16 THE ROLES OF BACTERIA AND MICRO AND MACRO ALGAE IN ABALONE AQUACULTURE: A REVIEW

THE ROLES OF BACTERIA AND MICRO AND MACRO ALGAE IN
ABALONE AQUACULTURE: A REVIEW
Author(s): SABINE DAUME
Source: Journal of Shellfish Research, 25(1):151-157.
Published By: National Shellfisheries Association
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Journal of Shellfish Research, Vol. 25, No. 1, 151–157, 2006.

THE ROLES OF BACTERIA AND MICRO AND MACRO ALGAE IN ABALONE
AQUACULTURE: A REVIEW
SABINE DAUME
Research Division, Department of Fisheries Western Australia, PO Box 20, North Beach,
WA 6920, Australia
ABSTRACT Abalone aquaculture is dependent on cultured algae to induce larval settlement and as a food source for the early life
stages of abalone until formulated feed or macroalgae such as Macrocystis sp., Porphyra sp. and Ulva sp. are introduced into the
growout system. In the natural environment, abalone larvae settle on coralline red algae, which provide one of the strongest and most
consistent settlement cues available for abalone larvae. However, propagation of coralline red algae is not practical commercially.
Abalone farms in Japan successfully settle abalone larvae (Haliotis discus hannai) on the green alga Ulvella lens. U. lens also proved

to be suitable to enhance settlement of cultured southern Australian abalone species (Haliotis laevigata, H. rubra). Most abalone farms
in Australia are now growing U. lens for that purpose. U. lens is easy to culture, no specific facilities are needed and the alga can be
grown on PVC settlement plates in commercial nursery tanks. However, U. lens has limited value as a feed for young postlarvae.
Instead, cultured diatoms can be added after larvae successfully settle and start feeding. Juvenile abalone (>3 mm in shell length) can
consume U. lens and grow rapidly on this alga. Diatom cultures and biofilms developing on settlement plates are not axenic and the
role of bacteria in early postlarvae feeding is poorly understood. It has been suggested that bacteria may perform metabolic activities
in the undeveloped gut of young postlarvae. At later stages of the nursery phase it becomes increasingly difficult to maintain adequate
feed on the plates and this is still regarded as a significant bottleneck for the abalone aquaculture industry. Recent investigations have
indicated that sporelings of macroalgae like Ulva sp. or diatoms that can provide more biomass may provide a suitable additional food
source for juveniles (>3 mm in shell length).
KEY WORDS:
lens

abalone, abalone eggs, antibiotics, algae, bacteria, diatoms, growth, larval quality, lipids, settlement, Ulva sp., Ulvella

INTRODUCTION

Abalone fisheries (Haliotis spp.) produce high value, exportorientated products with about 50% of the world supply being
provided by Australian fisheries in 1999 (Gordon & Cook 2001).
The worldwide catch from abalone fisheries has declined by about
30% over a 10-year period from ca. 14,000 mt in 1989 to 10,000
mt in 1999, and consequently the interest in aquaculture products
has increased substantially. The world production of abalone from
aquaculture in 1999 was approximately 7,775 tonnes (Gordon &
Cook 2001). Future production from the numerous farms and sites
established, under construction or approved in several countries
including Australia, could be even more substantial if the technology is improved.
In an aquaculture environment, abalone larvae are produced by
spawning recently collected wild broodstock, or wild or farmed
abalone broodstock that have been held in conditioning systems

for extended periods. The nonfeeding larvae have a short larval
phase (e.g., 7 days at 17°C for Haliotis rubra Leach and Haliotis
laevigata Donovan). When larvae are ready for settlement they
actively seek a suitable surface. In the natural environment, abalone larvae settle on coralline red algae (Shepherd & Daume
1996); however on farms the surface is typically vertical, spaced
plastic plates that have been colonized by a variety of different
algal species. Abalone aquaculture in most countries is dependent
on cultured algae at least for the early life stages, to induce larval
settlement and as a food source for postlarvae and juveniles, until
formulated food is introduced into the growout system. As provision of algal supplies decline, the juveniles may be weaned onto
formulated foods. They can be transferred to various land-based
tanks or sea-based systems (Freeman 2001). In several countries
around the world (e.g., South Africa) even the growout depends

Corresponding author. E-mail:

solely on algae; macroalgae that are harvested from the ocean are
fed to the abalone in specialist growout systems. A large component of the cost of producing juveniles is the provisioning of live
food in a manner suitable for a grazing herbivore. This review
examines the roles of bacteria, micro and macroalgae during the
nursery phase of abalone aquaculture and emphasizes research
conducted by the author with postlarval and juvenile H. laevigata
and H. rubra in Australia. It complements earlier reviews by Roberts (2001) on larval settlement and by Kawamura et al. (1998c) on
postlarval growth and survival by highlighting the applicability of
bacteria and algae for commercial abalone hatcheries and nurseries. Their roles are considered in the context of the main areas of
research undertaken to improve juvenile production efficiency: (1)
presettlement larvae quality; (2) larval settlement; (3) dietary requirements for postlarvae and juveniles.
Pre-settlement Larvae Quality

