Effect of feeding attractants on the behaviour and performance of juvenile penaeus monodon fabricius r hartati, m r p briggs

Aquaculture and Fisheries Management 1993, 24, 613-624

Effect of feeding attractants on the behaviour and
performance of juvenile Penaeus monodon Fabricius
R. HARTATI & M. R. P. BRIGGS Institute of Aquaculture, University of Stirling,
Stirling, Scotland

Abstract. A study was conducted to obtain a measure of the potency of some potential and
commercially used feeding attractants for Penaeus monodon Fabricius. Behavioural trials
monitoring the feeding response of the shrimp were used to gauge the attractant qualities of the
substances, A growth trial recording the feed intake, feed assimilation, growth, food
conversion and survival rates of the shrimp was used to assess further the feeding stimulant
properties of the substances.
Replicate groups of juvenile shrimp were fed semi-purified diets containing 1-5% by weight
of a range of potential feeding attractants.
In the behavioural trial, diets containing taurine and a yeast extract were found to be
significantly preferred to the control and all other diets,'However, none of the substances
appeared to act as potent feeding stimulants, producing statistically similar feed intake and
assimilation rates to the control diet. However, taurine and an amino acid mixture designed to
mimic a clam extract promoted the best performance of the attractants tested in terms of growth
rate and feeding efficiency.
Overall, the behavioural response of the shrimp to the feeding attractants was found to show
similarities to the effects of attractant supplementation of feed on subsequent ongrowing
performance, but not significantly so.

Introduction

Feeding attractants can play an important role in enhancing food detection and feeding
stimulation. Their role can be especially important in slow-feeding crustaceans, reducing
leaching of essential water-soluble nutrients present in the diet (Deshimaru & Yone 1978;
Heinen 1980) by increasing the rate of food consumption.

Many studies have been made of the factors involved in stimulating feeding in
crustaceans, particularly with lobsters, crabs and freshwater prawns. However, little is
currently known about the effect of attractants on the feeding behaviour and ingestion rate of
Penaeus monodon Fabricius, one of the most widely cultivated crustacean species.
Most of the studies on feeding attractants to crustaceans have analysed the effects of a
range of concentrations of various potential attractants on behaviour, particularly with
regard to their effects on the crustacean chemosensory apparatus (Carr 1978; Carr &
Thompson 1983; Carr, Netherton & Mistead 1984; Carr & Derby 1986; Harpaz, Kahan,
Galun & Moore 1987; Harpaz & Steiner 1987, 1990; Nakamura 1987). In addition, growth
trials have been conducted with crustacean species using both commercial (Farmanfarmaian,
Lauterio, Tejada, Manger, Moore & Park 1979; Murai, Sumalangkay & Pascual 1981; El
Hag 1984; Costa-Pierce & Laws 1985) and semi-purified diets (Deshimaru & Yone 1978;
Pascual 1980) containing various attractants.
Correspondence: Mr M, R, P, Briggs, Institute of Aquaculture, University of Stirling, Stirling FK9 4LA,
Scotland,
613


614

R. Hartati & M. R. P. Briggs

Olfactory stimulation by a variety of chemicals has been demonstrated by many authors in
decapod crustaceans (Heinen 1980). Substances (usually of low molecular weight, <1000)
which were found to be strong stimuli when tested singly on decapods were amino acids,
nucleotides and quaternary ammonium compounds, while sugars, alcohols, starches, fatty
acids and several other compounds have usually given lesser responses or none at all (Heinen
1980; Carr & Derby 1986).
The aim of the present work was to obtain a measure of the potency of various potential
and commercial feeding attractants to P. monodon by way of both behavioural and

