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Agri. Review, 36 (2) 2015 : 100-112
AGRICULTURAL RESEARCH COMMUNICATION CENTRE
www.arccjournals.com
Print ISSN:0253-1696 / Online ISSN:0976-0539
Dietary essentiality of trace minerals in aquaculture-A Review
S. Chanda, B.N. Paul*, K. Ghosh1 and S.S. Giri2
Regional Research Centre,
Central Institute of Freshwater Aquaculture, Rahara, Kolkata-700 118, India.
Received: 26-08-2014
Accepted: 12-05-2015
DOI: 10.5958/0976-0741.2015.00012.4
ABSTRACT
An element is considered as dietary essential when its absence or insufficiency in diet causes deficiency syndrome and
supplementation in normal level brings back normal health. An organism can neither grow nor remain healthy without the
element in question. The element should have a direct influence on the organism and involved in the metabolism. Effects of
the essential elements cannot be wholly or partly replaced by any other elements. All forms of aquatic inhabitants require
some inorganic elements in little or trace amounts for their normal growth and metabolism. Trace minerals do not exist only
by themselves but in combination with others. Therefore too much of one element may lead to imbalances in others resulting
in disease or adverse effect in metabolism. Factors such as diet, absorption ability, toxicities and drug-nutrient interactions
may play a role in maintaining a balance of trace minerals in the animal body. In comparison to the farmed animals, the
knowledge on dietary essentiality of minerals in fish is scarce being mainly restricted to iron, copper, manganese, zinc,
iodine, selenium, cobalt and chromium as components of body fluids, co-factors in enzymatic reactions, structural units of
non-enzymatic macromolecules, etc. Investigations in fish are comparatively complicated as both dietary intake and its
uptake from availability in water have to be taken in account for determining the total mineral budgets. The importance of
trace minerals as essential ingredients in diets, although in small quantities, is also evident in fish. Though requirements of
trace minerals have been studied in some species, still research work needs to be intensified for other freshwater fish species.
Key words: Copper, Fish nutrition, Iron, Manganese, Requirement, Selenium, Trace minerals, Zinc.
INTRODUCTION
All terrestrial and aquatic organisms require
inorganic elements or minerals for their normal life processes.
Fishes have the ability to absorb inorganic elements from
diets as well as from their external environment in both
freshwater and marine water. Minerals, which comprise the
ash of biological materials remaining after the organic
substances have been completely burnt or oxidized in the
body of all animals including fish, are known to be essential
for cellular metabolism. Freiden, (1984) described that an
element is considered essential when its deficient intake
produces an impairment of function and when restoration of
physiological levels of the element prevents or relieves the
deficiency. Organisms can neither grow nor complete its life
cycle without those inorganic elements. They should have a
direct influence on the organisms and involved in the
metabolism. Minerals, which are present in fairly large
quantities in biological materials, are those of calcium,
phosphorus, potassium, sodium, magnesium, etc. The
principle minerals present in micro quantities (or trace levels)
are iron, manganese, zinc, copper, cobalt, selenium, chromium
and iodine (Lall, 1979). There are approx fifteen trace
elements considered to be essential in animals. Among these
the physiological role of a deficiency of chromium, cobalt,
copper, fluorine, iodine, iron, manganese, molybdenum,
selenium and zinc is well recognized (Lall, 1989). The main
function of those essential elements in the body is formation
of skeletal structure, colloidal systems maintenance (osmotic
pressure, viscosity and diffusion) and regulation of acid-base
equilibrium. They serve as an important component of
hormones, enzymes and enzyme activators. Calcium and
phosphorus are required for the formation of the skeletal
structures of the body and scales. Sodium, potassium and
chloride along with phosphates and bicarbonates maintain
homeostasis and acid- base balance equilibrium. A fixed
number of specific trace minerals viz., iron, manganese,
copper and zinc are firmly associated with a specific protein
in metalloenzymes, which produce a unique catalytic function.
*Corresponding author: [email protected].
1
Aquaculture Laboratory, Department of Zoology, The University of Burdwan, Golapbagh, Burdwan–713 104.
2
Central Institute of Freshwater Aquaculture Kausalyaganga, Bhubaneswar-751 002, Odisha, India.
Volume 36 No. 2, 2015
Non-metal iodine is necessary for the biosynthesis of thyroid
hormones, which in turn greatly affects development and
metabolism in all vertebrates (Lall, 1979, 1989).
The concentration of minerals in the body of an
aquatic organism depends on the food source, environment,
species and stages of development and physiological status
of the animal. Most of the organisms accumulate and retain
minerals from the environment; however, their incorporation
is highly selective. In marine food chains, a unique transport
of trace minerals has been reported (Bernhard and Andreae,
1984). Body utilization efficiency of dietary mineral elements
depends on the availability of the element from a feed
ingredient or complete diet. Many mineral bioavailability
crisis are being increasingly recognized in humans and animal
nutrition. Several factors influencing bioavailability include
the level and form of the nutrient, particle size and digestibility
of the diet, physiological and pathological conditions of the
fish, waterborne mineral concentration and the species under
consideration. Among these factors, those related to the
chemical state are important because the element may assume
different molecular forms, valence state and ligands when
ingested from different diets (Forbes and Erdman, 1983).