Previously wild abalone broodstock that feed on a range of

macroalgae have been the main source of gametes for commercial
abalone hatcheries. Selection of broodstock is mainly based on
gonad size and appearance (Litaay & De Silva 2001), with abalone
judged to be ready for induced spawning and have mature eggs
based on the amount of swelling of the gonad. However, animal
selection based on these criteria shows variable results in spawning
success and produce offspring with large variability in larval and
postlarval survival. More recently there has been greater commercial and research interest in conditioning captive and farmed
broodstock using macroalgae or formulated foods (Grubert & Ritar
2003, Daume & Ryan 2004a, Freeman et al. this volume).
Lipids and protein in abalone eggs are known to fuel the development and metamorphosis of the larvae (Jaeckle & Manahan
1989a, 1989b, Litaay et al. 2001). Nelson et al. (2002) demonstrated that variations in lipid content and fatty acid profile of the
digestive gland coincided with variation in their macroalgal diets

151


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152

and are related to seasonal temperature fluctuations. Biochemical
variation in the diet may affect the composition of the eggs and
ultimately larval performance. However studies of changes in biochemical composition such as fatty acids in abalone eggs are
scarce. Litaay et al. (2001) demonstrated changes in biochemical
composition during larval development. Recently, Daume and
Ryan (2004a) showed high variability in proximate biochemical
composition and fatty acid profiles of abalone eggs between
batches derived from conditioned and wild broodstock as well as
between two consecutive spawning seasons. The relative proportions of some PUFAs in the broodstock diets were reflected in the

eggs and varied between batches of conditioned and wild broodstock, indicating that formulated diets designed to maximize
growth rates are not necessarily adequate to maintain viable, high
quality eggs and larvae from captive broodstock.
Other factors that can influence the quality and success of
larval culture are opportunistic pathogenic bacteria that can bloom
and cause deformities in and collapse of whole larval batches
under potentially stressful commercial growing conditions. Many
abalone hatcheries are using antibiotics like oxytetracycline prophylactically. Similarly they may be used in research projects.
Roberts (2001) suggested using antibiotics to eliminate bacterial
interference in settlement assay systems. Apart from the general
problem of development of antibiotic resistant strains of bacteria in
hatcheries, problems have been reported with certain antibiotics
when used with abalone during larval rearing or settlement assays.
Streptomycin at low doses of 5 ␮g mL−1 was toxic to Haliotis
diversicolor (Bryan & Qian 1998). Emitine caused abnormal loss
of velum that could have been confused with metamorphosis
(Fenteany & Morse 1993).
An experiment conducted to assess the effect of two antibiotics
(Ampicillin and Kanamycin at 50 ␮g mL−1) on the settlement of
H. rubra revealed no difference in settlement rate between treated
and untreated settlement substrate (Table 1). In this experiment 3
algal settlement substrata were tested (Navicula cf. jeffreyi, Ulvella
lens, Sporolithon durum) and compared with a negative control
(plastic square of commercial settlement plate without any algal
growth) all with and without antibiotics. The ratios of settlement
rates between treated and untreated substrates did not change over
time. In addition, the difference in settlement preferences between
specific substrates remained the same regardless if antibiotics were
used or not. The antibiotics were initially effective as indicated by
the higher survival of swimming larvae (in water column) in control jars treated with antibiotics. However, the settlement rate was

not higher in the antibiotic treatment, indicating that unfit larvae
might survive if treated with antibiotics but they do not settle
successfully. This result questions the need and usefulness of antibiotics in abalone hatcheries. Further studies are needed to assess
the effects of other antibiotics and earlier treatment with antibiotics
(e.g., during larval rearing). However, alternatives like probiotics
should be investigated to enhance larval survival safely.
Many antibiotics, including Kanamycin and oxytetracycline,
work by inhibiting or interfering with the protein biosynthesis by
targeting the bacterial ribosomes. The close similarity between
bacterial and mitochondrial ribosomes makes the latter (present in
all cells of the “treated” organisms) a potential target (Hart 2004).
Inhibition of mitochondrial protein synthesis or injuries in mitochondria of the treated organism have occurred and can lead to
various dysfunction; any cell type or tissue with a high aerobic
energy requirement is more likely to be affected when this organelle is injured (Hart 2004). The effects of antibiotics on abalone
larval settlement and postlarval performance however are not well
understood. The knowledge we have from other systems, however,
warrants extreme caution and highlights the danger of introducing
other, potentially detrimental factors. These may not be obvious
initially but may manifest themselves at later stages of larval or
postlarval development.
Larval Settlement

The term “settlement” in this review describes the permanent
attachment of abalone larvae to the substrate after shedding of the
velum to complete metamorphosis. In the natural environment,
abalone larvae, like many other invertebrate larvae, settle on coralline red algae. Daume et al. (1999a) revealed that settlement of
Haliotis laevigata larvae in response to three nongeniculate coralline red algae is species-specific. In that study the frequency of
occurrence of epiphytic bacteria and diatoms was assessed on all
coralline red algal species tested. However, no significant correlation was found indicating that the settlement induction is algal in

origin. The authors concluded that bacteria and diatoms may influence the settlement response of abalone larvae but they are not
the main driving force. Roberts (2001) referred to some of his
unpublished work and stated that bacteria can induce abalone larval settlement but that the response is slow, taking 1 week to reach
50% metamorphosis. In contrast, very rapid settlement was reported in small-scale laboratory experiments through the use of the
coralline red alga, Sporolithon durum, with the maximum rate

TABLE 1.
Percentage settlement of Haliotis rubra on different settlement substrates (Ulvella lens and Navicula cf. jeffreyi and a negative control), with
and without antibiotics, as well as Sporolithon durum (positive control) after 24, 48 hours, % settled and survived up to 1 week and % of
larvae in water column after 1 week (n = 6 ± SE). Data are from Daume (2003).

