JOURNAL OF THE Vol. 40, No.

5
WORLD AQUACULTURE SOCIETY October, 2009

Dietary Zinc Requirement of Hybrid Striped Bass, Morone

chrysops × Morone saxatilis, and Bioavailability of Two
Chemically Different Zinc Compounds
J. Alejandro Buentello, Jonathan B. Goff, and Delbert M. Gatlin, III1
Department of Wildlife and Fisheries Sciences, Texas A&M University System, College Station,
Texas 77843-2258, USA

Abstract
Zinc is a trace mineral element that plays an essential role in numerous biochemical processes,
and has been shown to affect growth and health of several fish species. However, the dietary zinc
requirement of hybrid striped bass has not been defined. Therefore, a feeding trial was conducted
to determine the dietary requirement for zinc by this fish and to compare zinc bioavailability of two
chemically different forms (zinc sulfate and zinc proteinate). Six experimental diets were formulated
with purified ingredients and supplemented with ZnSO4 to provide total zinc concentrations of 7,
12, 16, 26, 42, and 80 mg/kg diet which were determined by analysis. Each diet contained 32%
crude protein, 6% lipid, and approximately 14.2 kJ of digestible energy per gram. The experimental
diets were fed twice daily for 10 wk to triplicate groups of 15 hybrid striped bass initially weighing
0.86 ± 0.05 g/fish in 38-L glass aquaria, connected as a recirculating system. Finally, after the feeding
period, the fish were evaluated for weight gain, feed efficiency, protein efficiency ratio, and survival,
as well as blood serum zinc, bone zinc, and hematocrit. All fish thrived during the feeding trial and
not even the fish fed the basal diet displayed any apparent deficiency signs, although weight gain
steadily increased with escalating levels of dietary zinc up to 42 mg/kg diet. On the basis of the most
responsive indicators–bone zinc and serum zinc–the minimum dietary zinc requirement of hybrid
striped bass was determined to be 17.0 and 17.3 mg Zn/kg diet, respectively, based on broken-line
regression. This estimate is similar in magnitude to dietary zinc requirements reported for other fish
species. In addition, the bioavailability of zinc proteinate versus that of ZnSO4 was estimated by
deriving the ratio of the slopes of the regression lines fitted to bone zinc and serum zinc data. This
analysis indicated that hybrid striped bass utilized zinc proteinate ∼1.7 times more efficiently than
ZnSO4 .

The hybrid striped bass, Morone chrysops ♀
× Morone saxatilis ♂, possesses desirable traits
for commercial aquaculture such as aggressive
feeding behavior, rapid growth, and tolerance
to a wide range of environmental conditions
(Myers and Kohler 2000; Wang et al. 2006).
These characteristics along with an advanta-
geous selling price have made the hybrid striped
bass one of the fastest growing segments of the
finfish aquaculture industry in the USA (Harrell
1997).
The minimum dietary zinc requirements have
been established for a number of fish species
including rainbow trout, Onchorhynchus mykiss
(Ogino and Yang 1978), common carp, Cypri-
nus carpio (Ogino and Yang 1979), channel
1 Corresponding author.
© Copyright by the World Aquaculture Society 2009
catfish, Ictalurus punctatus (Gatlin and Wil-
son 1983), blue tilapia, Oreochromis aureus
(McClain and Gatlin 1988), red drum, Sci-
aenops ocellatus (Gatlin et al. 1991), and
Atlantic salmon, Salmo salar (Maage and
Julshamn 1993). Information concerning the
nutrition of hybrid striped bass has increased
in recent years (Gatlin 1997; Webster 2003)
to facilitate aquaculture production; however,
the dietary requirements for several critical
micronutrients including zinc have not been
determined.
Zinc is an essential micronutrient required for
normal growth and metabolic functions by ver-
tebrates and is essential for fish (NRC 1993). It
is well known that severe zinc deficiency results
in reduced weight gain and feed intake; thus,
exacerbating zinc deficiency as well as causing