Some biologically important compounds contain minerals as
an inherent part of their structure, e.g., hemoglobin and
vitamin B12. Fish maintains a delicate balance of the body
levels of the trace minerals by integrating the various
parameters of uptake, storage and excretion. Information on
nutritional requirements of freshwater fish for trace elements
is scarce particularly because many are needed in extremely
small amounts and these pose difficulty in analysis. The
soluble and non-absorbable substances which are thereby
formed in the gastrointestinal tract of the animal may either
prevent or aid the uptake, transport and metabolism of the
element. During these processes certain other inorganic
elements compete with the element under consideration for
favorable binding sites. In addition to all these diet-related
mechanisms, several aspects linked to the environment
influence mineral bioavailability (Watnabe et al., 1980).
Although the trace minerals are required by the animal in
very small quantities (usually less than 100 mg kg -1 dry diet),
they are absolutely required for normal growth. If excess
amounts of the elements are ingested and assimilated, toxicity
may develop. Their ability to regulate abnormal
concentrations varies with the species. Some finfish and
crustaceans are capable to excrete high proportion of
excessive metal intake and can consequently regulate the
concentration in the body at relatively normal levels. These
take place for essential elements like iron, copper and zinc
(Bryan, 1976). Sublethal effects of several metals on aquatic
organisms have been well demonstrated by Bryan (1979).
Most of the sublethal toxicity appears to be of a biochemical
origin and causes morphological, physiological (growth,
swimming performance, respiration and reproduction) and
behavioral changes (Bryan, 1976; Albaster and Lloyd, 1980).
The toxicity mechanisms of metal ions include blocking of
essential biological functional groups of enzymes, displacing
the essential metal ion in the biomolecule (enzyme or protein).
The trace minerals play immense role in cellular metabolism
so their requirement in diet plays crucial role in Aquaculture
Nutrition.
Iron: Iron (Fe) has an active part in oxidation and reduction
reactions and electron transport associated with cellular
respiration. It is found in complexes bound to proteins such
as haem, in enzymes such as microsomal cytochromes,
catalase, etc. and in non-haem compounds such as transferrin,
ferritin and flavin iron enzymes. Iron in form of haemoglobin
occurs in erythrocytes while transferrin form is found in
plasma (Watnabe et al., 1997). The haem molecule has ferrous
or ferric ions in the center of a porphyrin ring. Haemoglobin
contains 4 porphyrin groups per molecule. Myoglobin, a
similar type of compound present in the skeletal or heart
muscle, contains one ferrous porphyrin group per molecule
(Lall, 1979). The iron content of fish is very low compared
to that of mammals (Van Dijk et al., 1975). Though limited
information on absorption and metabolism of iron in fish is
available, but the process is generally the same as in other
vertebrates. Although the gill membrane absorbs iron to a
certain extent, the intestinal mucosa is considered to be the
major site (Watnabe et al., 1997). Addition of iron (Ferrous
sulphate) improved the growth of sword tail and platy fish
indicating a nutritional benefit from dissolved iron in water
(Roeder and Roeder, 1966). Among the different forms of
iron, on comparing the effectiveness in preventing anaemia
it was seen that ferrous and ferric chlorides were equally active
(Sakamoto and Yone, 1979). It is reported elsewhere that
addition of ferrous sulphate to trout diet shows significant
increase in oxidation of that diet (Desjardins et al., 1987).
As in other animals, ascorbic acid is involved in the
metabolism of iron and vice-versa in fish (Desjardins, 1985).
A recent study on the effects of dietary iron supplementation
on growth performances, fatty acid metabolism an d
composition has been reported in rainbow trout (Senadheera
et al., 2012). It shows that graded level of feed supplemented
dietary iron can have a positive effect on LC-PUFA (Long
Chain Polyunsaturated Fatty Acid) biosynthesis in fish. It is
also observed that dietary iron level have a positive effect on
n-3 LC-PUFA content and can also modulate the activity of
fatty acid desaturase enzymes (Table 2).
102
AGRICULTURAL REVIEWS
It is reported elsewhere that deficiency of iron
induces anemia in brook trout (Kawatsu, 1972), yellow tail
(Ikeda et al., 1973), red sea bream (Sakamoto and Yone,
1978a) and also carp (Sakamoto and Yone, 1978b) (Table
1). Iron deficiency is also known to cause yellowish white
liver condition in carp (Sakamoto and Yone, 1978b). In
channel catfish poor feed utilization and lowering of plasma
iron level and transferrin saturation is noted (Gatling and
Wilson, 1986). The hatching rate of rainbow trout eggs was
poor when the iron content was low in feed (Desjardines
1985). Dietary iron toxicity signs develop in rainbow trout
fed more than 1,380 mg Fe/kg (Desjardins et al., 1987). The
dietary iron requirement is presented in Table 3.
