This document discusses the development of techniques for cultivating Lessonia trabeculata in Chile. Small sporophytes of L. trabeculata were propagated in the laboratory from reproductive blades and cultured on different substrates like nylon cord and PVC tubes. The algae were grown in outdoor tanks and then transferred to long lines in the ocean, where individual plants reached up to 1.7 kg in mass. When tested, abalone grew as well on the cultured L. trabeculata as on naturally occurring algae, demonstrating its potential as a sustainable food source for abalone aquaculture.

1. Development of techniques for the cultivation
of Lessonia trabeculata Villouta et Santelices
(Phaeophyceae: Laminariales) in Chile
Mario E Edding & Fadia B Tala
Departamento de Biolog|¤a Marina, Facultad de Ciencias del Mar, Universidad Cato¤ lica del Norte, Coquimbo, Chile
Correspondence: Mario E Edding, Departamento de Biolog|¤a Marina, Facultad de Ciencias del Mar, Universidad Cato¤ lica del Norte,
Casilla117, Coquimbo, Chile. E-mail: medding@ucn.cl
Abstract
Large quantities of brown algae have traditionally
been exported from Chile as a raw material, of which
Lessonia spp. has amounted to over 130 000 tons an-
nually since1995. To the export demand has recently
been added the new demand for high-quality
Lessonia spp. as a foodstu¡ for the expanding abalone
culture industry in Chile. The present study is based
on e¡orts to produce signi¢cant quantities of Lessonia
trabeculata in long-line culture as food for tank-
cultured Haliotis rufescens Swainson and Haliotis
discus-hannae Ino, which accept it as an excellent
source of nutrition. Small sporophytes of L. trabecula-
ta were propagated in the laboratory from reproduc-
tive blades harvested by diving near Coquimbo
(301S). The best culture substrate was polyvinyl
chloride (PVC) in small pieces, inserted into the nylon
cord for ¢nal culturing. Enrichment of seawater with
agricultural-grade fertilizer produced no di¡erences
in growth and development of the Lessonia compared
with results obtained using Provasoli medium. Spor-
ophytes 1^2 cm in length cultured on 12-mm cord
were transferred to outdoor tanks with circulating
sea water and strong aeration where they were out-
grown to 15^20 cm length; at this size, they were
transferred to a 50-m long line in the ocean. In a
1-year period, individual plants reached up to 1.7 kg
in mass, with average values per cord of about 9 kg.
Total production from the long line was about
500 kg fresh weight of the alga. In comparative test-
ing, H. discus hannae grew as well on the cultured
algae as on naturally occurring L. trabeculata.
Keywords: abalone culture, algae culture, algal
propagation, Chile, Lessonia, long-line culture
Introduction
The use of marine algae as food for humans and mar-
ine organisms has an extensive history in Asia (Ohno
& Critchley1993). Harvesting of such algae from nat-
ural beds has been replaced by the arti¢cial culture of
numerous marine algal species on di¡erent coasts
around the world (de Oliveira & Kautsky 1990; Ohno
& Critchley1993). Some of the better knownalgal cul-
tures include Laminaria in China (Tseng 1987) and
Japan (Kawashima1984), Undaria inJapan and Korea
(Ohno & Matsuoka 1993), Porphyra in Japan, Korea
and China (Oohusa 1993), Eucheuma in the Philip-
pines (Trono 1993), Chondrus in Canada (Craigie &
Shacklock 1989) and Gracilaria in Chile (Santelices &
Ugarte1987). E¡orts to culture these algae are a con-
sequence of an increase in commercial demands and
concern for maintaining the natural equilibrium
within the ecosystem from which they are harves-
ted. Culture of Laminariales in Chile is recent
and has been developed in university research
with experimental cultures of Lessonia trabeculata
(Edding,Venegas, Orrego & Fonck1990).
The feeding of cultured herbivores, such as lim-
pets, sea urchins and abalone, requires di¡erent
types of algae as food, depending on the growth stage
of the grazer (Corazani & Illanes 1998; Serviere,
Go¤ mez & Ponce 1998). As their feeding apparatus
matures, herbivores switch from the initial stages in
which they graze on benthic microalgae to the stages
whenthey begintoaccept macroalgal thalli (Corazani
& Illanes1998; Serviere et al.1998).
