brought to
the Calbuco culture site (Figure 1) in March. The initial
culture conditions of the sporophytes in the field were
261
Figure 1. Map indicating the five collection sites of fertile tissues of Macrocystis pyrifera in southern Chile (Calbuco, Pargua, Metri, Pucatrihue
and Bah´ıa Mansa) and the experimental and pilot cultivation sites in Metri and Calbuco respectively.
1 mm plant length, and a mean density of 51 plants
mm−1
. The experiment was initiated in March using
the horizontal culture system at 1 m depth (Figure 3),
based on previous results (Buschmann, unpublished
data). The Macrocystis pyrifera fresh weight produced
per m of long-line was evaluated by taking 10 random
1.5 m rope samples. Total fresh weight was deter-
mined and then the blades, stipes and pneumatocyst
262
Figure 2. Scheme representing the pretreatment and sporulation induction procedure (A), and the seeding operation (B) for Macrocystis pyrifera
mass production. The procedure was modified from Merrill and Gillingham (1991).
Figure 3. Suspended culture method used in this study: horizontal
culture method adapted from Kawashima (1993) and Merrill and
Gillingham (1991).
and disposable parts (holdfast, necrotic, perforated,
and epiphyted tissues) of 22 plants obtained at ran-
dom, were separated and weighed individually. Fi-
nally, “substantiality” (the weight per frond area; sensu
Kawashima, 1993) of 30 randomly taken fronds was
determined by weighing 1 cm2
frond discs on a digital
balance (± 1 mg). Statistical analysis was performed
following the above-described protocol.
Results
Cultivation of different populations
In the hatchery, the mean number of sporophytes pro-
duced at 9–10◦
C, 40 μmol photons m−2
s−1
with a
variable photoperiod, varied between 1.0 and 12 indi-
viduals mm−1
of seeded rope (Figure 4A). The num-
ber of sporophytes produced was significantly higher
(p < 0.05) from fertile sporophytic tissues, which
were collected from Metri and Pucatrihue. Pargua and
Bah´ıa Mansa showed the lowest seeding success under
the same culture conditions, whereas Metri and Bah´ıa
Mansa showed the highest size increment, with mean
values of 2.85 mm in 44 days (4B).
Under field conditions, all the individuals produced
from plants collected in Bah´ıa Mansa and Pucatrihue
(wave exposed coast) showed a mortality of 100% after
only one month of cultivation (Figure 5). No significant
size difference (p > 0.1) was found between plants
originating from Calbuco, Pargua and Metri during the
first month of culture. After two months in culture, no
significant differences (p > 0.08) were found between
the studied populations (Figure 5), although Calbuco
plants showed a trend towards increased growth.
Pilot study
The pilot study carried out in Calbuco produced a
biomass of 14.4 kg m−1
(±4.8 kg m−1
; S.D.) after 7
months of culture, and showed a substantiality value
of the harvested fronds of 68.5 mg cm−2
(±1.6 mg
cm−2
; S.D.). The different tissue types of Macrocystis
pyrifera produced yields that varied from 1.3 kg m−2
(pneumatocyst) up to 4.1 kg m−2
(stipe) and 4.8 kg m−2
263
Figure 4. Cultivation of Macrocystis pyrifera under controlled conditions, initiated from fertile sporophytes collected in 5 sites of southern
Chile: Calbuco, Pargua, Metri, Pucatrihue and Bah´ıa Mansa. (A) Mean density (number mm−2) of sporophytes on nylon ropes; and (C) mean
length (mm) of sporophytes attached to nylon ropes after 44 days in the hatchery (9 replicates per location).
Figure 5. Cultivation of Macrocystis pyrifera at a depth of 2 m under field conditions, initiated from fertile sporophytes collected in 5 sites of
southern Chile: Calbuco, Pargua, Metri, Pucatrihue and Bah´ıa Mansa. Mean size (mm ± 1 SE) of sporophytes attached to nylon ropes after
one month (dotted bars) and two months (grey bars). Letters above the bars indicate no significant differences after a Tukey a posteriori-test
(p < 0.05). Asterisks indicate no survival of the germlings. Three samples were taken at each location.
