essentiality of dietary calcium supplement in redlip mullet liza haematocheila
TRANSCRIPT
Essentiality of dietary calcium supplement in redlip mulletLiza haematocheila
M.A. HOSSAIN & M. FURUICHI Fishery Research Laboratory, Kyushu University, Tsuyazaki, Fukuoka, Japan
Abstract
This study was conducted to determine the essentiality of
dietary calcium supplement to redlip mullet Liza
haematocheila. Juvenile ®sh were fed four puri®ed experi-
mental diets containing 2.0 g kg±1 Ca from calcium lactate
(diet 1), no supplemental Ca (diet 2), and 2.0 g and 25.0 g
kg±1 Ca from tricalcium phosphate (TCP, diets 3 and 4),
respectively. At the end of the 10-week experiment, growth
was signi®cantly lower in ®sh fed diet 2 than ®sh fed all other
diets. This suggests that redlip mullet do not obtain adequate
Ca from sea water. Fish fed diets 3 and 4 showed growth
performances similar to ®sh fed diet 1. However, dietary TCP
negatively a�ected bone mineralization of Zn, Mn, K and Fe.
The Ca, Zn and Fe levels in liver were low in ®sh fed TCP-
supplemented diets. From these ®ndings, it may be concluded
that a dietary Ca supplement is necessary for redlip mullet.
Although this species can use dietary TCP as a Ca source for
growth, an easily digestible Ca (monobasic or dibasic)
supplement to a TCP-rich diet is also essential to maintain
normal mineral levels in tissues.
KEY WORDS:KEY WORDS: bone mineralization, calcium, feed utilization,
growth, redlip mullet, tricalcium phosphate
Received 9 December 1998, accepted 31 May 1999
Correspondence: M. Amzad Hossain, Fishery Research Laboratory,
Kyushu University, Tsuyazaki, Fukuoka 811±3304, Japan.
E-mail: [email protected]
Introduction
Mullets are widely spread throughout tropical and temperate
seas as well as brackish waters and are cultured intensively in
several countries (Nash & Shehadeh 1980). Among the mullet
group, redlip mullet Liza haematocheila (Temminck et
Schlegel) is characterized by a relatively short snout and
small head and di�ers sharply from grey mullet Mugil
cephalus in lacking an adipose lid. Redlip mullet can attain a
total length of 1 m. Adult ®sh are omnivorous and feed on
diatoms, blue-green algae and detritus (Okada 1966). In
recent years, the early development (Yoshimatsu et al. 1992a;
Yoshimatsu 1996), dietary protein (Arakawa et al. 1980;
Yoshimatsu et al. 1992b) and lipid requirements (Yoshi-
matsu et al. 1993) of this species have been reported.
However, there has been little reported investigation of the
mineral requirements of this species, especially the Ca
requirement. A dietary Ca supplement may not be necessary
for some ®sh (Ogino & Takeda 1976, 1978; Shim & Ho 1989)
since ®sh can easily absorb Ca from the surrounding water
(Lovelace & Podoliak 1952; Ichikawa & Oguri 1961; Templ-
eton & Brown 1963; Love 1980; Ichii & Mugiya 1983).
However, waterborne Ca has not satis®ed the requirements
of some species reared in low-Ca water (Arai et al. 1975a,
1975b; Robinson et al. 1984, 1986, 1987). It is generally
accepted that sea water contains su�cient amounts of ionized
Ca that are readily absorbed by marine ®sh and a Ca
supplement to the dietmay not be necessary formarine species.
Sakamoto & Yone (1976) reported that a Ca supplement was
unnecessary in a puri®ed diet for red sea bream Pagrus major.
In previous studies, however, we found that tiger pu�er
Takifugu rubripes could not absorb su�cient Ca from sea
water (Furuichi et al. 1997; Hossain & Furuichi 1998). Ca
from dietary tricalcium phosphate (TCP) was also unavailable
to tiger pu�er (Hossain&Furuichi 1998). Recently, El-Zibdeh
(1996) reported poor growth in redlip mullet when Ca was
excluded from puri®ed diets. The purpose of the present study
was to determine the essentiality of a dietary Ca supplement by
investigating whether redlip mullet can absorb su�cient Ca
from sea water or from dietary TCP for optimum growth and
bone mineralization.