In the feeding of commercially cultured organisms
such as abalone, signi¢cant amounts of algae are
required, as they may consume between 10% and
30% of their body weight in algae daily (Corazani
Aquaculture Research, 2003, 34, 507^515
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2. & Illanes 1998; Serviere et al. 1998). Parallel harvest-
ing e¡orts are required to provide these amounts of
algae for invertebrate cultures. Experimentation in
Chile has demonstrated that the alga L. trabeculata is
one of the best sources of food for tank-cultured aba-
lone, producing high growth rates (Owen, DiSalvo,
Ebert & Fonck 1984; Maureira, Takeda & Martinez
1993; Castillo 2000). Unsustainable harvest of algal
resources leads to a decrease in their populations by
overexploitation, an example of which is the case of
Gracilaria chilensis Bird, McLachlan & Oliveira in
Chile, which occurred during the 1970s (Santelices
& Ugarte 1987). Flora and fauna co-occurring in the
macroalgal communities may also be at risk from
extensive algal harvesting.
In addition to protecting macroalgae as a renew-
able resource, the culture of seaweeds would be a
strategy for price stabilization at acceptable levels, as
the price of the algae harvested from natural beds by
local divers is governed by prices in the world algi-
nate market. In Chile, regulations are in place that
permit associations of local ¢shermen to exploit,
manage and culture marine resources within‘Artisa-
nal Fishery Reserve Areas’ (ARPA) from the beach to
5 miles o¡shore. Culture of algae could provide a
stable source of the product for invertebrate culture
systems with ¢xed demands, more stable employ-
ment for local workers and a more reliable product
in terms of quality and quantity. A stable source of
algae would also allow for better management of
feeding regimes for cultured organisms (Basuyaux &
Mathieu1999) making growth more predictable.
The present study evaluates the technical feasibil-
ityof carrying out large-scale culture of L. trabeculata
as a complementary step to abalone culture.
Materials and methods
Sporulation and seeding
Release of spores was carried out using the method
described by Edding et al. (1990) for L. trabeculata,
following the additional recommendations of Fonck,
Venegas,Tala & Edding (1998) for this species. Repro-
ductive fronds were collected from a population
located on the Tongoy Peninsula (301150
S; 711300
W)
in north-central Chile. The collection dates of the
algal material are presented below, and collection
was always carried out 1 day before initiation of cul-
tures. In all case, the fronds were transported to the
laboratory in humid conditions in the dark, at a
temperature similar to that of their habitat. Immedi-
ately on arrival at the laboratory, the fronds were
rinsed with fresh water and drained in the dark for
6 h at1571 1C.The fronds (n 5 70^100) were then in-
duced to sporulate in 3-L plastic trays containing1.5 L
of 0.45 mm for E90 min. About10 fronds were placed
horizontally in each tray with water just covering the
reproductive tissue. Spore-containing water was
decanted in the dark for 30 min at1571 1C; the lower
one-third of the liquid was discarded in order to
reduce contamination of the cultures by diatoms.
The spore-containing water was then ¢ltered
through sterile gauze to remove contaminants and
seeded over arti¢cial substrates described below to
allow settlement of spores. Spore concentration was
determined using a haemocytometer.
Inthe initial phase, settled spores were maintained
in a controlled environmental chamber at 1571 1C
with a12 h:12 h light/dark photoperiod and light irra-
diance of 90 mmol photons m^2
s^1
with constant
aeration. Sea water for the cultures was enriched
using 0.114 g L^1
agriculture-grade fertilizer (potas-
sium nitrate and diammonium phosphate) at a ratio
of 23:1 (N:P) as in Provasoli medium (Starr & Zeikus
1993); this water was replaced weekly. As these re-
agents were not of analytical grade, their trace ele-
ment content was unknown.
Development of the microscopic phase of L. trabe-
culata was observed in parallel with development on
arti¢cial substrates by culturing in Petri dishes with
spores from the same suspension used to seed the
substrates for the mass culture. Observations were
made weekly of developing microscopic stages of the
alga in both Petri dishes and scrapings from the arti-
¢cial substrates using a stereoscopic microscope at
200 Â magni¢cation.