264
Figure 6. Mean (± 1 SE) biomass production (A) and percent yield (B) of different tissues and low quality tissues of Macrocystis pyrifera after
seven months in a horizontal culture system installed in Calbuco. Twenty 1-m2 samples were taken.
Figure 7. Production schedule of Macrocystis pyrifera in southern Chile.
(frond) (Figure 6A). Almost 30% of the total biomass
was of lower quality and could not be used for food
(Figure 6B), as it was covered with hydrozoan and
bryozoan colonies, or necrotic tissues. Thus, Macro-
cystis pyrifera can be produced by collecting fertile
tissues from the natural environment during January
(summer), whereas rope seeding should be carried out
at the latest during February (Figure 7). The hatchery
phase takes at least 1.5 to 2 months, which means that
the suspended culture at sea can be initiated in April or
May (fall) and harvesting is possible in late November
or December (Figure 7).
Discussion
This study demonstrates that the cultivation of Macro-
cystis pyrifera in southern Chile is technically feasi-
ble and that a productivity of over 14 kg m−1
during
one production season can be obtained, which seems
very promising (Figures 8A and B). The comparison
of this value is not easy as most of the production esti-
mates are based on extrapolations from physiological
or growth measurements made on parts of kelp plants
(Neushul & Harger, 1987). Estimations of 7 g ash free
dry wt. m−2
day−1
(Neushul & Harger, 1987) and of
265
Table 1. Description of Macrocystis pyrifera food products developed.
Food product Product specification
Fresh Salted Frond (Figure 8 D) 1 × 3 cm salted blade pieces are washed in filtered
seawater and blanched at 100◦C for 1 min and then mixed with
salt (23%). The product has a green color and is placed in 15 or 10 kg plastic bags
Pneumatocyst Rings and Slices of pneumatocysts and 2-cm pieces of stipes are
Stipe Pieces (Figure 8C and E) washed and heated in vinegar with alcohol content not
higher than 1% and an acetic acid value not lower than 5%.
The products are packed in glass or packs as required.
The product must be maintained in cool conditions (2–6◦C)
Figure 8. (A) Pilot culture of Macrocystis in Calbuco, (B) harvesting Macrocystis and products developed for human consumption in Chile: (C)
stipe fragments; (D) blade strips; (E) pneumatocyst rings and (F) a general presentation plate of these M. pyrifera products.
4 wet kg per production period per m2
(Coon, 1982)
exist, suggesting that our results are encouraging. It
is important to mention that over 70% of the harvested
biomass can be used to produce high quality food prod-
ucts (Figures 8C, D, E and F; Table 1), which represents
a significantly higher success rate for the use proposed
in this study when compared to other uses. In addi-
tion, the introduction of massive kelp culture in Chile
may have other associated benefits such as providing a
means of removing nitrogen and phosphorus produced
by salmon farming (Buschmann et al., 2001a; Chopin
et al., 2001; Troell et al., 2003). It is important to re-
member that this area of Chile produces over 300,000
tonnesofsalmon,thuscreatingsignificantenvironmen-
tal impacts and conflicts (Buschmann, 2001).
To date, Macrocystis pyrifera has not been used
to produce food products in Japan. Traditionally the
species used as food in oriental countries are Laminaria
japonica (Kombu) and Undaria pinnatifida (Wakame)
(Kawashima,1993;Ohno&Matsuoka,1993).InChile,
the bull kelp Durvillaea antarctica has been tradition-
ally used as a food source, but has a very low price.
Because Laminaria and Undaria are not present in
Chile, the idea was to develop alternative products that
could be exported to oriental countries with a tradi-
tion in seaweed consumption (Abbott, 1996). However,
266
as the characteristics of color, texture, substantiality,
and mucilage content of Macrocystis cannot be com-
pared to those of Laminaria or Undaria we pursued
the commercial strategy of creating novel products
(Figure 8C to F), which received a positive market re-
sponse in Asia after a first dispatch and market tour.
Thus, because this is a developing market, it is believed
that the commercial cultivation of Macrocystis is pos-
sible in Chile. In addition, alternative use of this kelp
as abalone and sea urchin feed, or as organic fertilizer,
strengthens its economic feasibility. Lower grade parts
of Macrocystis can also be used for the other purposes
indicated above.