Materials and methods
Experimental diets
Four experimental diets were formulated with puri®ed
ingredients (Table 1). Vitamin-free casein was used as a
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dietary protein source. In the control diet (Diet 1), calcium
lactate was supplied to provide 2.0 g kg±1 Ca. Diet 2 was
without Ca supplement. Diets 3 and 4 included TCP to
contain low (2.0 g kg±1) and high (25.0 g kg±1) levels of
supplemental Ca, respectively. The high level was similar
to that derived from ®sh meal in a ®sh-meal diet.
Ingredients for each diet were mixed thoroughly with an
appropriate amount of distilled water and pelleted by
passing the moistened mixture through a laboratory pellet
mill. Diets were dried at 60 °C for 1 h in an air-drying
oven and stored frozen ()20 °C). Just prior to use, a small
quantity of each diet was thawed, broken mechanically
into smaller particles and kept refrigerated (4 °C) until fed.Proximate and mineral compositions of diets are presented
in Table 2.
Fish and rearing procedures
Juvenile redlip mullet (0.6 g average initial weight) were
produced at the Fishery Research Laboratory, Kyushu
University. Prior to initiating the experiment, ®sh were
acclimatized to the indoor rearing conditions and fed the
control diet (diet 1) for 2 weeks. The ®sh were then weighed
individually, selected and distributed into 150-L rectangular
tanks (30 ®sh per tank, two replicates per treatment), and
reared for 10 weeks. Sand-®ltered sea water (salinity 32±
34 g L±1) was supplied continuously to each tank at a ¯ow rate
of 2±3 L min±1. Supplemental aeration was provided to
maintain dissolved oxygen near saturation. Ca content of the
rearing water was determined to be »400 mg L±1. A controlled
photoperiod of 12 h light/12 h dark was maintained through-
out the experiment. Rearing temperature was within the range
26.5±29.0 °C.Each experimental diet was provided at a rate of 5±7%
(equal among treatments) of the biomass daily. Daily rations
were divided into three equal portions and were fed to ®sh at
0900, 1300 and 1600 h. Fish in each tank were weighed
collectively once per week and rations were adjusted accord-
ingly. After weighing, ®sh in the respective tank were given
a bath of sodium nifurstyrenate (sodium salt of 5-nitro-2-(p-
carboxy styryl)-furan) to prevent bacterial infection from
handling stress (Sugimoto et al. 1976).
Sample collection and analytical methods
At the end of the feeding trial, feed was withheld from ®sh for
16±20 h to allow digestive tracts to completely empty. After
anaesthetizing ®sh in a solution of MS 222 (3-aminobenzoic
acid ethyl ester, 0.1 g L±1), standard length and weight of all
®sh in each tank were recorded. Livers were removed from all
®sh in each tank, weighed, and stored for further analyses.
Remaining viscera were subsequently removed and whole-
body carcasses were washed with distilled water and pre-
served at )20 °C prior to bone collection.
Proximate compositions of diets and liver were analysed as
follows. Lipid content was determined by ether extraction,
protein was measured by Kjeldahl analysis, ash was quan-
ti®ed after combustion at 560 °C for 12 h and moisture
content was determined after heating at 105 °C to constant
weight.
Table 1 Composition of the experimental diets for redlip mullet
Ingredient (g kg)1) Diet
1 2 3 4
Casein 450 450 450 450a-Starch1 100 100 100 100Dextrin 150 150 150 150Pollack liver oil 100 100 100 100Vitamin mix2 30 30 30 30Mineral mix3 50 50 50 50Carboxymethylcellulose 50 50 50 50Ca-lactate 15.4 ö ö öCa3(PO4)2 ö ö 5.2 65.2a-Cellulose 54.6 70 64.8 4.8
1Gelatinized potato starch.2Halver's vitamin mixture (1957)+a-cellulose (Halver1957).3Mineral mixture (g kg)1). Major elements: KCl 76.8; MgSO4á5H2O 81.6;NaH2PO4á2H2O, 685.2; Fe-citrate, 24.0; a-cellulose, 126.9; Minorelements: AlCl3á6H2O, 0.9; ZnSO4á7H2O, 2.64; MnSO4á5H2O, 1.753; CuCl,0.157; KI 0.037; CoCl2á6H2O, 0.013.