Substrate types and con¢gurations
The ¢rst spore seedings were carried out ontwo types
of spore collector, the ¢rst including rectangular
polyvinyl chloride (PVC) tubing frames of 60 Â80
cm, each of which was wrapped with 6-m lengths of
12-mm-diameter nylon cord. A total of10 PVC collec-
tors were prepared, each supporting10 nyloncords to
give a total of 600 m of spore-seeded cords. The sec-
ond type of collector was a galvanized wire frame
60 Â70 cm (n 510) covered with plastic mesh with
1-mm openings. Seeding was carried out on these
substrates placed horizontally in seeding tanks.After
an initial seeding period of 24 h, the sea water was
changed, and culture was initiated in tanks with the
collectors in the vertical position.
Lessonia trabeculata cultivation M E Edding & FB Tala Aquaculture Research, 2003, 34, 507^515
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3. Another experiment was conducted to determine
whether spores were capable of settling on a solid
substrate such as PVC. The experiment was carried
out on a small scale in six 5-L sea-water aquaria.
Each aquarium received six 20-mm-diameter PVC
tubes, each 20 cm in length.This seeding was carried
out in three aquaria containing the tubes in a hori-
zontal position, whereas in another three aquaria,
the tubes were exposed to seeding in the vertical
position. After sporulation, each aquarium received
725 mL of spore suspensioncontaining 21000 spores
mL^1
, and the aquarium was then ¢lled with ¢ltered,
sterilized sea water as above. After a 24-h seeding
period, water in the aquaria was changed, and all
tubes were maintained in the vertical position for
culture. Based on the success of this experiment (see
Results), subsequent spore settling was carried out in
20-L aquaria with 90 PVC tubes (as above) placed
horizontally.
Experiments with varying nutrient
enrichment
As the initial tests on culture of Lessonia produced
slow development and growth of the sporophytes,
an experiment was carried out to evaluate di¡erent
nutrient regimes for the procedure in June 1998.
Comparative testing of spore growth and develop-
ment was carried out using the following nutrient
conditions:
sea water control (no enrichment);
sea water enriched with 100% Provasoli medium
(Starr Zeikus1993);
sea water enriched with 100% Provasoli medium
plus iodine;
sea water enriched with agricultural fertilizer
(0.114 g L^1
).
Sea water enriched with agricultural
fertilizer plus iodine
The source of iodine was KI at 0.4% (wt/vol) made up
in Provasoli stock solution before its addition to the
sea water. The agricultural-grade fertilizer was
potassium nitrate and diammonium phosphate, with
an N:P ratio of 23:1.
The experiment was carried out with spores re-
leased using the same methodology as described
above with seeding into Petri dishes. Variables
recorded included percentage germination after
7 days, sexual di¡erentiation of the gametophytes
after15 days and percentage fertilityof the female ga-
metophytes on day 25 (Fonck et al.1998).
The same experiment was carried out using juve-
nile sporophytes (o2.5 mm frond length) produced
in culture, using 30 individuals for each nutrient
regime, maintained under the same culture condi-
tions as the spores. Total lengths of plantlets were
measured at the beginning of the experiment and
after 15 days in culture, using an ocular micrometer
in a stereoscopic microscope. Daily growth was
expressed as a percentage compared with the mean
initial size.
Culture experiment in outdoor tanks
Once sporophyte plantlets were grown under con-
trolled conditions and were visible to the naked eye
(E3 months), nylon cords with juvenile sporophytes
were placed in 1000-L outdoor circulating sea-water
(150 L h^1
) tanks with constant aeration in prepara-
tion for culture at sea. This treatment favoured the
development of the algae, primarily in length of the
fronds and attachment to the substrate.Time of culti-
vation in these tanks varied with development of
the plantlets, with the substrates removed to sea
at irregular intervals according to their degree of
development.