Recently, it has been demonstrated that Chilean
stocks of M. pyrifera show small genetic differences
from other Southern Hemisphere species, but stronger
differences from Californian stocks (Coyer et al.,
2001). Despite this low genetic diversity, plants col-
lected at various sites in southern Chile showed dif-
ferent potential for use in aquaculture practices. The
plants from the most exposed areas cannot be used
for farming in the inner seas, of southern Chile, al-
though they presented higher growth under hatchery
conditions. In contrast, marginal differences were ob-
served between M. pyrifera populations from the inner
seas. In contrast to an earlier report that identifies M.
laevis in this region (Aguilar-Rosas et al., 2003) and
indicates that some morphologically distinct plants ex-
ist, our six years of observations and experimentation
strongly suggest that this smooth bladed plants corre-
spond to M. pyrifera. Nevertheless, these results rec-
ognize that to some extent each farmer must consider
site-specific characteristics of his own licensed loca-
tion and the morphological characteristics of the parent
sporophytes, before starting commercial activities.
Despite the promising production results obtained
here, Macrocystis production in Chile still needs more
research. Several factors remain to be studied, but two
aspects are especially important, as has been demon-
strated for other algal resources (Wikfors & Ohno,
2001). Firstly, strain selection cannot be overlooked.
The manipulation of the growth capabilities of this
resource is important, but given that it is required to
produce food products, it is important to study as-
pects that can be used to produce plants with specific
morphological characteristics. Unpublished results in-
dicate that morphological variation and life-history
variability of M. pyrifera in southern Chile is high
(Buschmann et al., 2004), but we still do not know
how to manipulate morphological variation or how to
maintain certain morphological characteristics under
cultureconditions.Secondly,wewishtohighlightman-
agement requirements during the field phase that will
allow a successful commercial operation. During this
phase aspects such as plant density, harvesting strate-
gies, and disease control are extremely relevant, as can
be seen for example, during the intensive cultivation
of Laminaria japonica (Kawashima, 1993). All these
aspects remain unknown and will undoubtedly pro-
vide excellent material for future studies in southern
Chile.
Acknowledgments
This study was financed by a FONDEF grant
(D901/1101) and FONDECYT 1010706. The authors
acknowledgethehelpofMariamHern´andez-Gonz´alez,
Luis Fil´un, Ren´e Reyes, Rodrigo Mart´ınez and Ricardo
Ce˜na. The collaboration of Marcelo Brintrup is espe-
cially acknowledged, as well as the constructive criti-
cism to this manuscript given by D. Varela, R. Stead,
D. M. Luxton and an anonymous reviewer.
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Farming of the giant kelp macrocystis pyrifera in southern chile for

  • 1. Journal of Applied Phycology (2006) 18: 259–267 DOI: 10.1007/s10811-006-9025-y C Springer 2006 Farming of the giant kelp Macrocystis pyrifera in southern Chile for development of novel food products Alfonso Gutierrez1 , Tom´as Correa1 , Ver´onica Mu˜noz1 , Alejandro Santiba˜nez2 , Roberto Marcos3 , Carlos C´aceres4 & Alejandro H. Buschmann1,∗ 1 i∼mar, Universidad de Los Lagos, Camino Chinquihue km 6, Puerto Montt, Chile; 2 Departamento de Gobierno y Empresas, Universidad de Los Lagos, Casilla 557, Puerto Montt, Chile; 3 Productos del Pac´ıfico, S.A. de C.V., Ensenada, M´exico; 4 Consultora Los Lagos, Poblaci´on Varmontt Nr. 4, Puerto Varas, Chile ∗ Author for correspondence: e-mail: abuschma@ulagos.cl; fax: +56 65 322418 Key words: Macrocystis pyrifera, farming, food products, Chile Abstract This study explores the potential cultivation of the giant kelp Macrocystis pyrifera (L.) C.A. Agardh in southern Chile, for the development of novel food products. The study demonstrates the importance of considering the collection site of the parent sporophytes for successful cultivation. This study also shows that the ropes must be seeded with 10,000 to 40,000 spores ml−1 , depending on the culture method used. We also demonstrated that under environmental conditions in southern Chile, the seeded ropes must be put at sea at the latest during autumn (April) in order to reach the