687
688 ALEJANDRO BUENTELLO ET AL.
general malnutrition in zinc-deprived animals
(Eder et al. 2000), including fish (Gatlin and
Wilson 1983). Through complex interactions
with other minerals, such as magnesium, cop-
per, and selenium, and various metalloenzymes,
zinc also prevents free radical oxidative dam-
age to tissues and participates in the organism’s
immune system (Sahin et al. 2005). This trace
mineral is a cofactor in more than 20 metalloen-
zymes including those involved in lipogenesis
and lipolysis (Eder et al. 2000); therefore, it
significantly affects the metabolism of various
nutrients. As a cofactor, zinc performs catalytic,
structural, and regulatory functions. In addi-
tion, zinc also influences the conformation of
polysomes and the synthesis of nucleic acids
(Rhodes and Klung 1993). It stabilizes biolog-
ical membranes and functions as a free ion.
Because normal zinc levels in freshwater (Spry
et al. 1988; Gatlin and Phillips 1989) and in
seawater (Willis and Sunda 1984) are know to
be suboptimal to meet metabolic requirements
of rapidly growing fish species, the present
experiment was conducted to determine the
minimum dietary zinc requirement of juvenile
hybrid striped bass.
In addition, because of the similarity in phys-
iochemical attributes of zinc and other cations,
there often exist interactions among these min-
erals including competition for absorption by
the gastrointestinal tract. Because the presence
of excessive levels of dietary zinc can disrupt
transport and absorption systems, thus affecting
the nutritional status of other metals includ-
ing iron, copper, and cadmium, it is desirable
to avoid unnecessarily high supplementation of
zinc which also may limit loading of miner-
als in the aquatic environment. Organic forms
of several trace minerals have been shown
to be more bioavailable than inorganic forms
to various monogastric animals (O’Dell 1984;
Ashmead 1992) including fish (Paripatananont
and Lovell 1995, 1997). Organic forms of zinc
include amino acid-chelated zinc, zinc picoli-
nate, and zinc gluconate. Therefore, a second
objective of the present experiment was to esti-
mate the bioavailability of two zinc sources,
namely, zinc sulfate and zinc proteinate.
Materials and Methods
Experimental Diets and Design
The experimental diet formulations are pro-
vided in Table 1. Crude protein level was for-
mulated from casein, gelatin, and crystalline
amino acids to provide 32% of diet to minimize
intrinsic zinc. This protein level was slightly
below the optimum level for hybrid striped
bass (Brown et al. 1992) but previously deter-
mined to support adequate growth performance
(Jaramillo and Gatlin 2004). The protein-to-
energy ratio was estimated to be 44.4 MJ/g
protein, based on physiological fuel values of
the purified ingredients (Webster et al. 1995;
Keembiyehetty and Wilson 1998). Menhaden
oil was used as the sole lipid source. All
other known nutritional requirements of hybrid
striped bass (Gatlin 1997) were met by the
experimental diets. The basal diet was analyti-
cally determined to contain 7 mg/kg of intrinsic
zinc and was supplemented with graded lev-
els (0, 5, 10, 20, 40, or 80 mg/kg diet) of zinc
sulfate heptahydrate (ZnSO4 ·7H2 O, Sigma, St
Louis, MO, USA) resulting in total zinc con-
centrations of 7, 12, 16, 26, 42, and 80 mg/kg
diet which were determined by analysis.
Diets were blended in a V-mixer (twin shell
dry blender, Patterson-Kelly, East Stroudsburg,
PA, USA) and cold-pelleted using a commercial
meat grinder (model A-200; Hobart, Troy, OH,
USA), and then air-dried to approximately 90%
dry matter and stored at –20 C until fed. A
second set of diets in which zinc proteinate
(15.9% Zn; Chelated Minerals Corporation, Salt
Lake City, UT, USA) was supplemented in
place of ZnSO4 to provide 0, 5, or 20 mg Zn/kg
diet and was used to contrast the bioavailability
of zinc from these two chemically distinct
forms.
Fish, Feeding, and Experimental Conditions
Animal care and experimental protocols were
approved by the Texas A&M University Sys-
tem Animal Care and Use Committee. Recip-
rocal cross hybrid striped bass, M. chrysops
♀ × M. saxatilis ♂, was provided by a com-
mercial producer (Keo Fish Farms, Keo, AR,
 DIETARY ZINC REQUIREMENT OF HYBRID STRIPED BASS 689

Table 1. Composition of the experimental diets on a dry-weight basis.