The rate of elongation of the tank-cultured plants
was measured using the method of Parker (1948) as
modi¢ed by Edding et al. (1990). The tanks were
enriched weekly with 0.114 g L^1
agriculture-grade
fertilizer (N:P 23:1) and shaded with sunscreen
netting, which reduced light intensity by about 50%
to prevent excessive development of epiphytes.
Culture experiment at sea
Culture at sea was carried out using a long-line sys-
tem, based on the design described by Kawashima
(1984) for Japanese Laminariales and later modi¢ed
for L. trabeculata by Edding et al. (1990). A 50-m line
was used (20 mm diameter) with hanging seaweed
cultivation ropes (12 mm diameter,6 m length) every
1 m. The main line was held at 1-m depth, using
plastic buoys.
During March 1998, 26 cords carrying a total of
296 juvenile sporophytes of L. trabeculata (1278
plants per cord) with frond lengths from 5 to
150 mm were installed within the marine concession
managed by the UCN in Herradura Bay, Coquimbo
(291590
S). This ¢rst group of cords was not precul-
tured in outdoor tanks. A second group of 30 cords,
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4. whichwas inoculated inJune1998 and had remained
in outdoor tanks during the summer of 1999, was
placed at sea in three stages (March, April and July
1999). A third group of 10 cords was maintained
within the outdoor tanks for comparison with algae
placed at sea. Algae maintained at sea were routinely
measured for elongation of fronds according to the
methods described above. Cords and algae were peri-
odically cleaned of fouling, and algal tissue densely
covered with epiphytes was removed.
Experiment on the feeding of abalone with
L. trabeculata
Juveniles of the abalone Haliotis discus hannai were
fed with laboratory-propagated L. trabeculata in order
to compare the nutritional value of the cultured al-
gae with that of naturally harvested L. trabeculata.
Test abalone were obtained from cultures at the UCN
shell¢sh hatchery. Three feeding regimes were tested
on groups of 80 juvenile abalone with two replicates
each, all maintained in separate 60-L sea-water
tanks. All abalone were from the same production
cohort, with an average initial size of 9 mm maxi-
mum shell length and 6 mm width, with a fresh,
drained weight of 85 mg each. The feeding regimes
included: (a) L. trabeculata harvested from natural
beds (control); (b) the same species from long-line
culture; and (c) L. trabeculata cultured in outdoor
tanks and enriched with agricultural fertilizer. Feed-
ing was carried out once a week, using a quantity
equal to 50% of the body weight of the abalone test
population. The experiment had a duration of 2
months, in which all abalone were measured and
weighed at the beginning and end of the test period.
Growth was expressed as the monthly percentage in-
crement with respect to initial measurements for
each variable.
Results
Substrate types
Initial settlement of sporophytes on the cord and
plastic mesh substrates from the ¢rst culture carried
out in May 1997 were unfavourable, based on the
high degree of detachment of the microscopic game-
tophyte and sporophyte embryos. A high degree of
contamination by microorganisms, mainly diatoms
and Protozoa, was observed, which overgrewor even
consumed the gametophytes.
When PVC sections were used as a substrate in
March1998, their surfaces appeared brown in colour
1month after inoculation, indicating the presence of
the microscopic phases of settled L. trabeculata. After
1.5 months, the ¢rst microscopic sporophytes could
be detected. After 3.5 months a sample of 150 sporo-
phytes had a mean length of 1.6970.57 mm. PVC
tubes seeded in the vertical position demonstrated
poor ¢xation of sporophytes, whereas those exposed
in the horizontal position produced a high degree of
settlement.
Based on the preceding results, a culture was car-
ried out inJune1998 using 20-cm PVC tubes that had
been seeded and cultivated inthe horizontal position.
The tubes were placed undisturbed on the bottoms of
the aquaria. Settlement of spores was apparentlyuni-
form on the PVC tubes, as sporophyte plantlets devel-
oped uniformly over the tube surfaces. Subsequently,
during October1998, PVC tubes set with L. trabeculata
were cut into subsamples 3 cm in length.These rings
were threaded onto the ¢nal culture lines, at 30 cm
intervals, placing seven or eight pieces on lines 6 m
in length. About 90 such lines were prepared and
maintained in outdoor culture tanks during the
three summer months. During this period, the plants
grew to sizes of 0.5^3 cm in length. Over the culture
period, the PVC tubes became adherent to the line be-
cause of the growth of the algae.