harvesting season in December. However, several other management aspects must be considered to improve the quality of the product. Our final estimation indicates that over 14.4 kg m−1 of rope (fresh weight) can be produced and from this total production, over 70% can reach the quality to produce different food products that are already being introduced in oriental countries. The remaining 30% can be used for abalone feeding and is also available for the organic fertilizer industry located in Chile. Introduction California kelp beds started to be harvested as a source of potash during the first decade of the 20th century and commercial interest in the giant kelp Macrocys- tis pyrifera expanded significantly between the 1970s and 1980s (Neushul, 1987; Druehl et al., 1988). This interest was primarily for the production of alginates, but also to produce biomass as a feedstock for methane production as a consequence of the energy crisis at that time (North et al., 1982; Gerard, 1987; Neushul & Harger, 1987). Nevertheless, M. pyrifera commer- cial cultivation for methane production was never a reality. At present, the supply of M. pyrifera biomass for the alginate industry relies exclusively on restora- tion practices and management of natural beds to ob- tain a sustainable production (North, 1979; McPeak & Barilotti, 1993; V´asquez & McPeak, 1999). After the energy crisis and because of the low price of alginates, farming research on Macrocystis declined sharply. On the other hand, other brown algal species be- gan to be commercially cultivated in Japan, China and Korea, mainly for human consumption (Tseng, 1987; Kaneko, 1999; Hanisak, 1998) while kelp-farming at- tempts for this purpose have also proved technically feasible in other regions (e.g. Druehl et al., 1988; Kain, 1991; Merril & Gillingham, 1991). Interestingly, the demand for brown algae is also increasing due to the introduction of new uses such as fertilizers, cultivation for bioremediation purposes, and abalone as well as sea urchin feeding among others (Petrell et al., 1993; V´asquez & Vega, 1999; Buschmann et al., 2001c; Ugarte & Sharp, 2001; Chopin et al., 2001). In Chile, despite the commercial importance of various algal species, aquaculture is still limited to the red alga Gracilaria chilensis (Buschmann et al., 2001b).
  • 2. 260 In Chile, Abalone and sea urchin cultures, organic fertilizer production and novel seafoods have created a new niche market for the giant kelp Macrocystis pyrifera. Increased harvesting is already causing some deterioration of different kelp populations (V´asquez & Vega, 1999). Considerable information on Macrocys- tis cultivation has been published in the past (North, 1979). However, some basic knowledge necessary to run a successful commercial activity is still lacking, especially with regard to the different environmental conditions and complex morphological and reproduc- tion variability between populations, that can have im- portant commercial consequences. Considering this new market scenario, the potential impact on natural populations and the lack of biological knowledge necessary to produce a high quality prod- uct, this paper deals with the cultivation of M. pyrifera in southern Chile. Specifically, the effect of the ori- gin of the parental plants on the survival and growth of young sporophytes cultivated on ropes was tested, in both hatchery and field conditions. Finally, a pilot cultivation was established to determine the potential yields of M. pyrifera in southern Chile and we describe some of the food products developed. Materials and methods Study sites Fertile sporophylls of Macrocystis pyrifera were col- lected at six localities in southern Chile: Metri (41◦ 35 S; 72◦ 42 W), Pargua (41◦ 47 S; 73◦ 25 W), Calbuco (41◦ 46 S; 73◦ 08 W), Pucatrihue (40◦ 33 S; 73◦ 43 W), Bah´ıa Mansa (40◦ 34 S; 73◦ 44 W) and Cu- raco de Velez (42◦ 26 S; 73◦ 35 W) (Figure 1). Site se- lectionwasbasedonthepresenceofabundantkelppop- ulations and different water movement conditions. The plants were collected by scuba divers and transported, within 6 h, on ice to the seaweed culture laboratory in Metri. All field cultivation experiments were carried out in Metri and the pilot culture in Calbuco (Figure 1). Cultivation of different populations The sporophylls collected in Metri, Pargua, Calbuco, Pucatrihue and Bah´ıa Mansa, were washed under tap water and UV treated filtered seawater (0.2 