Species

Antibiotics

% Settlement
24 Hours

% Settlement
48 Hours

% Survival Up
to 1 Week

% in Water Column
After 1 Week

Ulvella lens
Ulvella lens
Navicula cf. jeffreyi

Navicula cf. jeffreyi
Control
Control
Sporolithon durum

−
+
−
+
−
+
−

30 ± 8.1a
22 ± 4.4a
5 ± 1.4b
0.3 ± 0.3b
0 ± 0b
0 ± 0b
39 ± 3.7

35 ± 7.6a
36 ± 5.3a
3 ± 2.0b
1 ± 0.3b
1 ± 0.6b
0.3 ± 0.3b
50 ± 4.6

12 ± 1.5

17 ± 1.1
4 ± 1.2
2 ± 0.3
0.5 ± 0.3
0.6 ± 0.3
16 ± 2.6

0±0
5 ± 1.6
8 ± 3.5
30 ± 4.2
3 ± 1.1
54 ± 6.7
0±0

* Means with different superscript letters are significantly different (P < 0.05).


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being reached after 24 h (Daume et al. 1999a) indicating that
nongeniculate coralline red algae are strong settlement inducers.

This result coincides with disproportional high numbers of recruits
found on S. durum in the natural environment (Shepherd & Daume
1996).
Historically, benthic biofilms, consisting of bacteria and mixed
diatom species growing on PVC settlement plates, have been used
in abalone hatcheries worldwide to induce larval settlement. Diatoms, brought in by the incoming seawater, colonize clear plastic
sheets arranged in commercial nursery tanks. This process is unpredictable and larval settlement rates can be low (1% to 10% of
larvae) (Daume 2003). In both experimental and commercial systems, to achieve more control and consistency, films dominated by
single algal species can be generated (Daume et al. 2000, Daume
& Ryan 2004b). H. rubra did not respond to films of any diatom
species tested, but settled on the nongeniculate coralline red alga
Phymatolithon repandum (Daume et al. 1999b). In contrast, H.
laevigata settled comparably well on the diatom Navicula ramosissima and on the coralline S. durum. Roberts (2001) reviewed
data on settlement cues including diatoms and other biofilms.
Overall it is apparent that coralline red algae provide more consistent and reliable settlement cues, whereas settlement on diatoms
can be highly variable. However, propagation of coralline red algae is not practical at a commercial scale.
Abalone hatcheries in Japan successfully settle abalone larvae
(Haliotis discus hannai) on the green alga Ulvella lens (Takahashi
& Koganezawa 1988). U. lens is also suitable for enhancing settlement of both cultured southern Australian abalone species (H.
rubra and H. laevigata) (Fig. 1). Most abalone farms in Australia
are now growing U. lens for that purpose (Daume et al. 2000,
Daume et al. 2004, Daume & Ryan 2004b). The earlier study
established settlement preferences of H. rubra for U. lens at laboratory scale whereas the later studies focused on commercial scale
experiments. Both species (H. rubra, H. laevigata) showed a clear
preference for older rather than for younger U. lens (Table 2, Table
3) even with similar percentage cover, indicating that the developmental stage of the alga and not percentage cover per se is
important in settlement induction (Table 3). Settlement was also
significantly higher in the combined U. lens treatments (old and
young) compared with 2 diatom treatments (Navicula cf. jeffreyi
and Cocconeis sp. demonstrating the suitability of U. lens to improve the settlement of Haliotis laevigata larvae on commercial

scale (Table 3). No significant difference between high and low
larval release densities was found with H. rubra in the nursery
(Table 2) confirming earlier findings at laboratory scale with H.
laevigata larvae that settlement of abalone larvae is not gregarious
when tested with larvae of the same batch (Daume et al. 1999a). In
contrast, settlement was found to be gregarious in response to
conspecific postlarvae as young as 7 days (Daume et al. 1999a)
and older conspecific juveniles and adults and their grazing mucus
is believed to be responsible (Seki & Kan-no 1981, Slattery 1992).

Figure 1. Sequence from settlement cue to potential food items proposed for Australian temperate abalone species, in commercial farming systems, as they grow.

IN

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TABLE 2.
Percentage settlement (±SE) of Haliotis rubra in the nursery 3 days
after larval release (n = 32). Data from Daume et al. (2004).
Larval
Density
High
Low

Ulvella lens

Per U. lens
Treatment

Total
per Tank

Old (18 days 31% cover).
Young (4 days 57% cover).
Old (18 days 31% cover).
Young (4 days 57% cover).