ongrowing studies. This two-phase approach also allowed an assessment as to whether the
behavioural responses to feeding attractants can predict the effects of attractant supplementation on subsequent ongrowing performance. This measure is important as the addition of
feeding attractants might elicit an increase in appetite and subsequently food intake and
growth (Lindstedt 1971) and also improve shrimp survival and food conversion (Heinen
1980).
Materials and methods
Stage 1: Diet choice behavioural trial
The experimental shrimp were obtained from the University of Stirling Institute of
Aquaculture hatchery and reared in a holding tank until they reached the desired weight.
Prior to the trials the shrimp were held in a common holding tank equipped with water
recirculation and were fed twice daily with chopped mussel and green beans at the rate of
20% wet body weight/day. Following each behavioural trial, each group of shrimp was kept in
individual cages made of mesh-covered PVC pipe measuring 16cm in length and 11cm in
diameter. All cages were placed in the holding tank so that they could be used for subsequent
trials after allowing at least 2 days for shrimp recovery.
A semi-purified diet was used as a medium for testing the potential feeding attractants as
has been used in earlier studies with Macrobrachium rosenbergii De Man (De Proenca 1990),
Penaeus japonicus Bate (Deshimaru & Yone 1978) and P. monodon (Pascual 1980).
The composition and proximate analysis of the basal diet is presented in Table 1. Crude
protein was determined by Kjeldahl analysis, expressed as total N x 6-25, crude lipid by the
Soxhlet method, ash by combustion at 450°C for 12h and carbohydrate as the remainder of
the dry weight. Moisture was determined by drying at 105°C for 24h and gross energy by
combustion in a micro-bomb calorimeter. All results were based on the mean of triplicate
analyses.
The attractants, included at a rate of l-57o in each diet (except the control) were as
follows:
Diet 1 (T)
Diet 2 (B/G)
Diet 3 (AA)

Diet 4 (AMP)
Diet 5 (TMAH)

: Taurine
: A mixture of 50% betaine and 50% glycine
: A mixture of L-amino acids based on the composition of an extract
of short-necked clam {Tapes japonica: Deshayes) as described by
Deshimaru & Kuroki (1974) (Table 2)
: Adenosine 5'-monophosphate
: Trimethylamine hydrochloride


Ejfect of feeding attractants on P. monodon

piet 6 (F)

Diet 7 (YE)
Diet 8 (C)

615

: Finnstim, a commercial chemoattractant developed by the Finnish
Sugar Co. Ltd. The product is based on betaine and a mixture of
stimulatory amino acids
: Yeast extract, a natural flavour enhancer
: Control diet, using 1-5% a-cellulose in place of an attractant.

All shrimp were starved for 48h prior to the trials. Each of the seven test diets and the
control were tested against each of the other diets in paired comparisons (Table 3). Each set
of two diets was tested against one another for the relative attractiveness of the diets. Every

combination of diets was replicated five times for statistical comparison. A total of 28
combinations of diets was tested. Tests consisted of a single group of eight shrimp being
introduced into an aquarium and being offered a choice of two diets.
Five replicate groups of eight juvenile shrimp, mean weight 35 ± 5mg, were tested in
separate perspex tanks each measuring 36 x 19-5 x 28cm. One end of each tank was divided
into two freely available, equal compartments (X and Y) by a piece of black acrylic plate
measuring 12 cm^. A section of black PVC pipe was fixed to the bottom of the tanks at the
opposite end as a shelter. The tanks were half-filled with 91 of sea water at 25%o salinity. The
Table 1. Basal composition of the diets
Ingredient
Casein (vitamin free)
Gelatin
Dextrin
Astaxanthin
Vitamin premix*
Mineral premix*'
Cod liver oil
Lecithin
Cholesterol
a-cellulose
Attractant/a-cellulose***
Source""
Proximate analysis
Protein
Lipid
Carbohydrate
Ash
Fibre
Moisture
Total energy (Kcal/g)

% dry weight
31-0
10-2
35-7
0-1
2-0
4-0
6-0
3-0
0-5
6-0
1-5

.

40-8
9-1 •
40-2
3-8
6-1
8-0
4-9

* Vitamin premix (in lOOg premix): thiamine-HCl 150mg,riboflavin500mg, niacinamide 2000mg, pantothenic acid
750 mg, inositol 10000 mg, biotin 15 mg, folic acid 37-5 mg, pyridoxine-HCl 150 mg, P-aminobenzoic acid 1000 mg,
choline chloride 20000mg, ascorbic acid 25000mg, a-tocopherol acetate lOOOmg, menadione (sodium bisulphide)
lOOmg, p-carotene 100mg, cholecalciferol 15mg, cyanocobalamine lmg, a-cellulose powder 39181-5mg.
** Mineral premix (in lOOg premix): CaHPO4.2H2O 75-0276g, MgSO4.7H2O 12-75g, NaCI 6 Og, KCl 5-Og,
FeSO4.7H2O 0-25g, ZnSO4.7H2O 0-55g, CUSO4.5H2O 0 0785g, MnSO4.4H2O 0 2538g, COSO4.7H2O 0-0478g,

CaIO2.6H2O 0 0295g, CrCl3.6H2O 0 0128g.
*** a-cellulose used in control diet instead of attractant.
**** All chemicals and other dietary ingredients obtained from Sigma Chemicals Ltd except lecithin from Lucas
Meyer Ltd.


