Aquaculture Research, 2005, 36, 906^911
Feeding level-scaled retention efficiency, growth and
energy partitioning of a marine detritivorous fish, redlip
mullet (Liza haematocheila T. & S.)
Kang Bin1,2 & Wei Wei Xian2
1
Fishery College, Ocean University of China, Qingdao, China
2
Key Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Academy of Sciences,
Qingdao, China
Correspondence: W Wei Xian, Key Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Acad-
emy of Sciences, No. 7, Nanhai Road, 266071 Qingdao, Shandong Province, China. E-mail: wwxian@ms.qdio.ac.cn
Abstract
The e¡ects of feeding level on growth, retention
e⁄ciency, faeces production and energy partitioning
of redlip mullet were studied. A practical diet was
used and fed at six levels from starvation, 1%, 2%,
3%, 4% of body weight (BW) to satiation for 3 weeks.
The temperature was kept at 24  1 1C. Reducing
the feeding amount resulted in signi¢cantly lower
weight gain, and retention e⁄ciency was signi¢-
cantly a¡ected by feeding levels and attained the
maximum at maximum feeding intake. Feeding 2%
BW was the minimum required for ¢sh to maintain
growth. Fish carcass composition under di¡erent
feeding levels could be divided into three groups: (1)
starvation and FL1; (2) FL2 and FL3 and (3) FL4
and satiation, with signi¢cant di¡erences among the
groups but no di¡erences in the groups except that
ash content remained at constant value. Body com-
position of ¢sh of group 2 was close to initial ¢sh.
The thermal-unit coe⁄cient was 0.0381 at satiation,
and signi¢cantly increased with increasing feed-
ing levels. In order to accurately estimate basal
metabolism (HeE), another trial on the relation-
ship between HeE (kJ) and BW (g) was carried
out. An exponential curve as HeE 5 0.1255BW0.8386
explained this relationship. Intake energy (IE) in-
creased from 11.30 to 63.08 kJ per ¢sh, matching
with di¡erent feeding levels. Energy allocated to
growth of IE decreased with reducing feeding
amount. There was a linear relationship between me-
tabolism energy and retention energy in percentage.
906
doi:10.1111/j.1365-2109.2005.01300.
Keywords: Liza haematocheila, feeding level, reten-
tion e⁄ciency, growth, energy
Introduction
Fish with di¡erent feeding habits have di¡erent ways
of transporting nutrients. By feeding on benthic or-
ganic detritus, detritivorous ¢sh release the nutrients
up into the water column in soluble forms that can be
utilized by phytoplankton (Schaus & Vanni 2000), af-
fecting the relative abundance of detritus, dissolved
inorganic nutrients, phytoplankton, zooplankton and
other ¢shes (Ray & Starskraba 2001). Redlip mullet
(Liza haematocheila T. & S.) is the ¢rst species in mari-
culture in China and now a popular polyculture spe-
cies for its scavenging e¡ects (Xian,Yang & Liu 2003),
also as economic by-products.
Basal metabolism (HeE) represents the minimum
use of energy for maintaining life. In ¢sh it is clearly
related to both weight and temperature and normally
estimated by measuring fasting heat production
(Hef). It is mainly a function of the ¢sh body weight
(BW) at a constant temperature and can be described
by the general equation HeE 5 aBWb, where HeE is
the basal metabolism (kJ), BW is ¢sh BW (g), a is a con-
stant which is dependent on species and tempera-
ture, and the exponent b for ¢sh has been reported
to range from 0.50 to 1.00 (Bureau, Kaushik & Cho
2002). Considering the temperature, Cho and Kaush-
ik (1990) observed that the Hef (kJ) of rainbow trout
(Oncorhynchus mykiss W.) as a formula of water tem-
r 2005 Blackwell Publishing Ltd
Aquaculture Research, 2005, 36, 906^911
perature (T, 1C) and BW (kg) could be described as:
Hef 5 ( 1.0413.62 T 0.05 T2) (BW0.824) day 1.
Feeding delivers nutrition to animals for maintain-
ing normal growth, health and reproduction. Nutri-
ents requirements in ¢sh are often quanti¢ed by
dose^response relationships, where diets containing
graded levels of a nutrient are fed and the resulting
growth is measured. The feeding levels have e¡ects
on the e⁄ciency of feed utilization. It has been sug-
gested that maximum retention e⁄ciency of ¢sh is
attained at maximum feed intake and maximum
growth (Einen, Holmefjord, Asgard & Talbot 1995),
or at moderate feed restriction but optimum for main-
taining maximum growth of the ¢sh (Storebakken &
Austreng 1987; Alanara 1994), or at feeding level be-
low that required for maximum growth (Elliot 1976;
Brett & Groves 1979; Kolsater 1995).
There were several models describing the relation-
ship between growth and ration size. Rafail (1968)
suggested the model speci¢c growth rate (SGR) 5
a1b (FL ^ FLm)1/2, where FLm is the optimum
feeding level. Some authors have used the model
SGR 5 a1b In (FL1c) to describe the growth-feeding
relationship (Allen & Wootton1982; Singh & Srivasta-
va 1985; Cui & Wootton 1988). For Negaprion breviros-
tris P. (Cortes & Gruber 1994), the von Bertalan¡y
model SGR 5 a (1^e b(FL c)) was used, where a
can be interpreted to represent the maximum growth
rate and c to represent maintenance ration size.
Although many studies used SGR based on the nat-
ural logarithm of BW to explain the growing perfor-
mance, in view of the in£uence of temperature and
feeding period, a more accurate and useful coe⁄cient
for growth prediction in ration to water temperature
based on the exponent of 1/3 power of BW as ther-
mal-unit growth coe⁄cient (TGC) was recommended
under each feeding level (Iwama & Tautz 1981;
Cho 1990).
The objectives of this experiment were to study the
e¡ects of di¡erent feeding levels on growth, retention
e⁄ciency and energy partitioning of redlip mullet.
Materials and method
Fish and experimental conditions
Redlip mullet caught in Qingdao seacoast, China and
reared at the aquarium of Institute of Oceanology,
Chinese Academy of Sciences were used in this study.
Fish were transferred into 1000 L £ow-through tanks
2 weeks prior to the trial for acclimation. The tanks
were individually aerated and supplied with sand-¢l-
r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 906^911
E¡ect of feeding level on redlip mullet K Bin & W W Xian
tered natural seawater. During the acclimation peri-