31.9 ± 7.5
21.7 ± 6.8
44.0 ± 7.3
26.4 ± 7.6

53.6 ± 5.8

Average

70.4 ± 8.7
62.0

Recently alternative systems, to replace live algae as a means of
settlement and growing postlarvae, have been proposed in Japan
for H. discus discus and H. diversicolor (Stott et al. 2002, 2003,
2004a, 2004b). In the earlier studies, an alginate gel solution containing micro particulate diets was pasted onto settlement plates. In
more recent studies settlement plates are sprayed with a solution of
agar and one of the following: dried algal powder (Spirulina platensis, Chlorella vulgaris, Undaria pinnafifida), dried natural diatom powder, formulated diet and two different concentrations of
␥-aminobutyric acid (GABA), each with and without antibiotics,
and compared with negative (clean plates) and positive (living
natural diatom biofilms). In both recent studies there was no significant difference in settlement rates between the microalgae

powder treatments and the living natural biofim but both supported
significantly higher rates when compared with the negative control
and GABA treatments (Stott et al. 2004a, 2004b). The authors
demonstrated that pregrazing of plates by conspecific juveniles
covered with microalgal powder/ agar solution enhanced larval
settlement significantly (85% vs. 30% on grazed and ungrazed
plates respectively). This system shows some potential, however
mechanized and cost-efficient ways of spraying the plates need to
be developed before it becomes viable commercially.
Dietary Requirements

Post-larval abalone feed on benthic diatoms (Kawamura et al.
1995) and the diatom film on plates also provides the food for
growing postlarvae in commercial abalone nurseries. Commercial
farms traditionally rely on mixed species of diatoms as a food
source throughout the nursery period (settled larvae to 8–10 mm).
The film is maintained through passive seeding (new cells are
brought in with the incoming seawater), adding nutrients and manipulating the light intensity through shading. Without much control over composition and density of the biofilm species, the results
are very inconsistent and often very poor. Isolating particular diatom species and growing them in monoculture before inoculating
settlement tanks in the nursery affords greater control. This however has not been embraced by the industry and further investigations are needed to assess the effectiveness in larger scale systems.
However, a significant bottleneck experienced by industry is the
inability to maintain adequate food (both quantity and quality) on
the plates particularly at later stages of the nursery phase. Growth
rates of juveniles are influenced by the availability, digestibility
and nutritional composition of the algae (Kawamura et al. 1998b,
Roberts et al. 1999, Daume et al. 2003).
The Role of Bacteria in Postlarval Nutrition

Diatom cultures and biofilms developing on settlement plates
are not axenic and the role of bacteria in early postlarvae feeding

DAUME

154

TABLE 3.
Percentage settlement (±SE) of Haliotis laevigata 3 days after larval release (n = 3) when given a choice between 4 substrates. Data from
Daume and Ryan (2004b).

Treatments

Old U. lens
(8 weeks–97% cover)

Young U. lens
(6 weeks–82% cover)

Navicula sp.

Cocconeis sp.

Total
per Tank

% Settlement

61 ± 14

14 ± 1

7 ± 0.3

5 ± 0.5

87

and growth is poorly understood. Newly settled postlarvae ingest
diatoms but are often not able to digest the cell contents. This
suggests that bacteria and the extracellular material produced by
the diatoms, present in the biofilm, are a significant source of
nutrition for postlarval abalone (Fig. 1). Garland et al. (1985)
reported that postlarval H. rubra ingested bacteria growing on the
surface of coralline red algae. It has been suggested that bacteria
may perform metabolic activities in the undeveloped gut of young
postlarvae and are able to enhance the digestion efficiency of the
host by supplying polysaccharolytic enzymes (Garland et al. 1985,
Erasmus et al. 1997). Polysaccharolytic enzyme activity has been
reported in day 17 H. discus hannai postlarvae (Takami et al.
1998). Sawabe et al. (2003) detected the bacteria Vibrio halioticoli
in the gut of H. diversicolor aquatilis and suggested that this
bacterium may play a crucial role in converting alginate to acetic
acid. As part of the alternative systems proposed by Stott et al.
(2002, 2003, 2004a, 2004b), the authors observed that the growth
of postlarvae H. diversicolor aquatilis fed a formulated diet was
reduced when antibiotics were added and suggested that bacteria
that assisted in digestion became limiting. In a later study they
discovered that 5–10 times more bacteria (including Vibrio spp.)
were present on plates sprayed with the agar/formulated diet solution. These bacteria could have provided a substantial food
source to early postlarvae, which may have contributed to the

significantly better growth rates on these plates 1 week after settlement (Stott et al. 2004b). The authors suggest that for recently
settled postlarvae, bacteria might be a superior food source compared with diatom and abalone grazing mucus. All these studies
indicate that bacteria are ingested and play an important role in
early postlarvae nutrition and health, but further studies are needed
to elucidate their role and contribution.

grew faster on Cocconeis scutellum and Cylindrotheca closterium.
Both species were most efficiently digested. Transitions in postlarval feeding preferences and growth performances on different
algal species are reviewed in Kawamura et al. (1998c).
Alternative Food Sources for all Stages of Nursery Culture