616

R. Hartati & M. R. P. Briggs

Table 2. Composition of the amino acid mixture
Amino acid

% dry weight

Taurine
Aspartic acid
Threonine
Serine
Glutamic acid
Proiine
Glycine
Alanine
Cystine
Valine
Methionine
Leucine
Isoleucine
Tyrosine
Phenylalanine

Histidine-HCl
Lysine
Arginine

43-6
I.4
0-8
1.6
6-8
1-0
21-6
8-6
0-4
1-0
0-6
1'2
0-6
10
1-4
0-6
1-6
6-2

temperature of the sea water in the tanks was kept constant at 29 ± 1°C. All tanks were placed
in dim lighting conditions on a 12:12h light:dark cycle.
The diets were offered at the rate of 20% wet body weight at the beginning of each trial
either in compartment X or Y, while the same amount of control diet or another test diet was
placed in the other. Aeration was turned down during the trial. The trial was started and the
number of shrimp located in each compartment counted every minute for a total of 30 min.
The q^ttractivene^s of the diets is expressed in terms of feeding activity as shrimp time

units, i.e. the number of shrimp feeding on each diet, each minute for 30 min. These data
were converted to give a measure of the percentage of the maximum possible response to
each diet in order to gauge the overall attractiveness of the diets. The results were ranked and
Table 3. Test group comparisons used in the behavioural trials
Test diet

T

•j"

B/G
AA
AMP
TMAH
F
YE

.

B/G

AA

AMP

TMAH

F

•

•

•

•

•

.

.
•

,.
"

.
*

T — Test diet 1 (Taurine).
B/G — Test diet 2 (Betaine-Glycine mixture).
A A — Test diet 3 (Amino acid mixture).
AMP — Test diet 4 (AMP).
TMAH — Test diet 5 (TMAH).
F — Test diet 6 (Finnstim).
YE — Test diet 7 (Yeast extract).
C — Control diet (Without attractant).

YE

C


Effect of feeding attractants on P. monodon

617

analysed for significant differences between paired samples by the Wilcoxon-Mann-Whitney
two-samples test (Steel & Torrie 1980),
Stage 2: Feeding trial

The diets analysed in the behavioural trial were retested in a feeding trial. Results of the
behavioural trials were compared with these results to determine whether the subsequent
ongrowing performance of shrimp could be predicted by behavioural studies.
Two replicate groups of five shrimp were used in this stage of the study. Experimental
animals, rearing conditions and test diets were the same as those used in stage 1.
All shrimp were starved for 48 h prior to the trial. The feeding regime was 10% of the wet
body weight/day given in two equal portions at 0900 and 1600 h for 6 days/week with no
feeding on the weekly sampling day.
After l-5h (because it was found that shrimp started producing faeces 2h after first
feeding) all food remaining uneaten was siphoned, filtered through Whatman filter paper no,
54 and dried in the oven for 24 h to calculate the actual food intake based on the following
equation:
Daily feed intake (g feed/g shrimp/day)
Fl
[{wt, + Wt2)/2]d

One and a half hours later, the faeces produced were siphoned, filtered and dried in the
oven to determine feed assimilation efficiency based on the following equation:

Food assimilation efficiency (%)

Where
Fl

: Food ingested (g/wk)
= food given — food wasted (g)
FP : Faeces produced (g/wk)
vv^i : Mean wet body weight at day 1 (g)
Wt2 : Mean wet body weight at day n (g)
d
: Time (days)
All uneaten food and faeces were collected before both the morning and evening feedings
and the water lost during each siphoning was replaced. This resulted in a water exchange rate
of approximately 30%/day,
The feeding experiment was conducted for 28 days. The shrimp were weighed
individually and counted every 7 days and the feeding regime adjusted after each weighing.
The performance of each test diet was evaluated as specific growth rate (SGR), food
conversion ratio and survival rate.
One-way analysis of variance (arc-sine transformed for the percentage values) and
Duncan's multiple range tests (P < 0-05) (Steel & Torrie 1980) were employed to establish
the effect of feeding attractants on shrimp feed intake and assimilation, growth, survival and
feeding efficiency.