Table 2 Proximate and mineral compositions of the experimental
diets for redlip mullet
Diet
1 2 3 4
Proximate composition (g kg)1dry matter)Moisture 153 155 159 132Protein 447 434 411 431Lipid 84 73 83 73Ash 48 49 50 104
Mineral content (mg kg)1dry matter)P 8600 8900 9600 20 000Ca 2200 200 2200 24 500K 2100 2000 2100 1900Mg 310 340 350 350Fe 250 230 260 230Zn 56 53 56 53Mn 21 15 17 18Cu 11 10 13 10
M.A. Hossain & M. Furuichi
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For bone collection, whole carcasses were defrosted at
room temperature, and then steamed over boiling water.
Vertebral bone was separated from whole carcasses, washed
with distilled water to remove muscle and other tissues and
dried in an oven at 110 °C for 2 h. Dried samples for mineral
determination were ground to ®ne particles and stored at
)20 °C.Mineral determinations were performed on vertebrae, liver
and diet samples after wet digestion with a nitric acid±
perchloric acid mixture. All the minerals except P in the
digested sample were determined with an atomic absorption
spectrophotometer (Perkin Elmer 3300, Perkin Elmer Co.,
USA) using ¯ame absorption techniques. P was determined
colorimetrically using the molybdate method (Taussky &
Shorr 1953). Sea water was ®ltered with 0.45 l Millipore
®lter (Millipore Filter Corp., MA, USA), diluted to a suitable
range with deionized water, and Ca was determined by an
atomic absorption spectrophotometer.
Data, when applicable, were analysed by one-way ANOVAANOVA
and Fisher PLSD test in STATVIEWSTATVIEW 4.5 (Abacus Concepts Inc.
1996) for Macintosh to identify signi®cant di�erences
(P < 0.05).
Results and discussion
The results of growth and feed utilization are shown in Table 3
and Fig. 1. Mortality was low and was not related to dietary
treatment. The body weight of ®sh fed diet 2 (without Ca
supplement) gradually decreased from week 7 of the feeding
experiment compared with ®sh fed diet 1. Final body weight
was signi®cantly lower in ®sh fed diet 2 than that of the other
groups.Growth of ®sh fed theTCP-supplemented diets 3 and 4
was comparable to ®sh fed the calcium lactate supplemented
diet 1. Daily feed consumption and feed e�ciency were similar
in all groups. There were no signi®cant di�erences (P > 0.05)
in condition factors among ®sh fed the di�erent diets.
Normally, ®sh can easily absorb Ca from the surrounding
water (Lovelace & Podoliak 1952; Ichikawa & Oguri 1961;
Templeton & Brown 1963; Love 1980; Ichii & Mugiya 1983).
Dietary Ca was not needed when the amount of ionized Ca in
fresh water ranged from 20 to 23 mg Ca L±1 for common
carp Cyprinus carpio and rainbow trout Oncorhynchus mykiss
(Ogino & Takeda 1976, 1978) and also for chum salmon
Oncorhynchus keta held in fresh water (Watanabe et al.
1980). Lovell & Li (1978) reported no de®ciency signs in
®ngerling channel cat®sh Ictalurus punctatus fed puri®ed
diets containing only 0.50 g Ca kg±1 when reared in water
Table 3 Performance of redlip mullet
fed the experimental dietsDiet1
1 2 3 4
Average body weight (g)2
Initial 0.62 þ 0.09 0.62 þ 0.08 0.62 þ 0.08 0.62 þ 0.08Final 7.3 þ 1.8a 6.5 þ 1.8b 7.1 þ 2.1a 7.7 þ 2.2a
Weight gain (%) 1080a 950b 1050a 1140a
Daily feed consumption (% of body wt) 6.1a 6.3a 5.7a 6.1a
Feed efficiency (%) 47.8a 45.9a 48.7a 47.7a
Condition factor2,3 1.51 þ 0.09a 1.48 þ 0.09a 1.50 þ 0.06a 1.48 þ 0.08a
Survival rate (%) 91.7 86.7 93.3 96.7
1Values in the same row with di¡erent superscript letters are signi¢cantly di¡erent (P < 0.05, Fisher PLSDtest).2Means þ SD.3Condition factor = 100 3 (body weight in g) 3 (total length in cm))3.