od, the ¢sh were fed the experimental diet at a rate of
about 3% of BW daily. The temperature was con-
trolled as 24  1 1C. The illumination in the room in
which the tanks and ¢sh were held was 12 h light/
12 h dark with the light period from 08:00 to 20:00
hours. The salinity was 30 g L 1.
Basal metabolism trial
The relationship between basal metabolism and ¢sh
BW was studied. Before the trial, ¢sh were starved for
48 h and completely empty of diet. Fish weighing
1.88^14.02 g were randomly distributed to eight re-
spirometer tanks full of water (one ¢sh each respi-
rometer tank) and sealed with plastic ¢lm, and each
soaked in a 0.2 m3 glass aquaria to avoid air ex-
change. Two blank groups were kept for comparison.
The respirometer tank was glass tank (25  25 
40 cm, volume 50 L) with outlet (diameter 5 mm) for
water sampling. According to the sizes of ¢sh, trial
lasted for 3^6 h. Oxygen consumption was deter-
mined by the di¡erence of dissolved oxygen before
and after the trial. Dissolved oxygen was measured
using Winkler’s method. The oxycalori¢c coe⁄cient
of 13.54 kJ g O2 1 (Elliot 1975) was used to convert
oxygen consumption (g) into energy (kJ), estimating
the heat production of ¢sh (Hef).
Growth trial
The growth trial was carried out in a £ow-through
system of glass tanks (volume 50 L) with aeration.
Water £ow was 4 L h 1. Salinity was measured every
day and averaged 30  2 g L 1. The tanks were indi-
vidually aerated to keep dissolved oxygen above
5 mg L 1. Temperature was controlled at 24  1 1C
by temperature-controlling equipment. The ¢sh were
starved for 48 h before weighing. Fish weighing 1.03^
4.42 g were randomly distributed to 23 tanks (one
¢sh each tank) and divided into six groups of 5, 3, 3,
3, 3 and 6 ¢sh, corresponding with six feeding levels,
starvation, 1% (FL1), 2% (FL2), 3% (FL3), 4% (FL4) of
¢sh BWand satiation. To avoid mistakes in determin-
ing the actual feed requirement or studies under
poorly controlled experimental conditions (e.g., me-
chanical distribution of feed, variable temperature),
highly controlled conditions (careful hand-feeding
to avoid feed waste, controlled temperature) were
adopted. Fish in FL1, FL2, FL3 and FL4 were fed twice
a day (at 08:00 and 10:00 hours) and ¢sh at satiation
907
E¡ect of feeding level on redlip mullet K Bin & W W Xian
were freely fed. For redlip mullet showed continuous
feeding activity, we judged satiation on the follow two
factors: (1) intensity and frequency of feeding action
weakened; and (2) remains observed 30 min after
feeding. Ration size ranged from 5.76% to 7.27%
(mean 6%) BW was regarded as satiation. The unea-
ten feed in the satiation group was collected 30 min
after feeding. Feed was also placed in water without
¢sh for 30 min and then collected, dried and weighed
to determine the potential loss of uneaten feed by the
di¡erence to correct the amount of actual feed intake.
Faeces were collected twice a day (12:00 and 20:00
hours) and dried at 70 1C for 24 h. Eight ¢sh were
sampled from the original batch for measurement of
initial body composition. The trial lasted 21 days. At
the end of the trial, all ¢sh were weighed individually
after 2 days of starvation, then dried and reground
for carcass chemical analysis.
Fish BWgain and retention e⁄ciency (weight gain:
intake feed) were determined. Growth rate was cal-
culated using the thermal-unit growth coe⁄cient
(Cho 1992) as follows: (FBW1/3 ^ IBW1/3)/S(T  D) 
100 where FBW is the ¢nal BW (g), IBW is initial
BW (g), T is water temperature (1C) and D is number
of days.
Chemical analysis
Diet, faeces, initial carcass and ¢nal carcass samples
were analysed in duplicate. Dry matter was calcu-
lated by weight loss after drying under 70 1C to a con-
stant value. Crude protein was measured according
to the Kjeldahl procedure and multiplying N by 6.25.
Lipid was measured using ether extraction. Ash was
calculated from the weight loss after incineration of
samples for 24 h at 550 1C in a mu¥e furnace.
Energy partitioning
The energy partitioning scheme and notation pro-
posed by the NRC (1981) was used to describe energy
partitioning by the ¢sh. The gross intake energy (IE),
retention energy (RE) or faecal energy (FE) was cal-
culated by converting their chemical composition
into energy using the equivalent of 23.65 kJ g 1 for
protein and 39.50 kJ g 1 for lipid (Elliot 1976). The di-
gestible energy was the di¡erence of IE and FE. Non-
faecal losses (kJ) were calculated from the nitrogen
budget as follows: (IN RN FN)  24.83 where IN
is nitrogen intake (g), RN is retained nitrogen (g) and
FN is faecal nitrogen (g), 24.83 is a constant (kJ g 1).
908
Aquaculture Research, 2005, 36, 906^911
Basal metabolism was calculated from the rela-
tionship between ¢sh weight and HeE concluded
earlier. Dynamic live BW of the ¢sh was calculated
for each day of the experimental period by the TGC
growth model from earlier trial in this study.
Statistical analysis
The e¡ect of feeding level on carcass composition,
weight and energy gains, retention e⁄ciency, faeces
production, growth and energy budget components
were analysed using STATISTICA (A¢¢ & Clark 1999).
Turkey’s honestly signi¢cant di¡erence test (Steel,
Torrie & Dickey 1997) with Po0.05 was used to de-
tect signi¢cant di¡erences among the means.
Results
Basal metabolism
Fish weighing 1.88^14.02 g consumed 10.29^57.48 kJ
¢sh h 1. The relationship between BW (g) and Hef (kJ)
was curvilinear correlation (Fig.1) and could be writ-
ten as: HeE 5 0.1255BW0.8386 (n 5 8, R2 50.9770).
Diet and carcass composition
Diet and ¢sh carcass composition are presented in
Table 1. The results of ¢nal ¢sh carcass composition
seemed to be divided into three groups: (1) starvation
and FL1; (2) FL2 and FL3 and (3) FL4 and satiation,
with signi¢cant di¡erences among the groups but
1.6
1.4
1.2
1
HeE (kJ/g/d)
0.8
0.6
0.4
0.2
0
0 5 10 15
Body weight (g)
Figure 1 The relationship between fast heating produc-
tion (HeE) and body weight (g).
r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 906^911
Aquaculture Research, 2005, 36, 906^911 E¡ect of feeding level on redlip mullet K Bin & W W Xian

