MINERALS

Introduction
M in e r a ls a re in o r g a n ic s u b s ta n c e s th a t h a ve m a n y im p o r ta n t
fu n ction s in the anim al body. T h ey are required to m aintain m any
m etabolic processes and to provide m aterial for m ajor structures (e.g.
bones, teeth, exosk eleton o f crustaceans) o f aquatic animals. They
are also re q u ire d fo r m a in ten a n ce o f osm otic p ressu re, acid -b a se
balance (e.g., the regulation o f blood pH, hemolymph, urine and other
body fluids) and the proper functioning o f m uscles and nerves. Unlike
carbohydrates, fats, and proteins, they do not provide energy but they
may serve as com ponents o f enzymes, vitam ins, and hormones. Fish,
especially m arine fish species, live in an environm ent that contains
m any o f the m inerals that they need for growth and survival.
This section discusses the m acro, m icro, and trace m inerals; their
physiologic functions; and deficiency signs and symptoms. It also gives
a summary o f the mineral functions and mineral requirements o f fishes
and shrimp.

Classification of Minerals

A bout 20 inorganic elem ents are required to m eet the structural and
m etabolic functions o f living organisms. They are grouped into macro,
micro, and trace minerals.
T h e m acrom in erals calcium , phosphorus, m agnesium , potassium ,
sodium and chlorine are required in the diet in relatively larger quantities
than m icrom inerals.
T h e m ic ro m in e ra ls are ch ro m iu m , cop p er, co b a lt, iron , iod in e,
m anganese, m olybdenum , selenium, and zinc. So far, trace minerals
such as alum inum , arsenic, cadmium, lead, mercury, nickel, silicon,
tin, and vanadium have no known function in fish.

General functions of minerals

The biochem ical functions o f minerals in aquatic anim als are sim ilar to
those in land animals, w ith the exception o f osm oregulation. Minerals
m a y s e rv e as c o m p o n e n ts o f h a rd -tis s u e m a tric e s , s o ft tis s u e s ,
m etalloproteins, and as cofactors or activators o f enzymes. The more
s o lu b le m in e ra ls (ca lciu m , p h o sp h o ru s, sod iu m , p o ta ssiu m , and
chlorine) function in osm oregulation and in m aintenance o f acid-base
balan ce and m em brane potentials.
U n lik e v e rte b ra te s , th ere are severa l p h y sio lo g ica l requ irem en ts
u n iq u e to c ru sta cea n s, p a rtic u la rly in the m o ltin g cycle w h ich is
n e c e s s a r y in c r u s ta c e a n g ro w th . A lth o u g h som e m in e r a ls are
te m p o ra rily sto red in the tissu es, such as the h ep a to p a n crea s, a
sign ifican t am ount is lost during m olting or ecdysis. A sum m ary o f
functions o f m inerals is shown in Table 2.14.
58
F ig u r e 2 . 1 5
Calcium and phosphorus
deficiencies. Soft-shelled shrimp
mainly due to calcium and
phosphorus deficiencies has
caused losses in the shrimp
industry.
Source: Lavilla 2001
CHAPTER 2 NUTRITION IN TROPICAL AQUACULTURE
T a b le 2 . 1 4 Summ ary of mineral functions
| M in e ra l F u n c tio n a n d E s s e n tia lity
C a lc iu m S tru c tu ra l c o m p o n e n t o f h a rd tis s u e ; C o -fa c to r fo r e n z y m a tic p ro c e s s e s ; M u s c le
fu n c tio n a n d p ro p e r n e rv e im p u ls e tra n s m is s io n ; O s m o re g u la tio n
P h o s p h o ru s C o m p o n e n t o f h a rd tis s u e s a n d o rg a n ic p h o s p h a te s (e .g ATP, p h o s p h o lip id s ,
c o e n z y m e s , D N A a n d R N A ); B u ffe r fo r th e m a in te n a n c e o f n o rm a l p H o f in tra -
a n d e x tra -c e llu la r flu id s
P o ta s s iu m C a rb o h y d ra te m e ta b o lis m a n d p ro te in s y n th e s is ; O s m o re g u la tio n ; A c id -b a s e
b a la n c e ; P h o s p h o ry la tio n re a c tio n s
M a g n e s iu m M e ta b o lis m o f fa ts , c a rb o h y d ra te s a n d p ro te in s ; C e llu la r re s p ira tio n ; In tra - a n d
e x tr a c e llu la r h o m e o s ta s is ; P h o s p h a te tr a n s fe r a n d th ia m in e p y ro p h o s p h a te
re a c tio n s ; O s m o re g u la tio n
Copper F u n c tio n s in h e m a to p o ie s is a n d in n u m e ro u s c o p p e r - d e p e n d e n t e n z y m e s (e .g .
c y to c h ro m e c o x id a s e , fe rro x id a s e ); C o m p o n e n t o f h e m o c y a n in
C o b a lt S o u rc e fo r m ic ro b ia l s y n th e s is o f B 12 in in te s tin e
Iro n H e m e c o n t a in in g e n z y m e s (e .g . c y to c h r o m e s , o x id a s e s , p e r o x id a s e a n d
c a ta la s e s ); C o fa c to r fo r e n z y m e s ; H e m o g lo b in
Io d in e N e u r o m u s c u la r f u n c tio n s ; In te r m e d ia r y m e ta b o lis m ; S y n th e s is o f th y r o id
h o rm o n e s
M anganese C o fa c to r o f v a rio u s e n z y m e s y n th e s is
S e le n iu m C o m p o n e n t o f g lu ta th io n e p e ro x id a s e
Z in c C o fa c t o r in s e v e r a l e n z y m e s y s te m s ; C o m p o n e n t o f a la r g e n u m b e r o f
m e ta llo e n z y m e s
Source: Halver 1989
Mineral Availability
In general, bioavailability o f m inerals has been found to be positively
correlated with their solubility in water. Highly soluble salts appear to
be beneficial, but their leaching rates from diets have to be measured.
In addition to inorganic sources, organic chelates and com plexes o f
mineral elements are useful means of delivering minerals. Considering
these lim itations, w ater soluble inorganic salts or bioavailable organic
salts are p re fe ra b le for use in mineral premixes.
Macrominerals
Calcium
Calcium is mostly found in the skeleton and scales o f bony fish and in
the exoskeleton o f crustaceans. Fish scales are an im portan t site o f
calcium metabolism and deposition. Aside from its structural function,
calcium is im portant in physiological processes including metabolism ,
nerve and muscle contraction, nerve impulse transmission, maintenance
o f cell integrity, osmoregulation, and activation o f im portant enzymes.
Calcium and phosphorus deficiencies can cause soft-shelling in shrimps
(Figure 2.15).
Fish may totally or partially meet their calcium requirem ent through
absorption of calcium from the water via their gills, fins and oral epithelia.
The gills are the most im portant site o f calcium regulation. Generally,
 CHAPTER 2 Minerals 59
calcium from feed in gredien ts for exam ple, fish meal, m ay m eet the
requirements o f most fish, however, it is a common practice to supplement
the feed with calcium.

