JOURNAL OF

EXPERIMENTAL
MARINE BIOLOGY
Journal of Experimental Marine Biology and Ecology AND ECOLOGY
ELSEVIER 194 (1995) 189-199

Dietary value of benthic diatoms for the growth of

post-larval abalone Haliotis discus hannai
Tomohiko Kawamuraa*, Takahiro Saidob, Hideki Takami,
Yoh Yamashita
Tohoku National Fisheries Research Institute, Shinhama-cho, 3-27-5 Shiogama, Miyagi 985, Japan
bIwate Fisheries Technology Center, Heita, Kamaishi, Iwate 026, Japan

Received 16 January 1995; revision received 17 May 1995; accepted 31 May 1995

Abstract

The feeding behaviour and the growth of post-larval abalone (l-2 mm in shell length)
Haliotis discus fed on nine species of benthic diatom were examined in the
hannai
laboratory. Digestion efficiencies of abalone fed on the nine diatom species were also
measured. All abalone showed active feeding behaviour and diatom cells were observed in
their stomachs. However, the growth rates of abalone fed on four diatom species, which
were readily digested, were higher than those of abalone fed on the other five species of
diatom. The cell walls of the former four diatom species were easily broken open when
they were grazed by abalone because of their high adhesive strength (three species), or due
to the weakly silicified cell walls (one species with a low adhesive strength). The other five
species, whose adhesive strengths are low, were easily ingested by abalone without any
deformation of the cell walls and the majority of them were excreted whole and still alive.
It is considered that post-larval abalone cannot digest diatom cell contents in their
alimentary canal without first rupturing the cell wall with the radula. It is thought that the
post-larvae which ingested the latter five diatom species could not readily absorb the
diatom cell contents as the cell wall was intact and thus their growth rates were low.

Keywords: Benthic diatom; Dietary value; Feeding; Growth, Nutrition; Post-larval abalone

1. Introduction

Abalone is an economically important group of gastropods in many countries.

The culture of abalone has also made great advances, especially in Japan. In 1991,

* Corresponding author.

0022-0981/95/$09.50 0 1995 Elsevier Science B.V. All rights reserved

SSDI 0022-0981(95)00099-2
190 T. Kawamura et al. I J. Exp. Mar. Biol. Ecol. 194 (1995) 189-199

about 26 million juvenile abalone were produced in seed production hatcheries

throughout Japan and released into the natural environment to enhance the stock
(Japan Fisheries Agency and Japan Sea-Farming Association, 1993). However,
the basic ecology of abalone in their natural habitat, particularly the larval
ecology including trophic behaviour, has not been examined in detail (McShane,
1992).
Benthic diatoms are considered the principal food for post-larval and early
juvenile abalone below the size at which they can consume macroalgae (Ino, 1952;
Tomita & Tazawa, 1971). In the seed production hatcheries of abalone, naturally
occurring benthic diatoms on plastic plates have been used as an initial food for
larval abalone (Kikuchi, 1965; Seki, 1980; Ebert & Houk, 1984). However, details
of their feeding habits such as the selection of diatom species are restricted (Seki
& Kan-no, 1981a; Norman-Boudreau et al., 1986; Matthews & Cook, 1995), and
the dietary value of different diatom species has also received little attention (Uki
et al., 1981; Ohgai et al., 1991; Kawamura & Takami, 1995).
In our previous paper, we showed that the extracellular substances of diatoms
are important food for post-larval abalone up to the size of 800 pm in shell length
and that these larval abalone can grow well without high levels of absorption of
the diatom cell contents (Kawamura & Takami, 1995). This study examines the
feeding behaviour and the growth of post-larval abalone (1-2 mm in shell length)
Haliotis discus hannai fed on nine species of benthic diatom. Digestion efficiencies
of abalone feeding on the nine diatom species were measured by observing the
fecal material of post-larval abalone. Of the nine species of diatom, only four
supported high growth rates in post-larval abalone and we suggest that this is due
to the physical charasteristics of the diatom species as to whether the cell wall of
the diatom is ruptured on grazing or not. We discuss the dietary value of the
diatom species for the growth of post-larval abalone up to 2 mm.