Table 1 Composition of diet, initial ¢sh and ¢nal ¢sh in wet at di¡erent feeding levels (from starvation to satiation)

1
Sampling Feeding level Moisture (%) Protein (%) Lipid (%) Ash (%) Gross energy (kJ g )

Diet 2.88  0.08 50.38  0.09 13.29  0.05 11.01  0.14 17.16  0.09
Initial fish 76.34  0.11a 16.07  0.14a 5.48  0.16a 1.42  0.20a 5.58  0.19ab
Final fish Starvation 78.96  0.13b 14.39  0.26b 3.94  0.18b 1.60  0.13a 4.96  0.22a
1 79.14  0.15b 15.01  0.18b 3.50  0.18b 1.44  0.19a 4.93  0.20a
2 76.33  0.16a 16.46  0.23a 5.06  0.21c 1.24  0.14a 5.89  0.22b
3 75.27  0.09a 16.83  0.18a 5.65  0.25c 1.43  0.13a 6.21  0.21bc
4 72.41  0.17c 18.71  0.23c 6.27  0.13d 1.34  0.10a 6.90  0.18cd
Satiation 71.08  0.22c 18.91  0.23c 7.15  0.29e 1.53  0.24a 7.29  0.29d

Means followed by di¡erent letters showed signi¢cant di¡erence (Po0.05).