618

R. Hartati & M. R. P. Briggs

Results

Stage 1: Diet choice behavioural trial

The results of the paired sample comparisons were looked at in terms of feeding activity
(shrimp-time units), i.e. the number of shrimp feeding on each diet for each of the 30 min of
each trial. Results showed that most of the substances, particularly taurine, the yeast extract
and the amino acid mixture, were more attractive (P < 0-05) than the control while the
betaine/glycine mixture and adenosine 5'-monophosphate were not.
Results from the response of shrimp to each diet during the paired comparisons as a
percentage of the maximum possible response showed that there were significant {P < 0-05)
differences between diets (Table 4 and Fig. 1). The rank attractiveness of the test diets in
decreasing order were as follows (test diets with a common underline were not significantly
different):

T

YE

F

AA

TMAH

A M P

C

B/G

Overall, the shrimp response data indicated that taurine and the yeast extract were

significantly [P < 0-05) preferred over all other attractants and the control. In addition, the
amino acid mixture and Finnstim were preferred (P < 0 05) over the control, but that
TMAH, AMP and betaine/glycine were not.

Stage 2: Feeding trial

The daily feed intake results showed that there were no significant differences between
treatments and that the shrimp fed on the control diet had a similar daily feed intake (0-35 g/g
shrimp) to those fed the diets containing attractants (Table 5). Among the test diets which
contained attractants, the shrimp fed diet 4 (containing AMP) had the highest daily feed
intake followed by diet 1 (Taurine). Both of these diets also resulted in a fairly constant feed
ingestion rate, while that of shrimp fed any of the other diets tended to decline during the trial
period.

Table 4. Response of shrimp fed the test diets as a percentage of the maximum possible response
Test diet
Replicate

1
2
3
4
5
6
7
X
+SEM

T

B/G

AA

AMP

310

120

27-0
33-4
36-0
33-2
30-8
31-0

10-1

14-6
16-6
23-2
15-8
14-5
25-6
28-8
19-9
2-22

318

1-06

5-6
13-6
13-0
10-4

5-0
9-9
1-30

TMAH

F

YE

C

14-6

102

9-0

24-0
10-2
15-4
17-2
12-4

24-6
17-7
2-08

19-8
31:0
23-2
17-2
23-0
16-4
20-8
19-6
1-31

27-8
31-0
25:6
31 6
34-6
33-4
37-4
31-6
1-52

98
7-2

12-2
18-6
15-6

7-2
19-6
13-2
1-82

13-0
14-6

98
9-0
14-6

IM
110


Effect of feeding attractants on P. monodon

619

40 -I

E
3

30-

O

c
o
o.

20 -

10 c
c

at

5 .
4
Test Diet
Figure 1. Mean response (+SEM) of shrimp to test diets as a percentage of the maximum response.

The differences in feed assimilation efficiency between the test diets were insignificant (P
< 0-05). However, the highest feed assimilation was achieved by shrimp fed on diet 4 (AMP)
and the lowest by shrimp fed on diet 7 (YE) (Table 5),
Statistical analysis showed that weight gain of shrimp fed on diets 1 (T) and 3 (AA) was
significantly (P < 0-05) higher than that of shrimp fed on diet 2 (B/G), but not than those fed
the control or any other test diet. The SGR of shrimp fed on diets 1 (T) and 3 (AA) was also
significantly (P < 0-05) better than shrimp fed on diet 2 (B/G) (Table 5 and Fig, 2),
Table 5. Effect of the feeding attractants on the daily feed intake, feed assimilation efficiency, weight gain, specific
growth rate, food conversion ratio and survival of the shrimp (+SEM) over 28 days
Test
diet

Attractant

1

Taurine

2

4

Betaine/
Glycine
Amino acid
mixture
AMP

5

TMAH

6

Finnstim

7

Yeast
extract
Control

3

8

DFI

FAE

WG

SGR

FCR

SR

0-33"
(0-02)
0-28"
(0-05)
0-29"
(0-06)
0-35"
(0-05)
0-25"
(0-05)
0-28"
(005)
0-26''
(0-05)
0-35"
(0-07)

92-44'"'
(0-41)
92-75""
(1-22)
93-81"
(0-80)
93-90"
(0-74)
92-69""
(0-87)
92-27""
(0-15)
89-732"
(0-29)
93-45"
(0-14)

1-32"
(0-20)
0-45"
(0-10)
1-21"
(0-26)
0-79""
(0-10)
0-85"?
(0-03)
0-81""
(0-22)

0-89""
(0-13)
0-79""
(0-12)