The major factors influencing iron absorption are
the proportion of organic and inorganic components of the
diet, the amount ingested and the conditions of the digestive
TABLE 1: Deficiency syndrome of trace minerals in some fishes
Trace Minerals
Deficiency syndrome
Fish species
Reference
Iron (Fe)
Anaemic condition
Brook trout
Yellow tail
Red Sea Bream
Carp
Eels
Channel Catfishes
Kawatsu, 1972
Ikeda et al., 1973
Sakamoto and Yone, 1978a
Sakamoto and Yone, 1978b
Nose and Arai, 1979
Gatlin and Wilson, 1986
Rainbow trout
Channel catfish
Desjardins, 1985
Gatlin and Wilson, 1986
Channel catfish
Gatlin and Wilson, 1986
Common Carp
Satoh et al., 1983b
Atlantic salmon
Poppe et al., 1986
Rainbow trout, Carp,
Tilapia
Ishac and Dollar, 1968; Ogino
and Yang, 1980
Rainbow trout and
Carp
Satoh et al., 1983a,b.
Rainbow trout
Knox et al., 1982
Rainbow trout and
Brook trout
Rainbow trout
Rainbow trout
Takeuchi et al, 1981;
Lall, 1989
Ketola, 1979.
Ogino and Yang, 1978
Rainbow trout
Channel catfish
Satoh et al., 1983a
Gatling and Wilson, 1983
Common carp and
Rainbow trout
Brook trout
Salmonids
Rainbow trout
Carp
Channel catfish
Atlantic salmon
Rainbow trout
Catfish
Takeuchi et al., 1981
Common Carp
Hertz et al., 1989
Copper (Cu)
Manganese (Mn)
Zinc (Zn)
Iodine (I)
Selenium (Se)
Cobalt (Co)
Chromium (Cr)
Haemoglobin suppression with
transferrin saturation
Poor hatching rate
Reduced heart cytochrome c
oxidase
Reduced liver Cu-Zn superoxide
dismutase
Reduced growth and cataract
formation
Hitra disease (Coldwater bacterial
disease caused by Vibrio
salmonicida)
Reduced growth
Skeletal abnormalities
Cataracts
Dwarfism
Disturbance in bone metabolism
Low activity of Cu-Zn superoxide
dismutase and Mn superoxide
dismutase in cardiac muscle and
liver
Poor hatchability and low Mn
level of eggs
Cataracts
Reduced growth, high mortality,
eroded fins and skins,
Short body dwarfism
Dwarfism, low appetite, Zn –Ca
levels, and serum Zn
concentrations
Reduced egg production and
hatchability
Thyroid hyperplasia
Growth depression
Growth depression
Muscular dystrophy
Exudative diathesis
Reduces intestinal synthesis of
Vitamin B12
Improper Glucose utilization
Gaylord et al., 1914
Higgs and Eales, 1982
Hilton et al., 1980
Satoh et al., 1983b
Gatlin and Wilson, 1983
Poston et al., 1976
Bell et al., 1985
Limsuwan and Lovell,1981
Volume 36 No. 2, 2015
TABLE 2: Summary of research work done on these trace minerals
Trace minerals
Chemical form
used/Source
Model fish
Importance/Essentiality
Reference
Iron (Fe)
Ferrous sulphate (FeSO4)
Sword and platy
fish
Trout
Rainbow trout
Improved growth
Roeder and Roeder,
1966
Desjardins et al., 1987
Senadheera et al., 2012
Copper (Cu)
Manganese (Mn)
Zinc (Zn)
Selenium (Se)
Ferrous and ferric
chloride (FeCl2, FeCl4)
Copper sulphate
pentahydrate
(CuSO4)
Manganous sulphate
(MnSO4,4H2O) and
manganous chloride
(MnCl2)
Zinc sulphate (ZnSO4),
Zinc nitrate (ZnNO3),
Zinc chloride (ZnCl2),
Selenite, Selenate,
Selenometheonine,
selenocystene
All fish species
Rainbow trout,
carp, channel
catfish, marine
invertebrates
Carp
Atlantic salmon
Cobalt (Co)
Selenite and tocopherol
Rainbow trout
Cobalt chloride and
Cobalt nitrite
Carp
Tilapia
Chromium (Cr)
Chromic oxide, Chromic
chloride, High Cr-yeast,
Chromium nicotinate,
Chromium picolinate
Proper enzymatic activity,
metabolism, proper growth
Channel catfish
(young)
Rainbow trout and
brook trout
Rainbow trout
Salmon fry
Tilapia
Carp
tract. The inorganic iron is reduced to the ferrous state and
freed from the conjugate for absorption. The iron from animal
sources may occur as porphyrin, myoglobin and haemoglobin.
It may be in a complex form with phytin in cereals. The
availability of iron in the various feed stuffs for fish depends
on its form (organic and inorganic) (Watnabe et al., 1997).