μm) con- taining commercial iodine (0.5% for 10 s), packed in filter paper, covered with aluminum foil and stored at 15 ◦ C (Figure 2A). After 12 h, 10 to 15 sporophylls were placed in 20 L sterile plastic containers filled with filtered (0.2 μm) and autoclaved seawater (Figure 2B). Sporulation started in all cases after 25 to 35 min and, after 1 h the sporophylls were removed and the water was filtered with a 100 μm mesh. Eight PVC cylin- ders covered with a 1.5 mm nylon rope were introduced into each of the 20 L containers to allow for spore set- tlement (Figure 2B). After 12 h, the eight PVC cylin- ders were removed and placed in a 30 L glass tank filled with autoclaved, filtered, and Provasoli enriched seawater (McLachlan, 1973; Figure 2B). Culture was carried out at a photon flux density of 30–40 μmol m−2 s−1 ; a temperature of 9–10◦ C; a salinity of 30% and a pH of 7.8–7.9. Photoperiod was 16:8 (L:D) dur- ing the first week; 14:10 (L:D) during the second week; 12:12 (L:D) during the third week and 10:14 (L:D) thereafter (following a previously determined proto- col; Buschmann unpublished results). After 44 days, 3 cm pieces of rope were randomly cut off and plant density (number of sporophytes fronds per cm of rope fragment) estimated under a stereomicroscope. Fur- thermore, the maximum lengths of the juvenile sporo- phytic fronds were determined using an ocular microm- eter. The data were statistically analyzed by a one-way ANOVA after logarithmic transformation to ensure the normality and homocedasticity of the data. If signifi- cant differences were detected between treatments, a Tukey a posteriori-test (according to Steel and Torrie, 1985) was performed. As the data were obtained from independent tanks, seeded from independent plants and comparisons between times were not considered, no pseudoreplication exists (sensu Hurlbert, 1984). After 60 days in the hatchery (September), seeded ropes were attached to a 3 m long horizontal supporting rope (18 mm diameter) in groups of three placed at 2 m depth (Figure 3) in Metri (Figure 1). Three seeded ropes were used for each one of the five original populations. Plant density and length of the different Macrocystis pyrifera populations were estimated, after one and two months in the field, by random sampling of 5 cm seeded rope sections under a stereomicroscope. All data were analyzed as above. Pilot study Parent sporophytes were collected in Curaco de Velez (Figure 1) and seeded on ropes following the same methods and culture conditions described above. The ropes were seeded in mid January and were brought to the Calbuco culture site (Figure 1) in March. The initial culture conditions of the sporophytes in the field were
  • 3. 261 Figure 1. Map indicating the five collection sites of fertile tissues of Macrocystis pyrifera in southern Chile (Calbuco, Pargua, Metri, Pucatrihue and Bah´ıa Mansa) and the experimental and pilot cultivation sites in Metri and Calbuco respectively. 1 mm plant length, and a mean density of 51 plants mm−1 . The experiment was initiated in March using the horizontal culture system at 1 m depth (Figure 3), based on previous results (Buschmann, unpublished data). The Macrocystis pyrifera fresh weight produced per m of long-line was evaluated by taking 10 random 1.5 m rope samples. Total fresh weight was deter- mined and then the blades, stipes and pneumatocyst
  • 4. 262 Figure 2. Scheme representing the pretreatment and sporulation induction procedure (A), and the seeding operation (B) for Macrocystis pyrifera mass production. The procedure was modified from Merrill and Gillingham (1991). Figure 3. Suspended culture method used in this study: horizontal culture method adapted from Kawashima (1993) and Merrill and Gillingham (1991). and disposable parts (holdfast, necrotic, perforated, and epiphyted tissues) of 22 plants obtained at ran- dom, were separated and weighed individually. Fi- nally, “substantiality” (the weight per frond area; sensu Kawashima, 1993) of 30 randomly taken fronds was determined by weighing 1 cm2 frond discs on a digital balance (± 1 mg). Statistical analysis was performed following the above-described protocol. Results Cultivation of different populations In the hatchery, the mean number of sporophytes pro- duced at 9–10◦ C, 40 μmol photons m−2 s−1 with a variable photoperiod, varied between 1.0 and 12 indi- viduals mm−1 of seeded rope (Figure 4A). The num- ber of