Figure 1 Weekly growth of redlip mullet fed the experimental diets
for 10 weeks. The scale on the right shows the change in water
temperature (W.T.). Di�erent letters indicate signi®cant di�erence
(P < 0.05).
Ca supplement to redlip mullet diet
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containing 14 mg Ca L±1. Growth and feed utilization of
guppy Poecilia reticulata were found to be independent of
dietary Ca when the rearing water contained 40 mg Ca L±1
(Shim & Ho 1989). Sakamoto & Yone (1976) reported that
the dietary Ca supplement was dispensable for red sea bream
fed a puri®ed diet. A dietary Ca supplement was reported not
to be essential in tilapia Oreochromis mossambica reared in
arti®cial sea water (Boroughs et al. 1957). In the present
study, however, poor growth of redlip mullet fed diet 2
indicates that Ca uptake from sea water, which was the sole
source of available Ca, was not adequate for proper growth.
Similar results were obtained by the authors with another
marine species, the tiger pu�er (Hossain & Furuichi 1998).
This suggests that the ability to absorb Ca in water varies in
®sh species.
Ca and P exist in ®sh meal mainly as hydroxyapatite and/
or TCP which are di�cult to digest by ®sh. P from TCP was
found to be less available to ®sh than that from more soluble
mono- and di-calcium phosphate (Takamatsu et al. 1975;
Ogino et al. 1979; Shitanda et al. 1979). Ca in TCP also is
supposed to be unavailable to some ®sh. Tiger pu�er were
unable to utilize Ca from white-®sh meal (Furuichi et al.
1997) or from diets supplemented with TCP (Hossain &
Furuichi 1998). In the present study, redlip mullet fed the
calcium lactate-supplemented and low or high TCP-supple-
mented diets had similar growth performances, which indi-
cates that the Ca availability from TCP was similar to that of
calcium lactate to redlip mullet. Tiger pu�er were likely to be
unable to utilize Ca from TCP because they lack a true
stomach and gastric secretion (low pH), which decrease the
ability to digest minerals (Furuichi et al. 1997; Hossain &
Furuichi 1998). In contrast, redlip mullet have a well-
developed stomach with a long intestine which facilitates
the absorption of Ca from TCP.
No remarkable di�erence was observed between the lipid
contents of liver among ®sh fed the di�erent diets (Table 4).
Deletion of the Ca supplement to the diet increased Mn and
Cu contents of the liver. Increased Mn and Cu contents were
also found in the liver of tiger pu�er fed a diet without Ca
supplement (Hossain & Furuichi 1998). Liver ash content
was slightly lower in ®sh fed diets 3 and 4 than ®sh fed other
diets. Dietary TCP supplement (diets 3 and 4) decreased the
Ca, Zn, and Fe contents and increased Mn contents of the
liver. Deletion of Ca did not a�ect the hepatosomatic index.
In contrast, dietary TCP increased hepatosomatic index
(Table 4).
A lack of Ca in the diet (diet 2) did not a�ect the Ca, P,
Mg, Fe, Zn, Mn and K contents of bone (Fig. 2). El-Zibdeh
(1996) found no di�erences in Zn and Mn contents in bone
of redlip mullet fed Ca-free and Ca-supplemented diets.
Tiger pu�er also showed no di�erences in Ca, P, Mg, Fe,
Zn and Mn contents of bone when fed diets with or without
Ca supplement (Hossain & Furuichi 1998). Dietary Ca did
not a�ect bone Ca and P contents but did a�ect the growth
of channel cat®sh (Robinson et al. 1986). Bone Ca did not
re¯ect the Ca requirement of blue tilapia Oreochromis
aureus reared in Ca-free water (Robinson et al. 1987).