Feeds of animal origin other than milk by-products are good
source of iron. Common fish feed ingredients viz., fish meal
and meat meal contain 150 to 800 mg Fe/kg (approx.), cereals
grain ranges from 30 to 60 mg/kg and oil seed protein contain
iron ranges from 100 to 200 mg/kg approximately (Lall,
1989).
Copper: Copper (Cu) is an essential trace elements for all
animals including fish (O’Dell, 1984; Mertz, 1986; Lall,
1989). It plays a fundamental role in the activity of enzymes
Increased diet oxidation
Proper fatty acid metabolism
and biosynthesis of LCPUFA
Anaemia prevention
Good growth, increased
protein synthesis, prevented
fat synthesis in liver
Proper growth
Proper hatchability of eggs
Improve growth and reduce
cataract problem, feed
efficiency
Mortality noted in deficiency
and prevented when
administered
Prevention of Oxidative
cellular injury
Prevention of deficiency
signs
Enhances growth and
haemoglobin production
Higher growth and good
Protein efficiency ratio
Improved growth, energy
retention and liver glycogen
deposition
Improve glucose utilization
Sakamoto and Yone,
1979
Ogino and Yang, 1980;
Murai et al., 1981;
Lanno et al., 1985b,
Watnabe et al., 1997.
Satoh et al., 1987b
Gatlin and Wilson,
1984
Takeuchi et al., 1981,
Lall, 1989
Satoh et al., 1987a and
1987b.
Poston et al., 1976
Bell et al., 1987
Bell et al., 1985
Castell et al., 1986
Anadu et al., 1990
Shiau and Lin, 1993
Hertz et al., 1989
such as cytochrome oxidase, superoxide dismutase, lysyl
oxidase, dopamine hydroxylase and tyrosinase. In addition,
copper-proteins and chelates also have metabolic roles
(Watnabe et al., 1997). Copper metabolism revealed
similarities to mammals in the distribution of copper and
copper dependent enzymes (Syed and Coombs, 1982).
Copper levels are high in eyes (iris and chloroide), liver, brain
and heart. A copper- protein complex ceruloplasmin
exhibiting oxidative activity occurs in blood plasma (Watnabe
et al., 1997). Speruloplasmin appears to be enzymatic in
function though its specific character is unknown. Apart from
the enzymatic function, Cu proteins and chelates are involved
in various pivotal metabolic roles (Table 2). Marine
invertebrates, especially mollusks, possess a blue coloured
Cu containing complex in the haemolymph called the
104
AGRICULTURAL REVIEWS
TABLE 3: Dietary requirement of the trace minerals
Trace minerals
Model fish
Iron (Fe)
Atlantic salmon
Channel catfish
Red sea bream
Eel
Rainbow trout and Carp
Channel catfish
Atlantic salmon
Channel catfish
Rainbow trout and Carp
Atlantic salmon
Juvenile stage fish
Salmon and Trout
broodstock
Rainbow trout and
Common carp
Atlantic salmon
Channel catfish
Blue tilapia
Red drum
Atlantic salmon
Rainbow trout
Channel catfish
Gibel carp
All general fishes
Copper (Cu)
Manganese (Mn)
Zinc (Zn)
Iodine (I)
Selenium (Se)
Cobalt (Co)
Requirement level
(mg/kg diet)
33 – 100
30
150
170
3
5
5
2.4
12 – 13
7.5 – 10.5
2 – 15
>30
Reference
15 – 30
Ogino and Yang, 1978, 1979
37 – 67
20
20
20 – 25
4.5
0.15 – 0.38
0.25
1.18
0.05 – 1.0
Maage and Julshamn, 1993
Gatlin and Wilson, 1983
McClain and Gatlin, 1988
Gatlin et al., 1991
Lall et al., 1985
Hilton et al., 1980
Gatlin and Wilson, 1984
Han et al., 2011
Watnabe et al., 1997
haemocyanin. Copper also serves as an oxygen carrier in
haemolymph of these organisms (Lall, 1989). The
requirement for copper depends on physiological state of
animal, copper content of the water, and the levels of zinc,
iron, calcium and molybdenum, which are metabolic
antagonists of copper. Hence, those elements compete for
the same binding sites on proteins responsible for mineral
absorption and synthesis of metalloenzymes. However, the
details of such mechanisms in fish are little known. Afterwards
it was reported that in rainbow trout for absorption in the
intestinal tract, copper and zinc do not compete for the same
binding site (Knox et al., 1982; Lanno et al., 1985b).
The copper content of tissue declines when the diets
of carp and rainbow trout are devoid of copper (Ogino and
Yang, 1980). However, in the same fish copper deficiency
affected the activities of cytochrome oxidase in heart and
copper zinc superoxide dismutase in liver. The copper
requirement in different fish ranges from 3-5 mg/ kg dry diet
as reported (Watnabe et al., 1997). Higher Cu levels in diet
about 600 mg/kg shows no adverse effect in Rainbow trout.
But it is reported elsewhere that, 730 mg/kg Cu in diet reduced
growth and induces an aversion to food (Lanno et al., 1985b).