sporophytes produced was significantly higher (p < 0.05) from fertile sporophytic tissues, which were collected from Metri and Pucatrihue. Pargua and Bah´ıa Mansa showed the lowest seeding success under the same culture conditions, whereas Metri and Bah´ıa Mansa showed the highest size increment, with mean values of 2.85 mm in 44 days (4B). Under field conditions, all the individuals produced from plants collected in Bah´ıa Mansa and Pucatrihue (wave exposed coast) showed a mortality of 100% after only one month of cultivation (Figure 5). No significant size difference (p > 0.1) was found between plants originating from Calbuco, Pargua and Metri during the first month of culture. After two months in culture, no significant differences (p > 0.08) were found between the studied populations (Figure 5), although Calbuco plants showed a trend towards increased growth. Pilot study The pilot study carried out in Calbuco produced a biomass of 14.4 kg m−1 (±4.8 kg m−1 ; S.D.) after 7 months of culture, and showed a substantiality value of the harvested fronds of 68.5 mg cm−2 (±1.6 mg cm−2 ; S.D.). The different tissue types of Macrocystis pyrifera produced yields that varied from 1.3 kg m−2 (pneumatocyst) up to 4.1 kg m−2 (stipe) and 4.8 kg m−2
  • 5. 263 Figure 4. Cultivation of Macrocystis pyrifera under controlled conditions, initiated from fertile sporophytes collected in 5 sites of southern Chile: Calbuco, Pargua, Metri, Pucatrihue and Bah´ıa Mansa. (A) Mean density (number mm−2) of sporophytes on nylon ropes; and (C) mean length (mm) of sporophytes attached to nylon ropes after 44 days in the hatchery (9 replicates per location). Figure 5. Cultivation of Macrocystis pyrifera at a depth of 2 m under field conditions, initiated from fertile sporophytes collected in 5 sites of southern Chile: Calbuco, Pargua, Metri, Pucatrihue and Bah´ıa Mansa. Mean size (mm ± 1 SE) of sporophytes attached to nylon ropes after one month (dotted bars) and two months (grey bars). Letters above the bars indicate no significant differences after a Tukey a posteriori-test (p < 0.05). Asterisks indicate no survival of the germlings. Three samples were taken at each location.
  • 6. 264 Figure 6. Mean (± 1 SE) biomass production (A) and percent yield (B) of different tissues and low quality tissues of Macrocystis pyrifera after seven months in a horizontal culture system installed in Calbuco. Twenty 1-m2 samples were taken. Figure 7. Production schedule of Macrocystis pyrifera in southern Chile. (frond) (Figure 6A). Almost 30% of the total biomass was of lower quality and could not be used for food (Figure 6B), as it was covered with hydrozoan and bryozoan colonies, or necrotic tissues. Thus, Macro- cystis pyrifera can be produced by collecting fertile tissues from the natural environment during January (summer), whereas rope seeding should be carried out at the latest during February (Figure 7). The hatchery phase takes at least 1.5 to 2 months, which means that the suspended culture at sea can be initiated in April or May (fall) and harvesting is possible in late November or December (Figure 7). Discussion This study demonstrates that the cultivation of Macro- cystis pyrifera in southern Chile is technically feasi- ble and that a productivity of over 14 kg m−1 during one production season can be obtained, which seems very promising (Figures 8A and B). The comparison of this value is not easy as most of the production esti- mates are based on extrapolations from physiological or growth measurements made on parts of kelp plants (Neushul & Harger, 1987). Estimations of 7 g ash free dry wt. m−2 day−1 (Neushul & Harger, 1987) and of
  • 7. 265 Table 1. Description of Macrocystis pyrifera food products developed. Food product Product specification Fresh Salted Frond (Figure 8 D) 1 × 3 cm salted blade pieces are washed in filtered seawater and blanched at 100◦C for 1 min and then mixed with salt (23%). The product has a green color and is placed in 15 or 10 kg plastic bags Pneumatocyst Rings and Slices of pneumatocysts and 2-cm pieces of stipes are Stipe Pieces (Figure 8C and E) washed and heated in vinegar with alcohol content not higher than 1% and an acetic acid value not lower than 5%. The products are packed in glass or packs as required. The product must be maintained in cool conditions (2–6◦C) Figure 8. (A) Pilot culture of Macrocystis in Calbuco, (B) harvesting Macrocystis and products developed for human consumption in Chile: (C) stipe fragments; (D) blade strips; (E) pneumatocyst rings and (F) a general presentation plate of these M. pyrifera products. 