Experiment with varying nutrient enrichment
Table 1 shows that, at the microscopic scale,
fewer spores germinated and a low degree of sexual
di¡erentiation was obtained for L. trabeculata
gametophytes in the control treatment, di¡ering
signi¢cantly (one-way ANOVA, Po0.001) from the
other treatments. Signi¢cantly highest (one-way
ANOVA, P o 0.001) fertilities were observed using the
agricultural fertilizer as nutrients (Table1). Low den-
sities (o10%) of microscopic sporophytes could be
observed after 25 days of culture in the Provasoli 1
I, agricultural fertilizer, and agricultural fertilizer 1I
treatments.
The experiment using nutrients with juvenile
sporophytes did not show signi¢cant di¡erences
(Kruskal^Wallis, P40.05) in growth between treat-
ments (Table 2). Some juvenile sporophytes showed
negative growth because of loss of apical tissue.
Culture experiment in outdoor tanks
The growth of algae in tanks during the summer of
1999 produced a mean frond elongation of 1.670.8
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5. mm day^1
. Range in frond length varied between 5
and 10 cm and was 420 cm in 8 months (Fig. 2).
Growth of the fronds occurred primarily in the zone
between the stipe and the blade.
Algae maintained in tanks showed less frond
growth than those cultured at sea (Figs1and 2), and
elongation of the fronds in the tanks was stable over
time without showing the seasonal di¡erences
observed in frond growth at sea (Fig.1).
Culture experiment at sea
The main problem detected in culture at sea was
the high degree of biofouling observed on the lines
Table1 Meanand standard deviationvalues forgermination, gametophyte sex ratioand fertilityof Lessonia trabeculata in ¢ve
nutrient regimes
Germination Gametophyte proportions Fertility
Treatment (%) (female/male) (%)
Control 83.875.16 No differentiation 0
Provasoli 91.773.01 1.770.38 28.5377.52
Provasoli 1 I 90.174.19 1.670.26 51.85716.27
N* 90.274.18 1.870.25 60.62713.03
N* 1 I 90.673.35 1.570.27 60.0274.73
*Agricultural fertilizer (potassium nitrate 1 diammonium phosphate).
I, iodine (KI).
Table 2 Mean and standard deviation for lengths of Lessonia trabeculata sporophytes from laboratoryculture with ¢ve di¡er-
ent nutrient regimes, and growth [(100 Lf/Li)/15 days]
Treatment
Initial length
(mm)
Length after
15 days (mm)
Growth
(% day^1
)
Sea water 1.3570.49 1.6370.47 1.3371.31
Provasoli 1.6570.54 2.2870.75 2.3971.94
Provasoli 1 I 1.6470.43 2.3070.56 2.1371.75
N* 2.0270.53 2.4470.64 1.8771.54
N* 1 I 1.8770.68 2.4070.69 1.4871.31
*Agricultural fertilizer (potassium nitrate 1 diammonium phosphate).
I, iodine (KI).
Figure1 Frond elongation rate over time (mm day^1
) of
Lessonia trabeculata cultured in outdoor tanks (E) and on
long-line systems at sea (I, II and III) after tank culture
during1999.
Figure 2 Variation in average length of the fronds over
time (cm) in Lessonia trabeculata cultured in outdoor
tanks (E) and on long-line systems at sea (I, II and III)
after tank culture during1999.
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6. within 1 month of their installation. Smaller sporo-
phytes were overcome by the growth of ascidians
(mostly Ciona) and hydrozoans, which precipitated
the death and decomposition of the algae. Plants that
survived were those with the greatest length at the
time of their transfer to sea (410 cm), although even
these plants became heavily encrusted with epi-
bionts, which had to be removed each time the plants
were measured.
The group of algae placed at sea without inter-
mediate culture in outdoor tanks experienced high
mortality in their initial months of culture. Of a total
of 26 cords (plant n 5 296) installed on the long line
in March 1998, only 105 plants remained among se-
ven cords (65% mortality). After 5 months, only 22
plants remained on three cords, representing a mor-
tality of 79%. It was impossible to measure the
growth of these plants successfully because of the
high degree of epiphytism and mortality.