Copper toxicity cause damage to gills and necrosis to liver
and kidney (Table 1). Higher copper levels depress growth
and impaired feed conversion in channel catfish (Murai
et al., 1981). A differential effect of ascorbic acid on dietary
Bjornvic and Maage, 1993
Gatlin and Wilson, 1986
Sakamoto and Yone, 1976a, 1978b
Nose and Arai, 1979
Ogino and Yang, 1980
Gatlin and Wilson, 1986
Lall and Hines, 1987; Lorentzen et al., 1997
Gatlin and Wilson, 1984
Ogino and Yang, 1980; Satoh et al., 1987
Maage et al., 2000
Lall, 1985
and waterborne copper has been noted which was unexplained
to some extent. The toxicity and tissue retention of copper
contained in water was affected by dietary ascorbic acid in
carp and rainbow trout (Yamamoto et al., 1977, 1981). Later
it was found that ascorbic acid had little effect on either the
uptake and/or the metabolism of dietary copper in rainbow
trout (Lanno et al., 1985b).
Most feeds and the aquatic medium generally
contain copper in amounts adequate for fish. Plant and animal
protein feed ingredients contain 5-30 mg/kg of copper, whey
products and fish solubles are relatively rich sources of
copper. Moreover during processing of feed copper content
can be increased by using protein enriched sources. In
processed feed ingredients wide variation can occurs in
copper content due to metal contamination (Watnabe et al.,
1997).
Manganese: Manganese is one the important elements
required in fish. It is widely distributed in fish and other animal
tissue as well. Mainly manganese is act as co-factor for the
enzymes peptidase, arginase, succinic decarboxylase and also
activates specific enzymes such as glycosyltransferase and
non-specific enzymes such as kinases, transferases, hydrolases
and decarboxylases. The manganese also acts as an integral
part of metalloenzymes (Table 1). This element has a great
implication in performing oxidative phosphorylation.
Manganese content in mitochondria is higher than cytoplasm.
Volume 36 No. 2, 2015
It is responsible for normal functioning of brain and proper
lipid and carbohydrate metabolism. It has been further
demonstrated that manganese has an active part in the
activation of leucine aminopeptidase (Clark et al., 1987).
Mechanism of manganese uptake from water as well as
gastrointestinal tract was not well documented. Later on
Srivastava and Agrawal (1983) reported the mechanism of
water dissolved manganese uptake, but it is better absorbed
through diet intake. Manganese has very important role in
brood stock nutrition. The manganese content of the diet
influences its level and that of other trace elements in gonads
(Satoh et al., 1987a).
Manganese deficient supply in fish usually results
in retard growth. It has been reported that rainbow trout and
carp show poor growth when a diet with inadequate
manganese is fed (Ogino and Yang, 1980). Satoh et al., (1983
a,b) further reported that due to manganese deficiency,
dwarfism linked disturbances in bone formation and cataract
in eye lens was observed in rainbow trout and carp and again
a reduction in skeletal manganese content has been noted
corresponding to insufficient dietary supply of manganese.
In tilapia deficiency syndrome included reduction
in feed intake, loss of equilibrium, poor growth and increased
mortality rate (Ishac and Dollar, 1968). Fish meal diet devoid
of manganese significantly influences the mineral
composition in common carp gonads. It was reported
elsewhere that eggs of brook trout and rainbow trout
subsequently shows poor hatchability when fed manganese
deficient fish meal diets (Takeuchi et al., 1981 and Lall, 1989).
In carp manganous sulfate and manganous chloride and better
sources. Good growth was recorded in carp provided with
manganous sulfate in the diet and it also increased protein
synthesis and prevented fat synthesis in liver. The requirement
of manganese in fish is 2-20 mg kg / dry diet. It is reported
elsewhere that a supplement of 10 mg/kg manganese required
for proper growth without deficiency symptoms (Satoh
et al., 1987b). It was reported elsewhere that about 2.5 mg/
kg manganese required for proper growth in young channel
catfish (Gatlin and Wilson, 1984). The content of manganese
in fishmeal ranged from 4-38mg/kg, depending on the fish.
Herring meal contains only 4-12 mg/kg. Among the plant
sources, cereals contain 8-50 mg/kg and corn 4-11 mg/kg,
rice bran, wheat middling and corn distillers dried soluble
are good source of manganese (Watnabe et al., 1997). On
the basis of growth, appearance of dwarfism and the
manganese content in vertebrae, it can conclude that
manganese appendage was indispensable in diets formulated
using brown fish meal and sardine meal, as noted for white
fish meal based diets. However, the availability of manganese
in different fish meals to carp was very high and not affected
by tricalcium phosphate present in the diet (Satoh et al., 1989).
Zinc: Zinc is an important trace mineral in fish nutrition as it
plays key role in various metabolic pathways like
prostaglandin metabolism and structural role in
nucleoproteins. Zinc is an integral part of about 20
metalloenzymes such as alkaline phosphatase, alcohol
dehydrogenase and carbonic anhydrase. It also serves as a
specific cofactor of several enzymes and act as a catalyst for
regulating the activity of several Zn-dependent enzymes.