3-45"
(0-46)
1-52"
(0-31)
3-29"
(0-33)
2-45""
(0-17)
2-53""
(0-06)
2-46""
(0-53)
2-55""
(0-31)
2-25""
(0-28)

0-91"
(0-01)
2-75°
(0-50)
1-08"
(0-05)
1-60""
(0-08)

l-23b
(0-02)
1-49""
(0-54)
1-39""
(0-03)
2-05""
(0-52)

80"
(0)
60"
(0)
60"

(0)
60"
(0)

50°
(0)
60"
(20)
50"
(10)
70"
(10)

DFI — Daily feed intake (g),
FAE — Feed assimilation efficiency (%),

WG —Weight gain (g),
SGR — Specific growth rate (% body weight/day),
FCR — Food conversion ratio,
SR — Survival rate (%),
"h — For single criterion, mean values in the same column bearing the same superscript are not significantly different
{P < 0-05),


620

R. Hartati & M. R. P. Briggs

4 •

•a

3

T

T

3-

T

T
T3
O

T

T

_

'S

1-

LJ.

0-

3ntro

;<

nnsti

m

0.

MAH

1-

aurin

0)"'

E

T

^'

4
5
6
Test Diet
Figure 2. Mean SGR (+SEM) of shrimp fed the test diets.

Diet 2 (B/G) also produced the poorest FCR of 2-75, which was significantly (P < 0-05)
worse than the FCR for the diets containing taurine (diet 1) at 0-91, the amino acid mixture
(diet 3) and TMAH (diet 5) (Table 5). Although the survival rate of shrimp during the trial
was not significantly different between treatments or from the control, the diet containing
taurine as an attractant gave the best survival (80%) and the yeast extract and TMAH the
worst (50%) (Table 5).
Discussion

Diet choice
The diet choice comparison technique adopted in this study was able to reveal significant
differences between the behaviour of shrimp towards diets containing different feeding
attractants. Shrimp were shown to have a significant {P < 0-05) preference for diets
containing taurine and the yeast extract over all other diets, particularly the control,
demonstrating the attractant qualities of these substances. Additionally, the diets containing
these two attractants were seen to elicit the highest (P < 0-05) percentage response figures
(about 32% of the maximum possible) of the diets.

This evidence suggests that P. monodon might have a taurine-specific antennular
chemoreceptor similar to that found in Panulirus argus Travis (Fuzessery, Carr & Ache
1978). This may be related to the fact that taurine is the most abundant p-amino acid in the
prey of shrimp and in their own bodies (Watanabe & Konusu 1972; Fuzessery et al. 1978;
Smith, Miller & Mead 1987). The diet containing the amino acid mixture (43-6% taurine) as
an attractant produced a lesser response than taurine alone, but was still preferred (P < 0-05)
over the control diet. Similar results to these were found by Deshimaru & Yone (1978) with
P. japonicus although they also found glycine to be particularly effective.
In this study the response of shrimp fed a diet containing the betaine/glycine mixture was
not significantly different to those fed the control diet, whereas both glycine and betaine
alone have previously been shown to improve diet attractiveness for P. monodon (Murai et
al. 1981). In contrast, Finnstim, composed largely of betaine, proved more attractive (P <
0-05) to the shrimp than the betaine/glycine mixture and the control in this trial. It may


Effect of feeding attractants on P. monodon

621

therefore be that betaine and glycine are antagonistic when used together as attractants, as
has been found for proline, alanine, arginine and taurine in lobsters due to competitive
interaction for receptor sites (McLeese 1970; Johnson & Ache 1978; Ache, Gleeson &
Thompson 1986), Alternatively, it may be the concentration of betaine that is responsible for
its attractant qualities, although this could not be quantified due to the unknown
concentration of betaine in Finnstim.
A new feeding attractant based upon a yeast extract was tested in this study. Information
on the type of yeast that was extracted to produce this attractant was not available. However,
it apparently acted as a potent feeding attractant for P. monodon, being preferred (P < 0-05)
over all diets except the one containing taurine.
Feed intake and assimilation