Food Preferences for Postlarval Abalone

The green alga U. lens has limited value as a food for growing
postlarvae. Instead, cultured diatoms can be added after larvae
successfully settle and start feeding. Seki (1997) reported that
growth rates of postlarvae on U. lens were improved by the inoculation of cultured diatoms.
Recent studies showed that plates with a low cover of young
germlings of U. lens could be used for settlement induction of
Australian abalone species (H. rubra, H. laevigata) and followed
with inoculation of the cultured diatom Navicula cf. jeffreyi to
ensure sufficient food for the growing postlarvae (Daume et al.
2000, 2004, Daume & Ryan 2004b). The former study provided
crucial information on early development of H. rubra and established that growth rates on several diatom species are significantly
higher than on U. lens at laboratory scale (Fig. 2). In the more
recent study, at commercial scale, the type of substrate on which
larvae settled, light (which affected the food density) and the density of postlarvae all had very marked effects on growth (Daume et
al. 2004). The results also suggest that early growth is important in
determining later performance. Daume and Ryan (2004b) investigated settlement, growth, survival and size variability of the abalone H. laevigata on commercial scale. Both growth rate and size
variability increased over time until juveniles reached approximately 5 mm in shell length. Whereas postlarval abalone do not

grow well on U. lens (Fig. 2), juvenile abalone (>3 mm in shell
length) can consume U. lens and grow rapidly (80–110 ␮m day–1)
on this alga (Table 4).

Worldwide, several studies have examined postlarval feeding
and growth on different algal species (Ohgai et al. 1991, Ishida et
al. 1995, Kawamura et al. 1998a, Roberts et al. 1999). Studies
devoted to examining their feeding preferences and growth (Kawamura & Kikuchi 1992, Kawamura & Takami 1995, Kawamura et
al. 1995, Matthews & Cook 1995, Kawamura 1996, Takami et al.
1997, Daume et al. 2000, Takami & Kawamura 2003) have shown
that food requirements change as abalone grow (Fig. 1). Two to
three weeks after settlement, postlarvae become responsive to the
“digestibility” of the diatom strains and grow more rapidly on
effectively digested strains (Kawamura et al. 1998a, 1998b). Postlarvae 0.8–2 mm in shell length grow ca 40–60 ␮m day−1 on
“digestible” diatoms and only ca 15–30 ␮m day−1 on “indigestible” diatoms (Kawamura et al. 1998b). In addition, the diatom cell
size, attachment strength, frustule’s strength and postlarval size
can influence digestion. In a feeding trial covering the whole postlarval period, Roberts et al. (1999) showed that different diatom
food species affected survival and growth. After day 17, postlarvae

Figure 2. Early growth of H. rubra postlarvae feeding on different
algal species. Vertical bars indicate standard error; n = 4. Data from
Daume et al. (2000).


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155

higher nitrate medium. Searcy-Bernal et al. (2003) found that recently settled H. fulgens postlarvae grew and survived better under
Daily growth-rates (µm day ) of juveniles (Haliotis rubra) on plates
low light (6 ␮E) conditions, whereas a lower number of cells of the
52 days after settlement and shell length (mm) 114 days after
diatom Navicula incerta were available in the lower light treatsettlement (mean ± SE). Data from Daume et al. (2004).
ment. The authors suggested that oxygen supersaturation in the
boundary layer, particularly in high-density diatom films at high
−1
Daily Growth Rate (µm day )
Shell Length (mm) light levels (75 ␮E), could have caused high mortality in this
114 Days
U. lens 52–64 Days 64–94 Days 94–114 Days
treatment. In another study, the influence of light intensity on two
diatom species (Navicula cf. jeffreyi, Cocconeis sp.) as a food for
Old
79.4 ± 7.7 107.4 ± 4.2
82.8 ± 4.2
6.9 ± 0.2
juvenile H. laevigata (3–4 mm in shell length) was tested (Watson
Young

94.9 ± 8.4 115.3 ± 14.8
87.8 ± 8.2
7.4 ± 0.2
et al. 2004). In contrast to N. cf. jeffreyi, growth of Cocconeis sp.
was not inhibited at lower light levels making it a good candidate
At later stages of the nursery phase (>5 mm in shell length), it for culture in shaded nursery systems. Light was more influential
becomes increasingly difficult to maintain adequate food on the in juvenile grazing behavior (photophobic) than food availability.
plates and this is still regarded as a significant bottleneck for the Watson et al. (2005) examined the combined effect of manipulaindustry. Recent investigations have indicated that sporelings of tions in light intensity and nitrate concentrations on the nutritional
macroalgae like Ulva sp. may provide a suitable food source for value of the diatom Navicula cf. jeffreyi when fed to juvenile
juveniles (see Strain et al. this volume) (Fig. 1). Alternatively, abalone (H. laevigata). Under high light conditions Navicula cf.
chain forming diatoms, like Delphineis, offer a 3-D structure com- jeffreyi was lower in protein and higher in carbohydrates and fat.
pared with the 2-D structure of nonchain forming prostrate attach- Juveniles grazed larger numbers of diatom cells when the protein
ing species, like Navicula spp. and thus providing more biomass content was low, possibly compensating for the lower protein levfor the growing juveniles (Fig. 1). Kawamura et al. (1995) reported els. The authors reported elevated pH levels in higher light treatgrowth rates of 48 ␮m day−1 of H. discus hannai juveniles 1–2 mm ments and suggested that this could have caused high mortality.
in shell length, when feeding on the diatom Achnanthes longipes, These studies indicate that changes in light intensity and nitrate
which has a 3-D structure. More recently, Takami and Kawamura. concentration, under which the diatom species are cultured, can
(2003) found that juveniles 2.8–2.9 mm in shell length grew 100 have a dramatic effect on growth, grazing rates and particularly
␮m day−1 on this diatom species, which was comparable to growth survival of postlarval and juvenile abalone. This emphasizes the
rates achieved on juvenile sporophytes of the macroalga Lami- need for selecting the right light and nutrient level to achieve high
value food and conditions for optimal growth and survival of junaria japonica.
venile abalone in commercial nurseries.
Biochemical Composition and Nutritional Value of Algal Diets
This study reviewed three main areas of abalone research associated with abalone hatchery and nursery production. Further
The biochemical composition of microalgae, and therefore their studies are needed to find alternatives, such as probiotics, to the
nutritional value to herbivores varies between species (Brown et al. use of antibiotics in abalone hatcheries. Alternative cost effective
1996) and is greatly affected by harvest stage, light intensity foods, for broodstock and for the latter stage of the nursery still
(Thompson et al. 1993, Brown et al. 1996), nutrient concentrations need to be found that will increase larval quality and allow abalone
(Fábregas et al. 1996, Fábregas et al. 1998) and culture methods farmers to keep animals on the plates longer and thus reduce
(Otero & Fábregas 1997). It is known that the biochemical com- weaning mortality.
position of algae can be altered by changing the growing condiACKNOWLEDGMENTS
tions (e.g., Otero & Fábregas 1997, Thompson et al. 1993, Brown