mg Zn/kg diet
Ingredient (g/kg) 0 5 10 20 40 80

Caseina 90.0 90.0 90.0 90.0 90.0 90.0

Gelatina 18.0 18.0 18.0 18.0 18.0 18.0
Amino acid premixb 240.0 240.0 240.0 240.0 240.0 240.0
Dextrina 250.0 250.0 250.0 250.0 250.0 250.0
Menhaden oilc 60.0 60.0 60.0 60.0 60.0 60.0
Cellulosea 252.0 249.5 247.0 242.0 232.0 121.0
Zinc-free mineral premixd 40.0 40.0 40.0 40.0 40.0 40.0
Vitamin premixe 30.0 30.0 30.0 30.0 30.0 30.0
Carboxymethyl cellulosea 20.0 20.0 20.0 20.0 20.0 20.0
Zinc premixf 0.0 2.5 5.0 10.0 20.0 40.0
Proximate compositiong
Crude protein 319.3 332.5 315.8 322.9 310.7 305.7
Crude lipid 73.7 74.4 72.2 70.9 73.1 73.7
Crude ash 26.5 26.3 26.3 27.6 27.0 27.2
Moisture 104.5 90.6 101.5 95.6 86.7 130.2
a US Biochemical Corporation, Cleveland, Ohio, USA.
b Contained the following L-amino acids (mg/kg): arginine HCl, 54,200 mg; histidine, 28,500 mg; isoleucine, 56,200 mg;
leucine, 94,400 mg; lysine, 35,100 mg; methionine, 20,000 mg; cystine, 9200 mg; phenylalanine, 65,900 mg; tyrosine,
62,300 mg; threonine, 57,800 mg; tryptophan, 15,700 mg; valine, 68,700 mg; aspartate, 88,000 mg; proline, 88,000 mg;
glutamate, 88,000 mg; serine, 80,000 mg; and glycine, 88,000 mg.
c Omega Protein Corporation, Inc., Houston, Texas, USA.
d Same as Gatlin et al. (1991).
e Same as Jaramillo and Gatlin (2004).
f Contained (g Zn/kg premix): 2 g/kg from ZnSO · 7H O in cellulose.
4 2
g Analyzed values are means of three replicate samples.

USA), sorted to uniform size, and stocked into
38-L glass aquaria, arranged in a closed recir-
culating system. The water temperature in the
culture chambers was maintained at 26 ± 2 C,
with salinity levels kept at 2 ± 0.5 ppt using
sodium chloride and synthetic sea salts. Zinc
level in the water utilized in the culture sys-
tem was determined to be 9 μg Zn/L by flame
atomic absorption spectrophotometer. Supple-
mental aeration was provided to the aquaria via
airstones attached to a low-pressure blower to
maintain dissolved oxygen near saturation. Flu-
orescent lighting was set to simulate a 12 h
light : 12 h dark cycle. Fish were acclimated
to the system for 2 wk prior to initiation of the
feeding trial and fed the basal diet twice daily
to apparent satiation. Groups of 15 fish with a
mean initial weight of 0.86 ± 0.05 g/fish were
then stocked into each aquarium.
Each diet was then assigned randomly to
three replicate aquaria. Diets were initially fed
at a rate of 6% body weight/d and gradu-
ally decreased to minimize overfeeding while
maintaining a level close to apparent satiation
throughout the trial. Fish were fed twice daily
with approximately half of the ration dispensed
at each feeding. All fish within an aquarium
were quantified and group-weighed weekly to
adjust feed allowances. The feeding trial was
conducted for 10 wk.
Sample Collection and Mineral Analysis
Approximately 16 h after the last feeding, all
fish were collectively weighed and three fish per
tank were randomly selected for blood collec-
tion via venipuncture of the caudal vasculature.
Blood was allowed to clot at room tempera-
ture for 1 h, and then centrifuged for 15 min
at 1000g. Using disposable glass pipettes,
red cell-free serum was transferred into clean
Eppendorf centrifuge tubes and stored frozen
(–80 C) until further analysis. Serum zinc was
690 ALEJANDRO BUENTELLO ET AL.