2. Materials and methods

Larval abalone were reared in October 1991 and May 1992 following the
methods described by Uki & Kikuchi (1984) at Iwate Prefectural Nanbu Sea-
Farming Center. After about 4 wk of initial rearing on plastic plates covered with
naturally occurring benthic diatoms in running seawater at 20C the post-larval
abalone were transported still on the plates to Tohoku National Fisheries
Research Institute within 4 h.
Nine monocultured species of benthic diatom (Table l), which were originally
isolated from a culturing water tank of abalone at Tohoku National Fisheries
Research Institute and were grown in modified Jorgensens medium (Jorgensen,
1962) containing an addition of 0.05 pg/l of vitamin B,,. The diatoms were
cultured in 6-well polystyrene tissue culture plates (16.8 ml in volume and 9.4 cm2
in bottom surface area per well; Corning). The density of each diatom species on
the bottom surface of the wells at the beginning of the experiments is shown in
Table 1.
 3
3
i?
Table 1 z
Mean cell length, width, initial density, and the growth form of the nine benthic diatom species used in the experiments a
9
Species Cell length Cell width Initial density Growth form* b
?
(ym, mean *SE) (cells/cm*, mean L-SE)
Type Adhesive strength 5
Exp. 1 Exp. 2
P
P
Achnanthes brevipes var. intermedia 28.8 ? 0.6 10.7 k 0.4 (2.7 2 0.3) X lo4 E +++
Achnanthes longipes 61.5 2 1.4 29.0 k 2.0 (5.1 5 0.6) X lo3 E +++ 5
?
Amphora angusta var. ventricosa 38.8 2 0.6 14.6 2 0.4 (4.5 ? 0.2) x lo4 (6.1 ? 0.4) x lo4 A +
t0
Cocconeis scutellum var. parva 19.5 k 0.3 13.1 k 0.5 (2.4 2 0.1) x lo5 (5.1 ? 0.5) x lo* B +++ 6
?
Cylindrotheca closterium 70.5 k 1.5 5.2 2 0.3 (5.9 k 0.4) x lo4 A +
h
Navicula ramosissima 12.8 z 0.3 4.3 2 0.1 (3.6 k 0.2) x 10 (8.8 2 0.6) X 10 A + B
(1.9 5 0.2) x lo5 + -
Nitzschia sp. 46.1 k 0.6 6.0 2 0.2 A
Pleurosigma sp. 138.1 2 2.0 22.6 0.6 (1.1 k 0.1) x 10 A + %
Synedra investiens 53.3 2 0.6 10.1 k 0.4 (4.3 k 0.4) x lo4 (5.8 -c 0.6) x lo4 C ++ 2
2
* Growth forms of pennate benthic diatoms were classified into seven types, A-G, based on their mode of attachment, whether solitary or colonial forms, *
and by their motility and adhesive strengths (see Kawamura & Hirano, 1992).
k

_ ._
192 T. Kawamura et al. I J. Exp. Mar. Bid. Ecol. 194 (1995) 189-199

Experiments were carried out twice (Exp. 1: November 1991, Exp. 2: June
1992). In Exp. 1, 18 post-larval abalone (0.9-1.6 mm in shell length) in six
replicate wells (three individuals per well) were reared on each species of diatom
(seven species, Table 1) at 20C 3000 lux with a 12 h L:D photo-cycle. In Exp. 2,
12 post-larval abalone (1.1-1.6 mm in shell length) in six replicate wells (two
individuals per well) were reared on each diatom species (six species, Table 1)
under the same condition as in Exp. 1. The culture medium for the diatoms in the
wells was changed with filtered (Millipore HA filter, 0.45 pm) seawater prior to
the addition of the post-larval abalone. The feeding behaviour of the post-larval
abalone was observed and the shell length measured individually over a period of
11-14 days at intervals of 3-7 days using a monitor video camera system
connected to an inverted optical microscope. At the time of the observations, the
seawater in the wells was changed and all fecal material removed. Diatoms in all
the wells were abundant throughout the entire experiments. The daily growth
rates of abalone were calculated for each individual, except for specimens that had
died or consistently showed no feeding behaviour indicating that they were dying.
To measure the digestion efficiency of abalone fed on the nine benthic diatom
species, three abalone (1.5-2 mm) were reared on each diatom species in a Petri
dish (5 cm in diameter and 1 cm in height) at 20C 3000 lux with a 12 h L:D
photo-cycle. Diatoms were grown on the bottom surface of the Petri dishes with
modified Jorgensens medium. The culture medium for the diatoms in the Perti
dishes was changed with filtered seawater before adding the abalone. Just before
adding the abalone, the number of whole, live diatom cells and the number of
broken, dead ones in the Petri dishes were counted using the inverted optical
microscope. After a few days of sufficient feeding, the abalone were transferred to
individual Petri dishes filled with filtered seawater without diatoms. Feces
excreted by the abalone in the individual Petri dishes were picked up on to glass
slides as soon as possible, and the number of whole, live diatom cells and that of
broken ones counted under a light microscope. Digestion efficiency of a diatom
species was calculated as follows:
Digestion eficiency (%) =(l - L/L,) X 100

where L is the mean percentage of whole, live cells to the total diatom cells in the
feces and L, is the mean percentage of whole, live cells to the total diatom cells in
the diatom community on the bottom surface of the Petri dish before grazing.