Figures 1 and 2 show the rates of elongation and
sizes of fronds of plants placed at sea at di¡erent sea-
sons of the year after their culture in the outdoor
tanks. All plants were from a single cohort as de-
scribed in the Materials and methods. Elongation of
fronds occurred during the spring months and was
similar among the di¡erent groups of cords, with va-
lues ranging from 2 to 6 mm day^1
(Fig.1).The fronds
showed gradual growth, reaching lengths of 50 cm
during October (Fig.2). Complete harvest of the cords
(n 510) in November1999 gave a meanvalue for total
fresh drained weight of 972.6 kg per cord.
Formation of reproductive tissue in the fronds was
¢rst observed in August 1999 and did not depend on
the date when the individual cord was placed at sea.
This phenomenon was not observed in plants main-
tained in the outdoor tanks. Although quantitative
data were not obtained, an informal laboratory test
suggested that spores from the cultured algae were
viable and capable of producing sporophytes.
Experiment on the feeding of abalone with
L. trabeculata
Table 3 presents the monthly mean increments in
morphometric characteristics measured on abalone
fed three di¡erent diets of Lessonia described in Mate-
rials and methods. Analysis of variance (ANOVA)
showed no signi¢cant di¡erences (P40.05) among
the growth values measured in the abalone among
the food treatments.
Discussion
Early experimentation with microscopic gameto-
phytes and sporophytes produced mixed results
when seeding cords because of the high degree of
detachment. This may have resulted from the micro-
structure of the nylon cord used, as its ¢ne ¢lamen-
tous structure underwent constant £exion as the
cord was stressed under environmental conditions.
This cord did not present a stable base for the attach-
ment of the algal haptera. Settlement of the spores on
solid substrates such as PVC provided a viable
solution to this problem.
Agricultural nutrients used during the cultures
contained almost no trace elements or the vitamins
that are provided in the Provasoli medium that may
in£uence the development of the microscopic phase
of the L. trabeculata life cycle. Culture media are en-
riched with iodine in the culture of Laminariales in
Japan as it appears to be an essential element in frond
elongation in these algae (Kawashima 1984; Lobban
Harrison1994). Based on the results of the present
study showing lack of statistical di¡erences, and
because of the relatively high cost of large-scale
production of Provasoli medium and the small e¡ect
of iodine addition on the growth of L. trabeculata, it
was concluded that agricultural nutrients provided
the most viable alternative for our tank cultures.
Table 3 Comparative increases (mean and standard deviation) in three growth characteristics of the abalone Haliotis discus
hannai cultured for 2 months using Lessonia trabeculata from three di¡erent origins
Increase per month (%)
Treatment Length Width Weight
L. trabeculata natural stand 6.4970.11 5.7070.23 24.0176.69
L. trabeculata long line 7.8070.77 6.8270.29 27.8470.26
L. trabeculata outdoor tank 6.9070.25 5.9170.43 26.5573.69
Initial sizes of abalone were L 51571mm,W 51071mm,Wt fresh 5 419784 mg (n 5180).
ANOVA analysis showed no signi¢cant di¡erences between treatments at P40.05.
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7. Once the sporophytes reached a size visible to the
naked eye under controlled conditions, it was recom-
mended to transfer them into intermediate culture in
outdoor tanks. During this stage, the shading of the
tanks, as described in Materials and methods, suc-
cessfully decreased the proliferation of microalgae
and epiphytes that otherwise grew on the cultured
algae and reduced their growth. Also, disintegration
of tissue in the fronds decreases with decreasing am-
bient light. It must be considered that L. trabeculata is
a subtidal alga, ranging from 1 to 20 m deep (Edding,
Fonck Machiavello1994), and is adapted to low light
occurring at1-m depth inthe culture tanks. For exam-
ple, during November (spring) in the middle day, the
light intensity can average up to 1400 mmol photons
m^2
s^1
in air; it is reduced to 35% at 1 m,65% at 3 m
and to 90% at 7 m depth (Edding et al.1990).