Recent research of zinc-gene interactions has revealed the
basic role of zinc in controlling growth (Chesters, 1991). Zinc
has a structural role in nucleoproteins and involved in
prostaglandin metabolism (Lall, 1979). Though a little data
is documented related to this but it is evident that some clinical
features of zinc deficiency may arise from disturbances in
metabolism of protein and nucleic acid. Intake of zinc in fish
occurs from both feed and water, but dietary supplement is
more efficiently utilized. Gills and gastrointestinal tract are
the main sites of zinc intake in fish (Pentreath, 1973;
Lovegrove and Eddy, 1982). In winter flounder fish entire
digestive tract is capable of absorbing zinc, but the uppermost
position of intestine has the highest capacity and the stomach
has the lowest (Shears and Fletcher, 1983). However Hardy
et al. (1987) reported that in rainbow trout gills play a major
role in excretion of dietary zinc. It is also documented that
generally zinc is excreted through kidney and by chloride cells
of gills (Bryan, 1976). It is reported elsewhere that even when
an optimum dietary zinc is supplemented, uptake of water
dissolved zinc is also occurs (Spry et al., 1988). Deficiency
syndrome of zinc in fish is summarized in Table 1.
Deficiency of zinc results in poor growth, lowered
digestibility of protein and carbohydrate, probably due to
reduced carboxypeptidase activity (Ogino and Yang., 1978).
Zinc deficiency also recorded eye lens cataract and erosion
of fins and skin, and sign of short-body dwarfism (Ogino and
Yang, 1979; Hughes, 1985). In catfish, diets low in zinc
resulted reduced appetite, low growth, low bone zinc and
calcium levels and serum zinc concentration (Gatlin and
Wilson, 1983). The mineral composition of common carp
gonads was significantly affected by zinc depletion from the
mineral supplementation in fishmeal diet (Satoh et al., 1987a).
Fortification of brood stock diets of salmon with zinc
increased the content of this mineral in the ovaries of female
Salmon (Hardy et al., 1984). However it is reported by Knox
et al., (1982) that in rainbow trout amplified levels of zinc in
diet (500 to 1000 mg/kg) cause reduction in haemoglobin,
haematocrit and hepatic copper concentration. In rainbow
trout zinc deficiency impairs immunological response (Kiron
106
AGRICULTURAL REVIEWS
et al., 1993). Deficiency syndrome of zinc is summarized in
Table 1.
up to 60-90 mg I kg-1. The iodine content in plant protein
concentrates is very low (Watnabe et al., 1997).
Zinc availability differs in plant and animal source.
Fish meal contains about 80 – 100 mg/kg of zinc. However
zinc concentration in vegetables proteins ranges from 40 –
80 mg/kg and in cereal grains is 15 – 30 mg/kg. (Watnabe
et al., 1997). However, Eisler (1980) have documented the
zinc sources of various marine invertebrates, vertebrates and
plants from several geographic locations throughout the world.
The requirement of zinc in fish is presented in Table 3.
Selenium: Selenium is an integral component of glutathione
peroxidase (Rostruck et al., 1973). The active level of this
enzyme in liver or plasma is indicative of Selenium supply to
the organism. Selenium along with vitamin E is essential to
prevent nutritional muscular dystrophy. Selenium is
implicated in the metabolism of tocopherol compounds. Both
act as an antioxidant substance in fish. Selenium compounds
are also capable of protecting heavy metal toxicity like
cadmium and mercury (Watnabe et al., 1997). A correlation
between selenium and copper has been observed in rainbow
trout and Atlantic salmon (Hilton, 1989). Selenium acts as
the principal factor in the protective mechanism against
oxidative cellular injury. In the liver the glutathione
peroxidase activity is lowered, while glutathione transferase
activity and pyruvate kinase activity in plasma were increased.
Poston et al., (1976) reported that salmon fry shows mortality
when diet with selenium deficient is supply and again
recovered when administered with selenium. In a study
reported by Bell et al., (1987) on Atlantic salmon it was
highlighted that selenium has a protective mechanism against
oxidative cellular injury. It was reported elsewhere that
maximum glutathione peroxidase activity noticed in the
plasma of rainbow trout when fed with diet fortified by
selenium in the range of 0.15 – 0.38 mg/kg (Hilton, 1980). In
an another report by Bell et al., (1985) it was showed that a
diet with a combination of selenium and tocopherol at 0.9
mg/kg and 41 mg/kg respectively can prevent deficiency signs
in rainbow trout.