et al. 1996). When microalgal cultures are grown in nitrogenThe author thanks Stephen Ryan, Sylvain Huchette, Ben Long,
limited media, the protein content of the cells decreases (Enright et Peter Crouch, Anton Krisnich, Sascha Brand-Gardner, Rob Day
al. 1986, D’Souza & Kelly 2000, Daume et al. 2003). Daume et al. and Bill Woelkerling who were involved in various parts of the
(2003) showed previously that juvenile H. rubra grew faster when work on H. laevigata and H. rubra. and Greg Maguire for many
feeding on the diatom Navicula cf. jeffreyi that was cultured in a useful comments.
TABLE 4.

−1

LITERATURE CITED
Brown, M. R., G. A. Dunstan, S. J. Norwood & K. A. Miller. 1996. Effects
of harvest stage and light on the biochemical composition of the diatom
Thalassiosira pseudonana. J. Phycol. 32:64–73.
Bryan, P. J. & P. Qian. 1998. Induction of larval attachment and metamorphosis in the abalone Haliotis diversicolor (Reeve). J. Exp. Mar.
Biol. Ecol. 223:39–51.
D’Souza, F. M. L., & G. J. Kelly. 2000. Effects of a diet of a nitrogenlimited alga (Tetraselmis suecica) on growth, survival and biochemical
composition of tiger prawn (Penaeus semisulcatus) larvae. Aquaculture
191:311–329.

Daume, S., S. Brand-Gardner & Wm. J. Woelkerling. 1999a. Settlement of
abalone larvae (Haliotis laevigata Donovan) in response to nongeniculate coralline red algae (Corallinales, Rhodophyta). J. Exp. Mar.
Biol. Ecol. 234:125–143.
Daume, S., S. Brand-Gardner & Wm. J. Woelkerling. 1999b. Preferential
settlement of abalone larvae: diatom films versus non-geniculate coralline red algae. Aquaculture 174:243–254.
Daume, S., A. Krsinich, S. Farrell & M. Gervis. 2000. Settlement, early
growth and survival of Haliotis rubra in response to different algal
species. J. Appl. Phycol. 12:479–488.

Daume, S. 2003. Early life history of abalone (Haliotis rubra, H. laevigata): settlement, survival and early growth. Final report for FRDC
project 1998/306. Department of Fisheries. Western Australia. Fisheries Research Contract Reports 3:1–110.

Daume, S., B. M. Long & P. Crouch. 2003. Changes in amino acid content
of an algal feed species (Navicula sp.) and their effect on growth and
survival of juvenile abalone (Haliotis rubra). J. Appl. Phycol. 15:201–
207.


156

DAUME

Daume, S., S. Huchette, S. Ryan & R. W. Day. 2004. Nursery culture of
Haliotis rubra: The effect of cultured algae and larval density on settlement and juvenile production. Aquaculture 236:221–239.
Daume, S. & S. Ryan. 2004a. Fatty acid composition of eggs derived from
conditioned and wild caught greenlip abalone broodstock (Haliotis laevigata). J. Shellfish Res. 23(4):967–974.
Daume, S. & S. Ryan. 2004b. Nursery culture of the abalone Haliotis
laevigata: Larval settlement and juvenile production using cultured
algae or formulated feed. J. Shellfish Res. 23(4):1019–1026.
Enright, C. T., G. F. Newkirk, J. S. Craigie & J. D. Castell. 1986. Growth
of juvenile Osterea edulis L. fed Chaetoceros gracilis Schuett of varied
chemical composition. J. Exp. Mar. Biol. Ecol. 96:15–26.
Erasmus, J. H., P. A. Cook & V. E. Coyne. 1997. The role of bacteria in
the digestion of seaweed by the abalone Haliotis midae. Aquaculture
155:377–386.
Fábregas, J., A. Otero, E. Morales, B. Cordero & M. Patino. 1996. Tetraselmis suecica cultured in different nutrient concentrations varies in
nutritional value to Artemia. Aquaculture 143:197–204.
Fábregas, J., A. Otero, E. Morales, B. O. Arredondo-Vega & M. Patino.
1998. Modification of the nutritive value of Phaeodactylum tricornutum for Artemia sp. in semicontinuous culture. Aquaculture 169:167–
176.
Fenteany, G. & D. E. Morse. 1993. Specific inhibitors of protein synthesis