Phosphorus
Phosphorus is another major mineral required by fish and is an important
constituent o f nucleic acids and cell membranes. It is directly involved
in the energy-producing cellular reactions and is a component of essential
molecules such as adenosine triphosphate (ATP). Feed is the main source
o f phosphorus for fish because the concentration o f phosphorus in natural
waters is low. Consequently, the need for a dietary source o f phosphorus
is more critical than for calcium because fish must effectively absorb,
store, mobilize, and conserve phosphate in both freshwater and seawater
environm ents.
Most fish species require a dietary inclusion of 0.5% to 1.0% available
phosphorus. Generally, phosphorus requirem ents are not affected by
dietary calcium levels. In controlled experim ents, the growth o f both
common carp and trout have been shown to be positively correlated with
dietary phosphorus levels but not w ith calcium levels. The apparent
phosphorus availability values for several phosphorus sources have been
estim ated. T h ey are: calcium phosphate m onobasic, 46%; calcium
ph osph ate dibasic, 14%; calcium phosphate tribasic, 5.7%; sodium
phosphate monobasic, 70%; and potassium phosphate monobasic, 68%.
Dietary calcium and phosphorus are important in the prevention o f soft-
shelling in shrimps.

Sodium, potassium, and chlorine

Sodium, potassium, and chlorine are the most common inorganic elements
found in fish. They are abundant electrolytes in the body and are essential
for a number of physiological processes. Appropriate levels of these ions
are required for proper functioning of cells and for maintaining nerve function.
Fish readily absorb these elements from the aquatic medium, especially in
m arine waters, and the tissue levels o f these ions are maintained as a
result of osmoregulation. They are abundant in sea water and in common
feedstuffs used in preparation o f fish diets. Thus, dietary supplementation
o f these elements is norm ally not required.