Iodine: Iodine metabolism is related to thyroid hormones
which help in the regulation of the level of metabolic activity
in fish. The hormones have wide influence on cellular
oxidation, neuromuscular control, circulatory dynamics,
nutrient metabolism and growth. Triiodothyronine (T3) is the
major hormone of the thyroid gland and also an active
precursor of thyroxine (T4) (Watnabe et al., 1997). Fish differ
from mammals in iodine utilization and in the extrathyroidal
metabolism of T3 and T4 (Higgs et al., 1982). T3 binds more
strongly to plasma proteins than T4, and its turnover rate in
trout is low compared to T4. Excretion of T3 and T4 occurs
mainly through bile, but the gills and kidney also involved
(Watnabe et al., 1997). A report of NRC (1983) noted that
thyroid hyperplasia (goitre) induced by iodine deficiency in
salmonid fish. Agrawal and Mahajan (1981) reported that
iodine uptake by thyroid tissue was reduced in ascorbic acid
deficient catfish. Iodine can be taken up from the surrounding
water via the gills, and the uptake rate is inversely dependent
on the calcium content of the water (Hunn and Fromm, 1966).
The plasma iodine level of freshwater fish ranges from 0.5 to
over 2000 µgdl-1 (Gregory and Eales, 1975). It is dependent
on the iodine content of the diet and water, on the iodine
level of tissue and on the iodine-binding capacity of plasma
proteins. Freshwater fish depend more on a dietary source
for iodine. The iodine from the diet is easily absorbed in the
digestive tract (Gregory and Eales, 1975).
As seawater contains more iodine than freshwater,
iodine deficiency signs appear more in freshwater fish. Age,
physiological state and stress factors considerably influence
the requirement for this mineral. Gaylord et al., (1914) noted
thyroid carcinoma results from the deficiency of iodine in
brook trout.
Marine plants and animals are rich sources of iodine.
Some seaweed contain up to 0.1% iodine. There is a wide
variation in the iodine content of feedstuffs; animal proteins,
excluding fish meals, contain only negligible amounts.
Normal herring meal and capelin meal have only 5-10 mg I
kg-1. On the other hand, Atlantic white fish meal may contain
Selenium deficiency results growth depression, loss
of appetite, lethargy, reduced muscle tone and mortality in
Atlantic salmon (Poston and Combs, 1979). Deficiency of
selenium and vitamin E are the probable factors responsible
for Hitra disease in farmed salmon (Fjoelstad and Heyeraas,
1985; Poppe et al., 1986). Apart from the deficiency
symptoms, it is observed that a reduction in tissue
concentration of vitamin E and selenium, low haematocrit
values and increased haemolytic rates (Watnabe et al., 1997)
and some deficiency syndromes are presented in Table 1.
Fish derive selenium from both diet and water. High
levels of Selenium (40 – 130 µg/l) in water are toxic to fish.
The usual concentration of selenium in water is less than 0.1
µg/l (Watnabe et al., 1997). Han et al., in 2011 reported that
the dietary selenium requirement is 1.18 mg/kg in case of
Gibel carp (Carassius auratus gibelio). Selenium is stored
in various tissues, except liver, in its inorganic form. Selenium
as selenite is effectively taken up through the gills. Selenium
Volume 36 No. 2, 2015
and vitamin E are complement to each other activity, and
protect biological membranes against lipid oxidation.
Fishmeal and marine by-products are the rich
sources of Selenium to fish. Plant materials vary widely in
their Selenium content. Among the source of selenium
selenomethionine was the most digestible (92%), and fishmeal
(47%) the least digestible sources of selenium. The
glutathione peroxidase selenium ration indicated that
selenium supplied as selenite or selenocystine was a better
source for plasma glutathione peroxidase than was selenium
from selenomethionine. Though the comparative availability
of Selenium is poor, still fishmeal-based diets generally
provide sufficient selenium to satisfy the nutritional
requirement of fish (Watnabe et al., 1997).
Cobalt: Cobalt is a component of cyanocobalamin (vitamin
B12), constituting nearly 4.5% of its molecular weight.
(Watnabe et al., 1997). Most animals need the element for
the synthesis of the vitamin B12 by intestinal microflora, and
such bacteria have also been isolated from the intestinal tract
of fish (Kashiwada et al., 1970). Cobalt as part of vitamin
B12 is associated with nitrogen assimilation, and synthesis of
haemoglobin and muscle protein. In addition, cobalt
influences certain enzymes. Cobalt binds to insulin
(Cunningham et al., 1955) and also reduces plasma glucose
levels (Roginski and Mertz, 1977). Dietary cobalt has been
found to be beneficial for haematology and growth of fish;
the erythrocyte count of young carp increased and rainbow
trout grew better (Frolova, 1960 and Sabalina, 1964; Steffens,
1989) and gold-spot mullet exhibited improved survival and
growth (Ghosh, 1975). Cobalt in carp diets resulted in better
growth (Sukhoverkov, 1967) survival of hatchlings (Khan
and Mukhopadhay, 1971) and improved synthesis of protein
(Bhanot and Gopalakrishnan, 1973). The uptake of cobalt
has been demonstrated in rainbow trout eggs at the embryonic
development stage (Kuenze et al., 1978). Hertz et al. (1989)
examined the effects of cobalt on glucose metabolism of carp.