The ¢rst experiments in culturing at sea were ad-
versely a¡ected by biofouling particularly when cul-
tures were initiated in the spring at a time of heavy
phytoplankton blooms and at the peak reproductive
period of many fouling species. Greater coverage of
sporophytes with fouling organisms when they are
placed at sea at an early stage reduces light absor-
bance of the fronds, and thus lowers their photo-
synthesis and growth. Moreover, repeated cleaning
of fouling organisms from growing plants may re-
move apical tissue, and thus reduce the growth in
length of the fronds.
The values for the elongation of fronds and their
seasonal trends during culture at sea (Fig. 3) were
Figure 3 Elongation rate (mm day^1
) of Lessonia trabe-
culata fronds cultured on long-line systems during1988
and1999.
Figure 4 Long line-cultivated Lessonia trabeculata, E10 months from initiation of culture on a long-line system at1-
m depth, as described by Kawashima (1984).
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8. similar to those measured previously for the same
species in culture (Edding et al.1990) and in natural
beds (Tala 1999). The main di¡erences in the magni-
tude of growth and loss of frond tissue occurred at
the beginning as a response to changed conditions
of the culture from the tanks to the sea.
Laminariales may be harvested whole or by prun-
ing; the fronds are the most useful part of the plant
for feeding abalone (Owen et al.1984; Corazani Ill-
anes 1998; Castillo 2000). Observations in the ¢eld
show that the typical morphology of the plants tends
to change with greater productionof the holdfast and
frond and lower production of stipe. Good develop-
ment of the holdfast permits more secure ¢xation to
the substrate, and the fronds contribute principally to
photosynthetic activity and reproduction. In culture,
as haptera develop, it is recommended that the hold-
fasts be tied to the cords to ensure theirattachment in
order to reduce the loss of plants as they increase in
weight.
Abalone cultures require large amounts of algae
as feed, particularly in advanced growth stages
(Maureira et al. 1993). Present results suggest that
L. trabeculata can be cultured throughout the year.
The reproductive characteristics of L. trabeculata
(Tala 1994) suggest there should be no problem in
¢nding reproductive material during any season of
the year in natural beds. Out-of-phase cultures could
thus be initiated throughout the year. However, the
quality of spores and the production of sporophytes
may vary over time, and it is therefore suggested that
autumnal reproductive tissue be preferred as this is
the time of the year when the greatest number of
spores are produced bysporophytes (Tala1994). In this
way, allowing for the time required for the develop-
ment of the microscopic phase in the laboratory, plant-
lets could be placed in tank culture during the spring
and summer months, followed by placement at sea
late in the summer or early the following autumn.
Results of the present experiments demonstrate
the feasibility of large-scale culture of L. trabeculata
(Fig. 4). The main problems include the detachment
of small plants from cords in the early phases of cul-
ture and biofouling of the fronds both in the labora-
tory and at sea. In terms of potential yield, it is
possible that each 50-m long line, supporting algae-
seeded cords at 1-m intervals, could produce 500 kg
fresh weight of material from the same cohort of
spores. Aculture of 500 000 juvenile abalone (length
20 mm) would require about 2 tons of algae in their
¢rst year of culture, equivalent to four successfully
seeded and cultured Lessonia long lines. Future
research needs to be directed at improvement of the
algal biomass yields from long lines, with cultures
complementing the harvesting of this species from
natural beds. The costs involved in these parallel
activities greatly in£uence the ¢nal pro¢tability of
invertebrate cultures. Additionally, harvesting of
Lessonia from natural beds should be subject to
resource management regulations in order to main-
tain them over time and protect natural commu-
nities associated with these beds. In view of the
general reproductive characteristics of the Laminar-
iales, it is possible that other species along the Chi-
lean coast, such as Lessonia nigrescens Bory and
Macrocystis spp., could be brought into culture using
the presently described methods.
Acknowledgments
The authors are grateful for support given to this pro-
ject by the Chilean National Commission of Scienti¢c
and Technological Research (CONICYT) as Project
FONDEF no.1102.
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