3. Results

All post-larval abalone fed on the nine species of diatom showed active feeding
behaviour from the initiation of the experiments. The diatom chromatophores
were observed in their stomachs and the ingestion of diatom cells were confirmed
by direct observations of feeding behaviour using the inverted optical microscope
apparatus. However, marked differences were observed between the growth rates
of abalone fed on the nine diatom species (Figs. 1 and 2). In Exp. 1, the mean
 T. Kawamura et al. I J. Exp. Mar. Biol. Ecol. 194 (1995) 189-199

0 2 4 6 8 10 120 2 4 6 8 10 120 2 4 6 8 10 120 2 4 6 8 10 12

Rearing time (days)

Fig. 1. Growth of individual post-larval abalone fed on seven species of benthic diatom in filtered
seawater at 2OC, 3000 lux with a 12 h LID photo-cycle (Exp. 1).

daily growth rates

of abalone fed on Cocconeis scutellum var. parva and
Cylindrotheca were significantly higher than those fed on the other five
closterium
species of diatom (comparisons for all pairs using Tukey-Kramer test, p < 0.05;
Fig. 1, Table 2). In Exp. 2, the mean daily growth rates of abalone fed on
Achnanthes brevipes var. intermedia and Achnanthes longipes were significantly
higher than those of abalone fed on the other four species of diatom (Tukey-
Kramer test, p < 0.05; Fig. 2, Table 2). The growth rate of the abalone fed on
Cocconeis scutellum var. parva was slower in Exp. 2 than in Exp. 1 (Table 2) and
there was no significant difference between the growth rates of abalone fed on
Cocconeis scutellum var. parva, Amphora angusta var. ventricosa and Navicula
ramosissima, contrary to the result of Exp. 1.
The digestion efficiencies of four diatom species, Achnanthes brevipes var.
intermedia, Achnanthes longipes, Cocconeis scutellum var. parva and Cylin-
drotheca closterium were high (Table 3). In contrast, those of the other five
species of diatom were low (Table 3) and many whole, live diatom cells were
observed in the fecal material of abalone fed on them. The cell walls of the former
four diatom species were observed to be easily broken open by the radula when
they were grazed by the abalone, whereas most of the cells of the latter five
species were ingested and excreted whole without any deformation of the cell
walls.

4. Discussion

The post-larval abalone fed on the four diatom species: Achnanthes brevipes
var. intermedia, Achnanthes longipes, Cocconeis scutellum var. parva and Cylin-
 P

Table 2
Mean daily growth rates of post-larval abalone fed on nine species of benthic diatom
b
Diatom species Daily growth rate Number of abalone* Rearing duration
(pm/day, mean *SE) e
(days)
Exp. 1 Exp. 2
Exp. 1 Exp. 2 Exp. 1 Exp. 2 ;
Achnanthes brevipes var. intermedia 57.2 2 2.4 12(0,0) 13 4

Achnanthes longipes 47.8 +- 3.8 lO(2,O) 13 $

Amphora angusta var. venlricosa 29.0 k 1.4 31.7 ? 3.5 18(0,0) 11(1,0) 12 13
Cocconeis scutellum var. parva 46.4 2 2.5 36.0 -c 2.2 13(4,1) 12(&O) 11 14 tC
-.
Cylindrotheca closterium 50.1 k 3.3 14(2,2) 11 :
Navicula ramosissima 21.1 2 1.3 24.4 + 1.3 13(&O) 9(3,0) 12 14 3
Nitzschia sp. 13.6 z 1.3 18(0,0) 12 e

Pleurosigma sp. 24.2 2 2.1 t6(L 1) 11 :

Synedra investiens 20.5 -c 1.9 15.7 2 2.1 16(2,0) lO(l,l) 11 13 2
* Number of abalone used for the calculation of mean daily growth rate, with the numbers of dead ones and dying individuals in parentheses, respectively. %
b
Total numbers of abalone used on each diatom species in Exp. 1 and Exp. 2 were 18 and 12, respectively.
5
 T. Kawamura et al. I J. Exp. Mar. Biol. Ecol. 194 (1995) 189-199