quantified by flame atomic absorption spec-
trophotometry according to the method of
Parker et al. (1967). The remaining animals
were anesthetized and immediately frozen for
subsequent analysis of bone and scale min-
erals and proximate composition of carcasses
(AOAC 1995). Five carcasses from each group
were thawed, scaled, and boiled in distilled
water for 10 min to obtain intact trunk and
caudal vertebrae. Scales were separated from
the epidermis of fish with a fine-mesh scrub-
bing pad and thoroughly rinsed to remove all
mucus. Both scales and bones were oven-dried,
extracted with anhydrous ethyl ether, and ashed
according to the previously described methods
(Gatlin et al. 1991) before being analyzed for
zinc using atomic absorption spectrophotometer
(AOAC 1995).
Statistical Analyses
Statistical analyses were performed using
SAS software (version 8.2, SAS Institute, Inc.,
Cary, NC, USA) with a significance level
set at P < 0.05. Each measured response was
subjected to one-way analysis of variance as
well as fitted to the SAS NLMixed broken-
line quadratic model with a random com-
ponent for asymptote as described by Rob-
bins et al. (2006). This model resulted in the
greatest fit (R 2 = 0.99) and accurately esti-
mated the minimum dietary zinc requirements
of hybrid striped bass. Separation of means was
conducted by Duncan’s multiple range test.
Regression analysis also was performed on
serum zinc and bone zinc data with the slopes of
the resulting lines used to compare the bioavail-
ability of dietary zinc from ZnSO4 or zinc
proteinate by deriving the ratio of the slopes
(Paripatananont and Lovell 1995; Tan and Mai
2001).
Results
Weight gain of hybrid striped bass fed the
various diets did not differ significantly (P >
0.05) at the end of the 10-wk period; however,
increasing zinc in the diet generally resulted in
dose-dependent increase in weight gain ranging
from 1.183% of initial weight for fish fed the
basal diet to a maximum of 1.528% for fish fed
the diet supplemented with 40 mg Zn/kg diet
(Table 2). Likewise, there were no significant
differences in survival, feed efficiency, or pro-
tein efficiency ratio of hybrid striped bass sub-
jected to various dietary treatments (Table 2).
There also were no significant differences (P >
0.05) in carcass proximate composition (mois-
ture, crude protein, lipid, and ash; data not
shown) of hybrid striped bass fed the various
levels of dietary zinc.
Serum zinc and bone zinc levels were
highly responsive to zinc sulfate supplemen-
tation (Table 3) such that broken-line models

Table 2. Percent weight gain (WG), feed efficiency (FE), protein efficiency ratio (PER), and survival
of juvenile sunshine bass fed diets containing various levels of zinc.a,b

Total zinc (mg Zn/kg diet) WG (%) Survival (%) FE (g gain/g fed) PERc %

7 1,183 82 0.4 1.3

12 1,268 87 0.4 1.3
16 1,329 82 0.5 1.4
26 1,456 82 0.4 1.3
42 1,528 76 0.4 1.4
80 1,413 71 0.4 1.4
ANOVAd
P >F 0.10 0.70 0.13 0.32
Pooled SE 89 7 0.02 0.05
a
Values are means of three replicate groups.
b
Average initial fish weight was 0.86 ± 0.05 g/fish, feeding trial lasted 10 wk.
c PER = (w − w )/D ; where: w , final weight; w , initial weight; and D , dry protein intake.
i o p i o p
d Significance probability associated with the F statistic.
 DIETARY ZINC REQUIREMENT OF HYBRID STRIPED BASS
Table 3. Analyzed bone zinc (BZn), serum zinc (SZn),
and hematocrit (Ht) of juvenile sunshine bass fed diets
containing various levels of zinc.
Total zinc
(mg Zn/kg diet) BZn (mg/kg) SZn (μg/dL) Ht (%)
7 72 1335
12 112 3900
16 156 6450
26 140 7914
42 156 5835
80 195 9575
ANOVA
P >F 0.0001 0.0074
Pooled SE 7.6 1246
were computed. The breakpoint in the regres-
sion line, which was considered to be the min-
imum dietary level for optimum response, was
17.3 and 17.0 mg Zn/kg diet for serum zinc and
bone zinc, respectively.
Figure 1 presents a contrast in the concen-
tration of zinc found in serum and bone of fish
fed graded levels of zinc from the two different
zinc compounds. The ratios of the slopes of the
linear equations were 1.81 (311.46/171.62) and
1.53 (4.6385/3.0363) for serum zinc and bone
zinc, respectively. On the basis of the amount
of zinc incorporated into bone and transported
in plasma, the zinc proteinate had a bioavail-
ability of about 1.7 times greater than that of
zinc sulfate.
Discussion
Diets used in this feeding trial supported sat-
isfactory growth of hybrid striped bass. After
10 wk of feeding, all groups grew relatively
well when compared with growth rates reported
for the same fish in other studies (Griffin et al.
1994; Keembiyehetty and Gatlin 1997; Gaylord
and Gatlin 2000; Twibell et al. 2003; Gaylord
et al. 2005). Data on concentrations of zinc
in bone and serum adequately fitted broken-
line models and thus, were used to estimate
the minimum dietary requirement for this min-
eral. Because throughout the experiment no
overt signs of zinc malnutrition were appar-
ent, other than a dose-dependent reduction in
weight gain (P > 0.05), these results suggest
691
that hybrid striped bass may be less suscep-
tible to dietary zinc deficiency than other fish
species such as channel catfish (Gatlin and Wil-
son 1983; Paripatananont and Lovell 1995), red
drum (Gatlin et al. 1991), Japanese flounder,
48.9
Paralichthys olivaceus (Wei et al. 1999), and
51.8 various salmonids (Ogino and Yang 1978, 1979;
44.8 Ketola 1979), for which overt signs of zinc defi-
40.2 ciency have been reported as early as 3 wk of
40.1 zinc deprivation. As in the present experiment,
47.8 McClain and Gatlin (1988) also reported no sta-
0.6
tistically significant differences in weight gain
5.2 of O. aureus fed diets containing graded levels
of zinc.
In the present experiment, the fact that
weight gain was not significantly different
(P > 0.05) among treatments but, both bone
and serum concentrations were strikingly dif-
ferent (P = 0.001 and 0.007, respectively),
provide a definite link and strong indication
of physiological mobilization of zinc reserves
from body tissues. A more extended feeding
period, however, would likely overwhelm this
adaptive ability resulting in typical zinc defi-
ciency signs (growth and appetite suppression,
dwarfism, hypogonadism, fin and skin erosion,
eye cataracts, and mortality) (NRC 1993).
In the present study, zinc from zinc proteinate
was incorporated into bone and transported by
plasmatic albumin in greater amounts (∼1.7-
fold) than zinc provided by the inorganic sulfate
(Fig. 1). Differences in bioavailability of zinc
from organic and inorganic sources observed
in the present trial are in line with studies on
abalone (Tan and Mai 2001), various fishes
(Paripatananont and Lovell 1995; Apines et al.
2001), and other vertebrates (Ashmead 1992;
Ashmead and Zunino 1992). These studies have
shown that chelation of various trace minerals
to compounds such as amino acids increases
absorption rates in the intestine due to the pre-
vention of insoluble complex formation which
facilitates zinc transport across the intestinal
mucosa (King and Keen 1994). Apines et al.
(2001) reported that a zinc chelate composed
of two amino acids binding the metal was even
more stable and bioavailable to rainbow trout
than chelates having only one amino acid. An
alternative mechanism that affects the enteric
692 ALEJANDRO BUENTELLO ET AL.