do not block RNA synthesis or settlement in larvae of a marine gastropod mollusk (Haliotis rufescens). Biol. Bull. 184:6–14.
Freeman, K. A. 2001. Aquaculture and related biological attributes of
abalone species in Australia—a review. Fisheries WA. Fisheries research report. pp. 128.
Freeman, K., S. Daume, M. Rowe, S. Parsons, R. Lambert & G. B. Maguire. Effects of season, temperature control, broodstock conditioning
period and handling on incidence of controlled and uncontrolled
spawning of greenlip abalone (Haliotis laevigata Donovan) in Western
Australia. J. Shellfish Res. (this volume).
Grubert, M. A. & A. J. Ritar. 2003. The effect of temperature and conditioning interval on the spawning success of wild-caught blacklip (Haliotis rubra) and greenlip (H. laevigata) abalone fed an artificial diet. In:
A. E. Fleming, editor. Proceedings of the 10th Annual Abalone Aquaculture Workshop, November, 2003, Port Lincoln, Australia. Fisheries
Research and Development Corporation, Abalone Aquaculture Subprogram, Canberra, Australia. pp. 5–25.
Garland, C. D., S. L. Cooke, J. F. Grant & T. A. McMeekin. 1985. Ingestion of the bacteria on and the cuticle of crustose (non-articulated)
coralline algae by post-larval and juvenile abalone (Haliotis ruber
Leach) from Tasmanian waters. J. Exp. Mar. Biol. Ecol. 91:137–149.
Gordon, H. R. & P. A. Cook. 2001. World abalone supply, markets and
pricing: historical, current and future. J. Shellfish Res. 20:567–570.
Hart, S. G. E. 2004. Mechanisms of mitochondrial toxicity. In: American
College of Veterinary Pathologists & American Society of Veterinary
Clinical Pathology. American College of Veterinary Pathologists &
American Society of Veterinary Clinical Pathology, editors. Proceedings of 55th Annual Meeting of the American College of Veterinary
Pathology and 39th Annual Meeting of the American Society of Veterinary Clinical Pathology, 2004. Middleton, USA. pp. 1–4.
Ishida, T., T. Akutsu & K. Torisawa. 1995. Effects of monocultured
benthic diatoms on the metamorphosis of veliger larvae and on the
growth of juveniles of abalone, Haliotis gigantean. Bull. Shizuoka Pref.
Fish. Exp. Stn. 30:17–21.
Jaeckle, W. B. & D. T. Manahan. 1989a. Feeding by a “nonfeeding” larva:
uptake of dissolved amino acids from seawater by lecithotrophic larvae
of the gastropod Haliotis rufescens. Mar. Biol. 103:87–94.
Jaeckle, W. B. & D. T. Manahan. 1989b. Growth and energy imbalance
during the development of a leicithotrophic molluscan larva. Biol. Bull.
177:237–246.

Kawamura, T. & H. Kikuchi. 1992. Effects of benthic diatoms on settlement and metamorphosis of abalone larvae. Suisanzoshoku 40:403–
409.
Kawamura, T., T. Saido, H. Takami & Y. Yamashita. 1995. Dietary value

of benthic diatoms for the growth of post-larval abalone Haliotis discus
hannai. J. Exp. Mar. Biol. Ecol. 194:189–199.
Kawamura, T. & H. Takami. 1995. Analysis of feeding and growth rate of
newly metamorphosed abalone Haliotis discus hannai fed on four species of benthic diatom. Fish. Sci. 61:357–358.
Kawamura, T. 1996. The role of benthic diatoms in the early life stages of
the Japanese abalone Haliotis discus hannai. In: Y. Watanabe, Y. Yamashita & Y .Oozeki, editors. Survival strategies in early life stages of
marine resources. Rotterdam: Balkema. pp. 355–367.
Kawamura, T., K. Okamura & H. Takami. 1998a. Growth of benthic
diatom Cocconeis scutellum var. parva, a suitable food for post-larval
abalone. Suisanzoshoku 46:509–516.
Kawamura, T., R. D. Roberts & C. M. Nicholson. 1998b. Factors affecting
the food value of diatom strains for post-larval abalone Haliotis iris.
Aquaculture 160:81–88.
Kawamura, T., R. D. Roberts, H. Takami & C. M. Nicholson. 1998c. A
review of the feeding and growth of postlarval abalone. J. Shellfish Res.
17:615–625.
Litaay, M. & S. S. De Silva. 2001. Reproductive performance indices
based on physical characteristics of female blacklip abalone Haliotis
rubra L. J. Shellfish Res. 20:673–677.
Litaay, M., S. S. De Silva & R. M. Gunasekera. 2001. Changes in the
amino acid profiles during embryonic development of the blacklip
abalone (Haliotis rubra). Aquat. Living Resour. 14:335–342.
Matthews, I. & P. A. Cook. 1995. Nutrition of newly settled larvae of the
South African abalone Haliotis midae. Mar. Freshwater Res. 46:545–
548.
Nelson, M. M., D. L. Leighton, C. F. Phleger & P. D. Nichols. 2002.