4 wet kg per production period per m2 (Coon, 1982) exist, suggesting that our results are encouraging. It is important to mention that over 70% of the harvested biomass can be used to produce high quality food prod- ucts (Figures 8C, D, E and F; Table 1), which represents a significantly higher success rate for the use proposed in this study when compared to other uses. In addi- tion, the introduction of massive kelp culture in Chile may have other associated benefits such as providing a means of removing nitrogen and phosphorus produced by salmon farming (Buschmann et al., 2001a; Chopin et al., 2001; Troell et al., 2003). It is important to re- member that this area of Chile produces over 300,000 tonnesofsalmon,thuscreatingsignificantenvironmen- tal impacts and conflicts (Buschmann, 2001). To date, Macrocystis pyrifera has not been used to produce food products in Japan. Traditionally the species used as food in oriental countries are Laminaria japonica (Kombu) and Undaria pinnatifida (Wakame) (Kawashima,1993;Ohno&Matsuoka,1993).InChile, the bull kelp Durvillaea antarctica has been tradition- ally used as a food source, but has a very low price. Because Laminaria and Undaria are not present in Chile, the idea was to develop alternative products that could be exported to oriental countries with a tradi- tion in seaweed consumption (Abbott, 1996). However,
  • 8. 266 as the characteristics of color, texture, substantiality, and mucilage content of Macrocystis cannot be com- pared to those of Laminaria or Undaria we pursued the commercial strategy of creating novel products (Figure 8C to F), which received a positive market re- sponse in Asia after a first dispatch and market tour. Thus, because this is a developing market, it is believed that the commercial cultivation of Macrocystis is pos- sible in Chile. In addition, alternative use of this kelp as abalone and sea urchin feed, or as organic fertilizer, strengthens its economic feasibility. Lower grade parts of Macrocystis can also be used for the other purposes indicated above. Recently, it has been demonstrated that Chilean stocks of M. pyrifera show small genetic differences from other Southern Hemisphere species, but stronger differences from Californian stocks (Coyer et al., 2001). Despite this low genetic diversity, plants col- lected at various sites in southern Chile showed dif- ferent potential for use in aquaculture practices. The plants from the most exposed areas cannot be used for farming in the inner seas, of southern Chile, al- though they presented higher growth under hatchery conditions. In contrast, marginal differences were ob- served between M. pyrifera populations from the inner seas. In contrast to an earlier report that identifies M. laevis in this region (Aguilar-Rosas et al., 2003) and indicates that some morphologically distinct plants ex- ist, our six years of observations and experimentation strongly suggest that this smooth bladed plants corre- spond to M. pyrifera. Nevertheless, these results rec- ognize that to some extent each farmer must consider site-specific characteristics of his own licensed loca- tion and the morphological characteristics of the parent sporophytes, before starting commercial activities. Despite the promising production results obtained here, Macrocystis production in Chile still needs more research. Several factors remain to be studied, but two aspects are especially important, as has been demon- strated for other algal resources (Wikfors & Ohno, 2001). Firstly, strain selection cannot be overlooked. The manipulation of the growth capabilities of this resource is important, but given that it is required to produce food products, it is important to study as- pects that can be used to produce plants with specific morphological characteristics. Unpublished results in- dicate that morphological variation and life-history variability of M. pyrifera in southern Chile is high (Buschmann et al., 2004), but we still do not know how to manipulate morphological variation or how to maintain certain morphological characteristics under cultureconditions.Secondly,wewishtohighlightman- agement requirements during the field phase that will allow a successful commercial operation. 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