2.4 Achnanthes brevipes l Achnonthes longipes

f 1.01
2 Cocconeis scutellum Naviculo ramosissima Synedra investiens
a 2,4

UT. porva
Z n=12
2.2
w
2.0 i

Rearing time (days)

Fig. 2. Growth of individual post-larval abalone fed on six species of benthic diatom in filtered
seawater at 2OC, 3000 lux with a 12 h L:D photo-cycle (Exp. 2).

drotheca cfosterium, which supported high growth rates (Figs. 1 and 2, Table 2)
also had high digestion efficiencies (Table 3). These results suggest that the
differences in growth rates were due primarily to the differences in the digestion
efficiency of the abalone fed on the diatom species. Kawamura & Hirano (1992)
classified the growth forms of benthic pennate diatoms into seven types based on

Table 3
Digestion efficiency of nine diatom species grazed by post-larval abalone (1.5-2 mm in shell length)

Diatom species Percentage as whole live cells Digestion efficiency*

(% mean *SE) (%)
Before grazing (L,)* In the fecal
material (L)*

Achnanthes brevipes var. intermedia 97.0 k 1.6 34.6 2 3.6 64.3

Achnanthes longipes 69.5 2 1.7 19.8 2 4.8 71.5
Amphora angusta var. parva 70.9 2 3.9 65.1 5 2.1 8.2
Cocconeis scutellum var. parva 74.6 2 5.6 11.7 2 0.8 84.3
Cylindrotheca closterium 100.0 -t 0 Ok0 100.0
Navicula ramosissima 72.8 k 2.4 60.8 +- 4.4 16.5
Nitzschia sp. 85.1 k 3.0 77.9 k 4.3 8.5
Pleurosigma sp. 97.8 2 0.1 94.1 2 0.8 3.8
Synedra investiens 98.9 k 1.1 93.4 k 0.7 5.6

* Digestion efficiency (% ) = (1 - L/1;,) X 100.

196 T. Kawamura et al. I J. Exp. Mar. Biol. Ecol. 194 (1995) 189-199

their mode of attachment, whether solitary or colonial forms, and by their motility
and adhesive strengths. Types of the nine species used in this study were all
solitary species, A, B, C and E (Table 1, Fig. 3). The adhesive strengths of types B
and E diatoms are high, whilst those of types A and C are relatively low (Table
1). The cells of Achnanthes brevipes var. intermedia, Achnanthes longipes and
Cocconeis scutellum var. parva, which have E, E and B growth forms, respectively
(Table l), are easily broken open when they are grazed by abalone because of
their high adhesive strengths. The types of growth form of the other six species
are A and C, and because of the low adhesive strength of these types, they are
easily ingested by abalone without any deformation of the cell walls with the
exception of Cylindrotheca closterium. As the cell walls of this species are only
weakly silicified (Reimann & Lewin, 1964) their cells are generally easily broken
open by the radula of abalone. From direct observations, we suggest that post-
larval abalone cannot digest the diatom cell contents in their alimentary canal
without first the splitting open of the cell wall by the radula. So we think that the
post-larvae which ingested the five species (Amphora angusta var. ventricosa,
Navicula ramosissima, Nitzschia sp., Pleurosigma sp. and Synedra investiens) could
not break open the cells and therefore could not absorb the diatom cell contents
leading to slow growth rates.
In addition to the diatom cell contents, the extracellular mucus of diatoms
appears to be an important food source for the early post-larval abalone up to the
size of 800 pm in shell length (Kawamura & Takami, 1995). Post-larval abalone

A B

Fig. 3. Schematic diagram of the four types (A, B, C and E) of diatom growth form shown by diatom
species used in this study. Growth forms are based on the seven types of structures defined by
Kawamura & Hirano (1992). Type A (gliding prostrate type): solitary cells with a prostrate form and
swift gliding movement, adhesive strength is low; Type B (adhesive prostrate type): solitary cells with a
prostrate form and slow movement, adhesive strength is very high; Type C (non-motile upright type):
non-motile, solitary cells or simple fan-shaped colonies standing upright on the substratum; Type E
(mucous thread solitary type): solitary cells or short belt-shaped colonies attaching to the substratum
with a mucous thread.
 T. Kawamura et al. I J. Exp. Mar. Biol. Ecol. 194 (1995) 189-199 197