Table 2 Feed intake (in wet), weight gain (in wet), retention e⁄ciency and thermal-unit growth coe⁄cient of redlip mullet fed
at di¡erent levels

Feeding Weight gain Feed intake Retention efficiency Thermal-unit growth

level (g fish 1) (g fish 1) (weight gain: intake feed) coefficient (TGC) (%)

Starvation 0.54  0.23a 0a – 0.0261  0.0011a

1 0.25  0.13a 0.66  0.19b 37.47  7.25a 0.0078  0.0016b
2 0.25  0.23b 1.08  0.41b 23.02  9.34a 0.0085  0.0021c
3 0.66  0.12c 1.92  0.37c 34.44  6.32b 0.0194  0.0016d
4 0.91  0.25d 2.31  0.45c 39.42  11.03b 0.0278  0.0034e
Satiation 1.50  0.51e 3.68  0.98d 40.80  17.56b 0.0381  0.0049f

Means followed by di¡erent letters showed signi¢cant di¡erence (Po0.05).

no di¡erences between two levels within a group ex- 1.4

cept that ash content remained constant. An excep-
tion was that the lipid content of FL4 was signi¢cant 1.2
lower than that in the satiation group. The protein
Faeces production (g/fish)

content ranged from 14.4% to 18.9%, and was posi- 1

tively related to increasing feeding levels. The lipid
content and gross energy also showed a positive rela- 0.8
tionship to increasing feeding levels. Conversely,
moisture content decreased with increasing feeding 0.6
level. Furthermore, the ¢sh composition of FL2 and
FL3 were the most similar to the initial values. 0.4

0.2
Retention e⁄ciency and growth
0
Retention e⁄ciency, faeces production and growth 0 3 6 9 12
performance are presented in Table 2. Reducing
Feeding level (% body weight)
the amount of feed signi¢cantly lowed weight gain.
There were signi¢cant di¡erences of retention e⁄- Figure 2 The relationship between faeces production
ciency among starvation, FL1 and FL2 groups.When and feeding level (% body weight).
the feeding amount was above 3% of BW, feeding le-
mean value, but the actual value of each ¢sh greatly
vel did not a¡ect retention e⁄ciency. Thermal-unit
£uctuated without a clear relationship (Fig. 2).
growth coe⁄cient was signi¢cantly a¡ected by feed-
ing level, showing a positive relationship with
Energy partitioning
feeding level and reaching a maximum of 0.0381 at
satiation feeding. There was an increased tendency Table 3 lists the energy partitioning of mullet under
of faeces production with increased feeding level in di¡erent feeding levels. The values of energy allocated

r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 906^911 909

E¡ect of feeding level on redlip mullet K Bin & W W Xian Aquaculture Research, 2005, 36, 906^911

Table 3 Energy partitioning of redlip mullet fed at di¡erent levels

Feeding level IE (kJ) DE (%) ME (%) RE (%) HeE (%) NFE (%) FE (%)

Starvation 0a – – 4.57  0.05 3.88  0.04 0.41  0.03 –

(kJ per fish 1) (kJ per fish 1) (kJ per fish 1)
1 11.30  0.21b 88.67  4.08a 76.55  4.23ab 2.45  0.03 58.85  0.19a 12.12  2.97a 11.33  2.35a
(kJ per fish 1)
2 18.46  0.28b 90.85  2.07ab 79.63  2.34b 6.88  1.28a 32.72  .5.41b 11.21  3.21b 9.15  1.88a
3 32.87  0.68c 91.91  1.85ab 82.23  2.49c 14.76  2.65b 22.24  6.38c 9.67  2.87a 8.09  2.18ab
4 39.65  0.88c 94.07  1.29bc 83.58  1.99c 22.35  4.83c 17.68  3.39cd 10.49  4.33a 5.93  1.33b
Satiation 63.08  1.51d 95.78  1.31c 86.10  2.03d 24.71  4.74d 14.03  2.57d 9.69  3.63a 4.22  1.87b

Means followed by di¡erent letters showed signi¢cant di¡erence (Po0.05).