Data on feed intake and assimilation can serve as a guide to whether the attractants were also
acting as feeding stimulants. The diet containing taurine promoted a high and consistent feed
intake. This diet was also shown to have excellent attractant qualities from the behavioural
study, Taurine thus seems to be effective as both a feeding attractant and as a mild feeding
stimulant for P. monodon, as was found for P. japonicus (Deshimaru & Yone 1978) and M.
rosenbergii (Smith et al. 1987),
The best overall feed intake and assimilation results, although not significantly different
(P < 0-05), were obtained for the diet containing AMP, which was shown to be a poor
attractant in the behavioural studies. AMP may thus have limited stimulant qualities, without
being attractive to P. monodon. In contrast, AMP has been shown to be a chemoattractant
for the caridean shrimp, Palaemonetes pugio (Carr & Thompson 1983), and M. rosenbergii
(Harpaz, Kahan & Galun 1987; De Proenca 1990), The diet stimulating the next highest
increase in food intake was that containing an amino acid mixture. A similar result was
achieved by Deshimaru & Yone (1978) with P. japonicus fed diets supplemented with the
same amino acid mixture. In previous trials, diets containing TMAH were found to result in a
30-38% increase in food consumption when fed to M. rosenbergii (Costa-Pierce & Laws
1985), This was suggested to be due to their strong faecal odour and the coprophageous
feeding habits bf caridean prawns. In this trial, however, TMAH was not found to increase
feed ingestion or growth rate significantly, although it did act as a mild attractant for P.
monodon.

Feeding trial
Differences in the growth rate of shrimp, although insignificant, followed similar trends as the
attractiveness of the diets during the behavioural trials, Taurine and the amino acid mixture
promoted the best growth rates of the attractants tested. Good growth was also achieved on
diets containing the yeast extract, TMAH, Finnstim and AMP, all greater than the control,
while the betaine/glycine mixture promoted the poorest growth.
These results indicate that in addition to the attractant qualities of taurine, the yeast
extract and the amino acid mixture, these substances may have stimulant qualities, resulting
not in increased ingestion or assimilation of food, but in increased growth. Similar results

have been achieved by Pascual (1980) and Murai et al. (1981) with P, monodon. These


622

R. Hartati & M. R. P. Briggs

authors found that both shrimp and mussel extracts containing high quantities of both taurine
and a mixture of amino acids were very attractive to P. monodon and increased growth and
survival rates due largely to enhanced diet acceptability.
The addition of protein and/or essential amino acids to a diet deficient in these nutrients
might have a role in metabolic processes as well as acting as incitants (Heinen 1980). The
ability of these diets to enhance growth rate may thus have been due to enhanced nutritional
quality, particularly because the test diets used purified ingredients which may lack essential
nutrients and attractive qualities. However, taurine is not regarded as an essential amino acid
(Coloso & Cruz 1980) and other attractants containing amino acids, such as Finnstim and the
betaine/glycine mixture, were not able to promote such a response. Additionally, the amino
acid mixture attractant, based on the short-necked clam (a known favourite food for P.
japonicus, Deshimaru & Yone 1978), contained a mixture of essential and non-essential
amino acids but was not able to promote a better growth or feeding response than taurine
alone.
Analysis of the FCR of shrimp fed the test diets revealed similar results to those with the
growth rate. This measurement thus seemed to be a more sensitive indicator of shrimp
feeding efficiency than assimilation efficiency, and again confirmed results from the
behavioural trial section of this investigation. Only the diet containing taurine promoted
better survival than the control diet, although there were no significant differences between
any of the diets.
A comparison of data from the behavioural and growth trials revealed similarities in the
ranking of the attractiveness of the test diets. This suggests that behavioural studies may be
able to predict subsequent ongrowing performance. Extending the duration of the feeding

trial stage of this study may have resulted in a closer correlation between the two stages of this
investigation and deserves further attention.
Acknowledgments

The authors wish to thank the following staff from the Institute of Aquaculture, University of
Stirling: Dr S. Smith, Mr D. Milroy and Mr A. Porter for their technical support, Dr B. J.
MacAndrew for assistance with the statistical analysis and Dr J. H. Brown, Dr C. J. Fox and
G. S. Haylor for their critical reviews of the manuscript.
We would like to thank the Ministry of Education and Culture, Indonesia and The British
Council for supporting R. Hartati during this work.
These trials were part of work funded by the Overseas Development Administration
(ODA), under Research Project Number R4443 NRG 522/832/8A.
References
Ache B.W., Gleeson R.A. & Thompson H.D. (1986) Mechanism of interaction between odorants at olfactory
receptor cells. Chemical Senses 11, 575.
Carr W.E.S. (1978) Chemoreception in the shrimp, Palaemonetes pugio: The role of amino acids and betaine in
elicitation of a feeding response by extracts. Comparative Biochemistry and Physiology 61 A, 127-131.
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