However, O'Connell & Gatlin (1984) observed that a
dietary Ca supplement increased bone Ca contents in the
same species held in low-Ca water. In the present study,
omission of Ca from the diet increased the Cu contents of
bone. An increase in bone Cu content in the absence of
dietary Ca was also observed in previous studies with redlip
mullet and tiger pu�er (El-Zibdeh et al. 1996; Hossain &
Diet1
1 2 3 4
Ash (g kg)1dry matter) 250a 254a 233ab 221b
Lipid (g kg)1dry matter) 585a 563a 584a 601a
Mineral contents (mg kg)1dry matter)Ca 1320a 1320a 1090b 1100b
P 575a 579a 576a 579a
K 5840a 6130a 5660a 5700a
Mg 450a 450a 410a 430a
Fe 270a 280a 240ab 230b
Zn 41a 41a 32b 33b
Mn 3.5b 4.1a 4.2a 4.3a
Cu 7.7b 8.4a 7.3b 7.5b
Hepatosomatic index2 2.70 þ 0.51bc 2.51 þ 0.51c 3.00 þ 0.52a 2.87 þ 0.52ab
1Values in the same row bearing di¡erent superscripts are signi¢cantly di¡erent (P < 0.05, Fisher PLSDtest).2Hepatosomatic index (mean þ SD) = (liver weight in g/body weight in g) 3 100.
Table 4 Liver ash, lipid and mineral
compositions and hepatosomatic index
value of redlip mullet fed the experi-
mental diets
M.A. Hossain & M. Furuichi
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Furuichi 1998) although the reason for this response is not
clear.
There was little variation in Ca, P and Mg contents of bone
among ®sh fed the control diet (diet 1) or diets with low (diet 3)
or high (diet 4) levels of TCP (Fig. 2). Dietary TCP did not
a�ect the bone Ca, P andMg contents in tiger pu�er (Hossain
& Furuichi 1998). The Fe content in bone of ®sh fed diet 3 was
similar to that of ®sh fed diet 1, but that in ®sh fed diet 4 was
lower than that in ®sh fed diet 1. These ®ndings indicate that
only excess dietary TCP supplement may a�ect bone Fe
content of redlip mullet.
Considerably lower bone Zn contents in ®sh fed diets 3 and
4 compared with ®sh fed diet 1 indicate that dietary TCP
decreases bone mineralization of Zn. A comparable result
was also found in tiger pu�er (Hossain & Furuichi 1998).
Dietary TCP has been reported to be one of the inhibitors of
Zn bioavailability in ®sh (Hardy & Shearer 1985; Satoh et al.
1987a, 1987b; Gatlin & Phillips 1989). Spinelli et al. (1983)
stated that excess Ca and Mg alone may reduce the growth
and decrease the availability of Zn in rainbow trout. Kaushik
(1995) showed that excessive dietary TCP reduced the
absorption of Mn, Zn and P by nearly 50% in common
carp. Conversely, dietary Ca at a level of 20.0 g kg±1 did not
a�ect Zn bioavailability in blue tilapia (McClain & Gatlin
1988). Satoh et al. (1989) also found no e�ect of supplemen-
tal dietary TCP on weight gain and Zn content in vertebrae
of channel cat®sh. Dietary TCP at low and high levels in diets
3 and 4, respectively, decreased the Mn contents in the bone
of redlip mullet. Excessive TCP in diet 4 also decreased the K
content of the bone. The Cu content of bone was higher in
®sh fed the low and high TCP-supplemented diets than in ®sh
fed the control diet.
TCP supplemented to a diet provides Ca to redlip mullet,
which is evident by the ®sh's growth and feed e�ciency.
However, dietary TCP resulted in poor bone mineralization
of Zn, Mn, K and Fe. TCP also decreased Ca and Zn
contents in liver.
From the present experiment, it may be concluded that Ca
uptake from seawater is not su�cient for proper growth of
redlip mullet thus, easily digestible Ca must be supplemented
to the diet. Vertebral bone Ca and P do not appear to be
good indicators of Ca de®ciency. Redlip mullet maintained
bone Ca, providing indequate Ca for other physiological
processes. It appears that Ca absorbed from seawater was
adequate for bone mineralization but not for normal growth.
The amount of TCP added to diet 4 was similar to that in a
diet containing white-®sh meal as a protein source. The
present study suggests the necessity of monobasic or dibasic
Ca supplement to a ®shmeal-based diet for proper growth
and maintenance of normal tissue mineral levels.
Acknowledgements
This paper is contribution no. 236 from the Fishery Research
Laboratory, Kyushu University. The authors express their
sincere thanks to Dr S. Matsui and Dr T. Yoshimatsu for
their suggestions and assistance in this study.
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