Based on glucose utilization, they suggested an increase in
insulin effectiveness caused by cobalt. They also mentioned
that the mineral may be associated with a protein-sparing
effect based on increased incorporation of labeled leucine
into white muscle proteins, and improved growth and protein
efficiency in cobalt fed fish. Very high doses of cobalt (0.1-5
g Co /kg body weight) were toxic to rainbow trout, resulting
in haemorrhages in the digestive tract and alterations in white
blood cells (Sabalina, 1968; Steffens, 1989). Cobalt
deprivation reduced the intestinal synthesis of vitamin B12 in
catfish (Limsuwan and Lovell, 1981).
Cobalt can be absorbed from the surrounding water
through the gills as well as from the diet. Several studies
with various trout by Phillips and coworkers (Phillips et al.,
1956, 1957, 1958, 1960) examined both modes of cobalt
uptake. The uptake of waterborne cobalt increased with a
rise in temperature and decrease in waterborne calcium. It
was found in brook trout that a large proportion of the mineral
absorbed from the diet was at first retained in the digestive
tract. The initial metabolism after absorption of cobalt occurs
in the pyloric caecum and intestine. The dietary cobalt demand
of fish has been put at 0.05 mg kg-1 diet (Lovell, 1979).
Generally, the cobalt content in fish feeds is in the range of 1
– 6 mg kg-1 diet (Tacon and De Silva, 1983). The mineral is
supplied as cobalt chloride. Cobalt chloride and cobalt nitrate
from feed and cobalt chloride from ambient water improved
growth and enhanced haemoglobin production in carp (Castell
et al., 1986). Cobalt chloride supplementation also resulted
in higher growth and protein efficiency ratio in tilapia (Anadu
et al., 1990). The feed composition has an appreciable effect
on cobalt supply as the demand for the trace element is very
low and can probably be derived from numerous feedstuffs
(Steffens, 1989).
Chromium: Chromium (Cr) shows essentiality for both
animals and humans. It is mostly occurs in oxidative states
viz., Cr (II), Cr (III) and Cr (VI). Cr (III) is required for
proper carbohydrate and lipid metabolism. The ability of Cr
to form co-ordination compounds and chelates, is an
important chemical characteristic that makes this essential
element available to living organisms. In food Cr is available
in inorganic form Cr (III) as a part of biologically active
molecule. Though the biological structure of the active form
of Cr is not fully characterized, it appears to be a
dinicotinatochromium (III) complex, stabilized with
glutathione or its constituent amino acids (Toepfer et al.,
1977). Biologically the function of Cr is closely alike with
that of insulin. Most of the Cr-potentiated reactions are also
insulin dependent as well. In humans Cr potentiates the action
of insulin in vitro and in vivo; maximal in vitro activity
requires a chemical form termed the glucose tolerance factor
and tentatively identified as Cr-nicotinic acid complex (Mertz,
1993). It was reported elsewhere that supplementation of Cr
with carp diets improves the glucose utilization by modulation
of endogenous insulin activity (Hertz et al., 1989). Again
Shiau and Lin, (1993) reported that Tilapia diet containing
glucose with chromic oxide showed improved growth with
better energy retention and liver glycogen deposition. But
Nag and Wilson in 1997 reported that there is no effect in
glucose utilization in Channel catfish supplemented with Cr
containing diets. No adverse effect is noticed in rainbow trout
fed with low Cr containing purified diet as reported by Tacon
and Beveridge (1982).
108
AGRICULTURAL REVIEWS
The dietary toxicity of Cr (III) is not well established,
though report has been made on the toxicological effects of
Cr (VI) in brook trout by Benoit (1976). The commonly used
Cr feed supplements are chromic chloride, high Cr-yeast, Crnicotinate, Cr-picolinate.
CONCLUSION
Trace minerals are essential for fish and is involved
in the normal metabolism and life processes. The trace
minerals requirement in fish are; Iron 30-170, Copper 3-5,
Manganese 2-30, Zinc 15-80, Iodine 4-5, Selenium 0.15 to 8
and Cobalt 0.05 to 1.0 (mg/kg diet). The minerals are required
in extremely small quantity in the diet but excess
supplementation through a continuous process can cause
severe toxicity. In deficient condition, growth is retarded with
abnormal metabolism in particular. It is also a matter of
concern about the potential causes of conditioned trace
element deficiencies such as food processing methods, dietary
interactions, disease conditions and genetic disorders. Fish
can absorb part of the required minerals directly from the
water through gills or even through their entire body surface.
The minerals absorbed from water do not meet the total
requirement and a certain supplementation through the diet
is required whether in the natural food or supplementary feed.
Definitive data on requirements should involve systematic
tests on the metabolic functions of the mineral. Precise
analytical procedure is essential for detection of ultra trace
element at tissue level, when present in minute quantity. There
is also insufficiency activity regarding trace minerals effect
on fatty acid metabolism specially PUFA. It should be taken
care of so that specific diet could be composed with
supplemented trace minerals to meet up the need. These
approaches should highlight future research and potential
strategies to maximize the activity on the requirement of trace
mineral nutrition in fish. Once this requirement is met, the
normal growth, survival and essential cellular metabolism
may be ensured.
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