Comparison of growth and lipid composition in the green abalone,
Haliotis fulgens, provided specific macroalgal diets. Comp. Biochem.
Physiol. B 131:695–712.
Ohgai, M., M. Wakano & S. Nagai. 1991. Effect of attached microalgae on
the settlement of larvae and growth of juveniles in abalone, Haliotis
discus hannai Ino. Suisanzoshoku 39:263–266.
Otero, A. & J. Fábregas. 1997. Changes in nutrient composition of Tetraselmis suecica cultured semicontinously with different nutrient concentrations and renewal rates. Aquaculture 159:111–123.
Roberts, R. 2001. A review of settlement cues for larval abalone (Haliotis
spp.). J. Shellfish Res. 20:571–586.
Roberts, R. D., T. Kawamura & C. M. Nicholson. 1999. Growth and
survival of post-larval abalone (Haliotis iris) in relation to development
and diatom diet. J. Shellfish Res. 18:243–250.
Sawabe, T., N. Setoguchi, S. Inoue, R. Tanaka, M. Ootsubo, M. Yoshimizu
& Y. Ezura. 2003. Acetic acid production of Vibrio halioticoli from
alginate: a possible role for establishment of abalone-V. halioticoli
association. Aquaculture 219:671–679.
Searcy-Bernal, R., C. Anguiano-Beltran & A. Esparza-Hernandez. 2003.
The effect of irradiance on the survival and growth of abalone postlarvae Haliotis fulgens fed Navicula incerta. Aquaculture 228:237–
248.
Seki, T. 1997. Biological studies on the seed production of the northern
Japanese abalone. Bull. Tohoku Nat. Fish. Res. Instit. 59:1–71.
Seki, T. & H. Kan-no. 1981. Induced settlement of the Japanese abalone,
Haliotis discus hannai, veliger by the mucous trails of the juvenile and
adult abalones. Bull. Tohoku Reg. Fish. Res. Lab. 43:29–36.
Shepherd, S. A. & S. Daume. 1996. Ecology and survival of juvenile
abalone in a crustose coralline habitat in South Australia. In: Y. Watanabe, Y. Yamashita & Y. Oozeki, editors. Survival strategies in early
life stages of marine resources. Rotterdam: Balkema. pp. 297–313.
Slattery, M. 1992. Larval settlement and juvenile survival in the red abalone (Haliotis rufescens): an examination of inductive cues and substrate selection. Aquaculture 102:143–153.
Stott, A., T. Takeuchi, Y. Koike, H. Yamakawa & O. Imada. 2002. Using
micro particle diets to replace diatoms for feeding postlarval abalone

Haliotis discus discus (Reeve). Fish. Sci. 68:1088–1093.
Stott, A., T. Takeuchi, Y. Koike & O. Imada. 2003. Settling and raising


ROLES

OF

BACTERIA

AND

ALGAE

post-larval abalone Haliotis diversicolor supertexta (Lischke) on microparticulate diets embedded in a layer of alginate. Aquaculture Research 34:561–567.
Stott, A. E., T. Takeuchi & Y. Koike. 2004a. Testing various substances
that have been bound to plastic plates with agar to induce larval settlement and metamorphosis of abalone Haliotis discus discus (Reeve).
Aquaculture 231:547–557.
Stott, A. E., T. Takeuchi & Y. Koike. 2004b. An alternative culture system
for the hatchery production of abalone without using live food. Aquaculture 236:341–360.
Strain, L., M. Borowitzka & S. Daume. Growth and survival of juvenile
greenlip abalone (Haliotis laevigata) feeding on germlings of the macroalgae Ulva sp. In comparison with a current commercial diet consisting of Ulvella lens plus the diatom species Navicula cf. jeffreyi. J.
Shellfish Res. (this volume).
Takahashi, K. & A. Koganezawa. 1988. Mass culture of Ulvella lens as a
feed for abalone Haliotis discus hannai. NOAA Technical Report
NMFS 70:25–36.
Takami, H., T. Kawamura & Y. Yamashita. 1997. Contribution of diatoms

IN

ABALONE AQUACULTURE

157

as food sources for post-larval abalone Haliotis discus hannai on a
crustose coralline alga. Molluscan Res. 18:143–152.
Takami, H., T. Kawamurra & Y. Yamashita. 1998. Development of
polysaccharide degradation activity in postlarval abalone Haliotis discus hannai. J. Shellfish Res. 17:723–727.
Takami, H. & T. Kawamura. 2003. Dietary changes in the abalone Haliotis
discus hannai, and relationship with the development of the digestive
organ. Japan Agricultural Research Quarterly 37:89–98.
Thompson, P. A., M. Guo & P. J. Harrison. 1993. The influence of irradiance on the biochemical composition of three phytoplankton species
and their nutritional value for larvae of the Pacific Oyster (Crassostrea
gigas). Mar. Biol. 117:259–268.
Watson, D., S. Daume, J. Prince, L. Beasley & B. Knott. 2004. The influence of light intensity on the density of different diatoms as feed for
juvenile greenlip abalone (Haliotis laevigata). Aquaculture 235:345–
359.
Watson, D., S. Daume, J. Prince, L. Beasley & B. Knott. 2005. The influence of culture conditions on the growth and biochemical composition
of algal feed species for juvenile greenlip abalone (Haliotis laevigata).
Molluscan Research 25:1–8.