(<BOO pm) fed on ZVavicuZa ramosissima, the digestion efficiency of which was
also low, grew as well as ones fed on Cylindrotheca closterium, this is probably
due to the ingestion of mucus (Kawamura & Takami, 1995). In contrast, the
results of this study showed post-larval abalone of l-2 mm in shell length require
high levels of absorption of diatom cell contents in addition to any extracellular
substances for favorable growth. The growth rate of the early post-larval abalone
(<BOO ,um) fed on Cocconeis scutellum var. parva was low (Kawamura & Takami
1995) this is because the abalone could not ingest the diatom cells owing to their
high adhesive strength at this stage, in addition to the relatively small quantities of
extracellular mucus exuded by the diatom. This study demonstrates that post-
larval abalone of l-2 mm in shell length can tear off and simultaneously break
open the cell wall of the highly adhesive diatom species such as Cocconeis
scutellum var. parva by the radula and are therefore able to absorb the cell
contents. Although slight variations were observed in the mean growth rates
between experiments, for example Cocconeis scutellum var. parva (Table 2, Figs. 1
and 2), possible differences in the quantity of ingested diatom cells due to some
factors such as the difference in the adhesive strength between diatom strains may
explain these variations. The results of this study, however do show clearly that
the physical charasteristics of the diatom species as to whether the cell wall of the
diatom is ruptured on grazing or not is one of the most important factors
determining its dietary value for the growth of post-larval abalone of l-2 mm.
It has been reported that diatom species with highly adhesive solitary growth
forms such as Achnanthes spp. and Cocconeis spp. are hard to be grazed by many
kinds of gastropod (Nicotri, 1977; Kesler, 1981; Steinman et al., 1989; Kawamura
et al., 1992) including adult and juvenile abalone (>l cm) (Ioriya & Suzuki, 1987;
Suzuki et al., 1987). Although it is not known yet whether adult and juvenile
abalone can readily digest the cell contents of loosely adherent diatoms, these
diatom species are selectively ingested by abalone (Ioriya & Suzuki, 1987; Suzuki
et al., 1987). This study, however, shows that post-larval abalone (l-2 mm) need
to graze on strongly attached diatoms for favorable growth. It thus appears
probable that post-larval abalone feed mainly on the diatoms which remain after
the grazing of the adult abalone and other gastropods. In seed production
hatcheries, plastic plates covered with diatom films which have previously been
grazed by juvenile abalone (>l cm) are used as a substratum for the settlement of
veliger abalone larvae, and the diatoms which remain on these plates such as
Cocconeis spp. are used as food for the post-larval abalone (Seki, 1980; Ebert &
Houk, 1984). Newly metamorphosed post-larval abalone on the plates feed
probably on the extracellular substances of diatoms (Kawamura & Takami, 1995)
and/or mucous trail secreted and left by the juvenile abalone (Seki & Kan-no,
1981b). Abalone from approximately 1 mm can then consume the diatoms of the
strongly adhesive solitary forms which remain after grazing by larger juveniles
(>l cm). Matthews and Cook (1995) also found from gut content analyses that
prostrate diatoms such as Cocconeis spp. which remain on plates after the grazing
of juveniles were the preferred food for post-larvae of Haliotis midae. However,
prostrate diatoms were also considered as good food for early post-larvae (7-14
198 T. Kawamura et al. I .I. Exp. Mar. Biol. Ecol. 194 (1995) 189-199

day old), this being contrary to the results for Huliotis discus hannai (Kawamura
& Takami, 1995).
In the natural environment, larval abalone are reported to settle exclusively on
crustose coralline red algae (Saito, 1981; Morse and Morse, 1984; Shepherd &
Turner, 1985; McShane & Smith, 1988). The surface mucus, cuticle and epithelial
contents of coralline algae and bacteria present on them are thought to be food
sources of early post-larval abalone (Morse & Morse, 1984; Garland et al., 1985).
In areas dominated by crustose coralline algae, diatom species with strongly
adhesive solitary forms such as Cocconeis spp. are observed to be the dominant
benthic diatoms (Kawamura et al., 1992). Larval abalone which settle and
metamorphose on coralline algae can probably find sufficient food sources up to
at least a size of 2 mm. However, the relationship between crustose coralline algae
and benthic diatoms growing on them have not been well studied, and further
studies are needed to understand the feeding ecology of larval and juvenile
abalone in the natural environment.

Acknowledgements

We wish to thank Dr. T. Seki for his valuable comments. The critical reading of
this manuscript by Drs. S.A. Shepherd and C.P. Norman are gratefully acknowl-
edged. This study was supported by a grant from the BIOCOSMOS project, the
Ministry of Agriculture, Forestry and Fisheries Agency, Japan.

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