IE, intake energy; ME, metabolism energy; RE, retention energy; HeE, basal metabolism; FE, faecal energy.

to growth were minus in starvation and FL1, then
signi¢cantly increased with feeding amount, and
reached 24.71% of IE at satiation. Basal metabolism
accounted for more than half of the IE in FL1, then
greatly decreased to about one third in FL2, and to
only 14.03% at satiation, with signi¢cant di¡erences
among groups. Faecal energy accounted for 4.22^
11.33% of IE, and was signi¢cantly reduced with in-
creased feeding. No di¡erences were found in non-FE,
showing independence of feeding level.
Discussion
The exponent of the relation between HeE and BW
was 0.8386, which agreed with conclusion that 0.8
can describe the change in metabolic rate with ¢sh
size accurately (Hepher 1988) and that metabolic
BW can be calculated as kg0.8 in practice.
Feeding levels had signi¢cant impact on retention
e⁄ciency, and retention e⁄ciency increased with in-
creasing feeding levels. This result was in agreement
with a hypothesis suggesting that maximum reten-
tion e⁄ciency was attained at maximum feed intake
(Einen et al.1995). However, this result was in contra-
diction to some studies suggested that maximum re-
tention e⁄ciency was obtained only at feeding levels
below that for required for maximum growth (Elliot
1976; Brett & Groves 1979; Storebakken & Austreng
1987). This result was also contrary to results with
Atlantic salmon (Salmo salar L.) (Storebakken & Aus-
treng 1987) which suggested that there were no sig-
ni¢cant di¡erences in retention e⁄ciency of ¢sh
fed at 50%, 75% or 100% of the ration required for
maximum growth and results with rainbow trout
(Alanara 1994) which showed feeding levels had no
a¡ects on retention e⁄ciency.
Increasing carcass lipid content and gross energy
in percent with increasing feeding intake was ob-
served in this study. In contrast to many studies (Stor-
ebakken & Austreng 1987; Shearer 1994), the protein
content also showed an obvious increase. This result
showed a high retention ability of redlip mullet at
high feeding levels. Fed at FL2 or FL3, ¢sh were simi-
lar in composition to initial ¢sh, and the higher feed-
ing amount resulted in higher retentive ability of lipid
than of protein for a signi¢cant di¡erence between
FL4 and satiation groups in lipid content but not in
protein content.
The relationship between faeces production and
feed consumption in ¢sh has been described by a lin-
ear (Van Dyke & Sutton 1977; Cui & Liu 1990) or
power equation (Allen & Wootton 1982; Cui & Woot-
ton 1988). There was also an irregular relationship in
Pseudorasbora parva T. & S. from data covering a wide
range of ration sizes because of the high individual
variability in absorption e⁄ciency at satiation (Cui &
Liu 1990). In this study, individual data of faeces
production was irregular although there was an in-
creased trend with increased feeding level.
Fish with di¡erent habits have di¡erent growth
performance. Diets with graded levels of essential
nutrients e¡ect growth. At 24 1C and over a 3-week
growing period, the value of TGC increased with in-
creasing feeding amount from 0.0085 to 0.0381, low-
er than rainbow trout, lake trout (Salvelinus fontinalis
M.), and Atlantic salmon in other studies (Iwama &
Tautz 1981; Azevedo, Leeson, Cho & Bureau 2004).
The relationship between TGC and feeding level of
redlip mullet was polynomial curve and could be
described as TGC 5 0.0015FL210.0195FL 0.0258
(R2 50.9991).
The basal metabolism of redlip mullet in this
study was 0.1255 kJ g 1 and decreased from 58.85%
to 14.03% of IE with increasing feeding amount.
Feeding at less than 2% BW was not enough to sup-
port energy consumption and resulted in loss of BW.

910 r 2005 Blackwell Publishing Ltd, Aquaculture Research, 36, 906^911

Aquaculture Research, 2005, 36, 906^911
This suggests that a feeding level of 2% BW was the
minimum required for juvenile redlip mullet at 24 1C.
The retained energy accounted for 6.88^24.71% of IE
from FL2 to satiation. The relationship between RE
and ME in percentage was linear, indicating feeding
level independence.
In this study, the measurement of feed intake
should be accurate. Weight gain was also measured
accurately. Although the ¢nal faeces production was
determined individually, the faeces collection meth-
od used could cause some error.
Acknowledgments
This study was supported by funding from National
Natural Science Foundation of China, No.40206001:
Impacts of detritivorous ¢sh on matter cycle in
ecosystem, and the Key Laboratory of Mariculture of
Ministry of Education, Ocean University of China,
No. 200404.
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