Magnesium
A large proportion o f m agnesium in fish is contained in the skeletal
tissue. Magnesium is an essential cofactor in many metabolic reactions.
These enzymes include the phosphokinases, thiokinases, phosphatases,
pyrophosphatases, and am ino acyl synthetases. It is also needed in
skeletal tissue metabolism, osmoregulation, and in maintaining muscle
tone.
General symptoms of magnesium deficiency include reduced weight gain
and poor feed conversion. In rainbow trout, magnesium deficiency leads
to renal calcinosis and flexibility o f the muscle, partly due to an increase
in extracellular fluid volume. M ost feed ingredients, especially those of
p la n t o rig in , are a b u n d a n t in m a g n e s iu m th u s, m a g n e s iu m
supplem entation in practical diets is generally not required.
60 CHAPTER 2 NUTRITION IN TROPICAL AQUACULTURE

Microminerals

Copper
Copper is an important component o f a number of metalloenzymes that are
involved in a wide variety of metabolic processes. It is associated with
cytochrome c oxidase o f the electron transport chain in cells. It is the
primary oxygen carrier in hemocyanin o f crustaceans and molluscs. Fish
appears to tolerate copper in the diet than dissolved copper from the
water. Concentrations of 0.8-1.0 mg copper per liter in water are toxic
to many fish species. Feed ingredients such as fish solubles, krill meal,
and yeast contain relatively high levels o f copper. Shrimp in general
cannot meet their physiological requirem ent for copper from seawater.
In white shrimp Penaeus vannamei, symptoms of deficiency include poor
growth, reduced copper levels in the carapace, hepatopancreas and
hemolymph, and enlargement o f the heart.

Iron
Dietary iron is essential in fish for blood formation, maintaining normal
hemoglobin content, hematocrit value, and cell size. A minimum dietary
concentration of 150µg per gram diet is required to prevent iron deficiency
resu ltin g in hypochrom ic, m icrocytic anem ia in red sea bream and
common carp. Dietary iron deficiency has not been observed in shrimp.
In contrast, dietary supplem entation o f shrimp feed w ith iron greater
th an 15 0 µg per gram has been found to give poor grow th . S ince
crustacean diets generally contain polyunsaturated fatty acids, excessive
supplementation of ferrous iron in the diet may affect diet stability through
increased lipid oxidation.

Manganese
Manganese is important either as a cofactor that activates metal-enzyme
c o m p le x e s or as an in te g ra l p a rt o f m e ta llo e n z y m e s in p ro te in ,
carbohydrate, and lipid m etabolism . The uptake o f m anganese from
water by fish has been demonstrated but it is more efficiently absorbed
from feed. However, due to the potential inhibitory effects o f phytic acid
on the b io a v a ila b ility o f m anganese, feed su p p lem en tation m ay be
desirable. Dietary deficiencies in fish have resulted in poor growth,
skeletal abnormalities, embryo mortalities, and poor hatching rates.

Selenium
Selenium is an integral component of the enzyme glutathione peroxidase
that protects cells and membranes from deleterious effects o f peroxides.
In conjunction w ith vitam in E, this enzym e functions as a biological
a n tio x id a n t w h ic h p ro te c ts b io lo g ic a l m e m b ra n e s a g a in s t lip id
peroxidation. Selenium imparts a protective effect against the toxicity
o f h e a v y m e ta ls su ch as ca d m iu m and m ercu ry . T h e s e le n iu m
requirement o f fish varies with the polyunsaturated fatty acid and vitamin
E content o f the diet. Both selenium and vitam in E are required to
prevent m uscular dystrophy in Atlantic salmon. In general, practical
diets containing fish meal contains adequate am ount o f selenium and
does not require supplementation.
 CHAPTER 2 Minerals 61
Zinc
Zinc is an integral com ponent o f m etalloenzym es
inclu ding dehydrogenases, aldolases, peptidases,
and p h o sp h atases. A p p ro x im a tely 20 d ifferen t
enzym es has been found to contain zinc. Many
metabolic functions are affected by zinc deficiency.
Fish can accumulate zinc from both water and feed
b u t d ie ta r y z in c is m ore e ffic ie n tly a b so rb ed .
P ra ctica l diets con tain feed stu ffs that are good
sources o f zinc; for example, fish meal. However,
zin c b io a v a ila b ility is g e n e ra lly v e ry low , thus
supplem entation is essential. Feedstuffs that are F ig u re 2 .1 6
Zinc deficiency cataract:
relatively high in phytate may further reduce zinc non-availability of zinc due to high levels of
bioavailability. Cataract is a common symptom of calcium and phosphorus in fish feed has
zinc deficiency in fish (Figure 2.16). caused losses in the fish farming industry.

Trace minerals
Information on the dietary requirements of other microminerals is limited.
A number o f trace elements are required for adequate fish nutrition but
th eir dietary deficien cies have not been reported. C obalt acts as a
component o f vitamin B 12. Sulfur is required for the synthesis of cysteine.
Chromium is important in normal glucose and lipid metabolism.