A μg/dL)
Serum Zn (μ
10000
9000
y = 311.46x + 1861.2
8000
R2 = 0.917
7000
6000
5000
4000
y = 171.62x + 2066.5
3000
R2 = 0.7732
2000
1000
0
0 5 10 15 20 25
Supplemental Zn (mg/kg diet)

B μg/g)
Bone Zn (μ
200
y = 4.6385x + 82.4
180
R2 = 0.8606
160
140
120
y = 3.0363x + 82.751
100
R2 = 0.7429
80
60
40
20
0
0 5 10 15 20 25
Supplemental Zn (mg/kg diet)

Figure 1. Analyzed levels of (A) serum zinc and (B) bone zinc of fish fed graded levels of supplemental zinc sulfate
(rhomboids) or zinc proteinate (squares).

transport of zinc is that of amino acids, in gen-
eral, and cysteine, in particular (through its thiol
group), which may upregulate gene expression
of metallothionein at the brush border, which
in turn may enhance the carrier-mediated trans-
port of zinc through the intestinal lumen (Kelly
et al. 1996; Wu et al. 2000).
Our results also indicate that a level of
17.0 mg Zn/kg diet is adequate to maintain opti-
mum performance of hybrid striped bass fed
purified diets. This zinc requirement estimate
is similar in magnitude to those reported for
other fish species (Ogino and Yang 1978, 1979;
Gatlin and Wilson 1983; McClain and Gatlin
1988; Gatlin et al. 1991; Maage and Julshamn
1993). However, hybrid striped bass fed practi-
cal diets are likely to have significantly higher
dietary zinc requirements due to the presence
of phytic acid and tricalcium phosphate derived
from practical feed ingredients which can inter-
act with dietary zinc and limit its absorption
(Gatlin and Wilson 1984; Satoh et al. 1987;
 DIETARY ZINC REQUIREMENT OF HYBRID STRIPED BASS
Gatlin and Phillips 1989; Lovell 1989; Porn-
Ngam 1995; Denstadli et al. 2006).
Acknowledgments
This research was supported by the Texas
AgriLife Research. Authors are grateful to Keo
Fish Farms (Keo, Arkansas, USA) for providing
fish for this study. We also thank Omega
Protein Corporation, Inc. for providing fish oil
used in the diets. The assistance of colleagues
at the Texas A&M University Aquacultural
Research and Teaching facility is gratefully
acknowledged.
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