Mineral Supplementation of Practical Fish Diets

Although the dietary m ineral requirem ents o f fish have not been well
established, practical diets are usually supplem ented w ith a m ineral
prem ix. Practical diets m ay norm ally contain substantial amounts of
en d ogen ou s m in erals, thus a com p lete m ineral prem ix m ay not be
necessary. E xcessive m ineral supplem entation increases the cost o f
feed and may reduce the bioavailability o f other minerals and increase
phosphorus pollution. Many commercial feed binders may also contain
high levels o f calcium and m agnesium that m ay reach undesirable
levels o f these minerals in the feed.
M o s t s tu d ie s on th e m in e r a l r e q u ir e m e n ts o f fis h h a ve b e e n
c o n d u c te d u s in g s e m ip u r ifie d d ie ts . D ie ta ry re q u ire m e n ts are
determ ined by feeding graded levels o f the elem ent being studied and
m easu rin g the p h ysiologica l respon se o f the test anim al. M inerals
req u ired by fish are calciu m , m agn esiu m , p h osph oru s, and m icro
elem ents such as copper, iodine, iron, m anganese, selenium and zinc.
In general, the dietary requirem ents for m inerals are poorly understood
b ecau se o f the d ifficu lty in d evisin g m in eral-d eficien t diets and the
need to deplete tissue m ineral stores.

Mineral Requirements of Fish

Although inorganic elements are required for the normal life processes
o f fish, know ledge o f their m ineral n u trition is still one o f the least
known areas o f fish nutrition. Unlike other nutrients, significant amounts
o f minerals can be absorbed from their external environment thus it is
62 CHAPTER 2 NUTRITION IN TROPICAL AQUACULTURE

d ifficu lt to control the dietary intake o f the m ineral bein g studied.

Calcium, sodium, potassium, iron, zinc, copper and selenium are generally
derived by fish from the rearing water. The exchange of ions from the
su rrou n din g w ater across the gills and skin o f fish com plicates the
m easurem ents o f mineral requirements. Interaction between minerals
further complicates the assessment of dietary requirements. So far, there
is very limited information on the dietary requirements for minerals by
fish.
A summary o f the known dietary mineral requirements o f various fish
species is presented in Table 2.15 and their deficiency sym ptom s are
summarized in Table 2.16.

Table 2.15 Summary of the mineral requirements of various fish and shrimp species
Channel C om m on Japanese N ile R a in b o w K u ru m a T ig e r
c a tfis h c a rp eel tila p ia tro u t s h rim p S h rim p

Ca 0 .2 7 % 1 .0 -2 .0 % 1 .0 %

P 0 .3 3 -0 .4 5 % 0 .6 - 0 .7 % 0 .5 8 % 0 .8 -1 .0 % 0 .7 -0 .8 % 1 .0 -2 .0 % 1 .0 %

Mg 0 .0 4 % 0 .0 4 -0 .0 5 % 0 .0 4 % 0 .0 5 -0 .0 7 % 0 .3 0 %

Cu 3 µ g /g 3 - 4 µ g /g 3 µ g /g

Fe 1 5 0 µ g /g 1 7 0 µ g /g

Mn 13 µ g /g 1 3 µ g /g 1 2 µ g /g

Zn 2 0 µ g /g 1 5 -3 0 µ g /g 10 µ g /g 1 5 -3 0 µ g /g

Source: Watanabe et al. 1988; Bautista and Baticados 1988

Table 2.16 Mineral deficiency symptoms in fish and shrimp

M in e ra ls | D e fic ie n c y s y m p to m s |

Ca P o o r g ro w th in c h a n n e l c a tfis h ; S o ft-s h e ll s y n d ro m e in c ru s ta c e a n s

P P o o r g ro w th & s k e le ta l a b n o r m a lity in c o m m o n c a rp & ra in b o w tro u t; L o w fe e d e ffic ie n c y a n d h ig h lip id

c o n te n t in c o m m o n c a rp ; L o w a s h in w h o le b o d y a n d v e rte b ra te in c o m m o n c a rp , ra in b o w tro u t a n d c h a n n e l
c a tfis h

Mg P o o r g ro w th a n d h ig h m o rta lity in c o m m o n c a rp a n d ra in b o w tro u t; S lu g g is h n e s s a n d c o n v u ls io n in c o m m o n

c a rp , ra in b o w tro u t a n d c h a n n e l c a tfis h ; H ig h C a c o n te n t in b o n e o f c a rp a n d ra in b o w tro u t; A n o re x ia in
c h a n n e l c a tfis h ; S k e le ta l a b n o rm a litie s a n d re n a l c a lc in o s is in ra in b o w tro u t

Cu P o o r g ro w th in c o m m o n c a rp a n d ra in b o w tro u t; D w a rfis m in J a p a n e s e e e l

Co P o o r g ro w th in c o m m o n c a rp

Fe A n e m ia in c o m m o n c a rp

I D w a rfis m in J a p a n e s e e e l

Al D w a rfis m in J a p a n e s e e e l

Zn P o o r g ro w th , h ig h m o rta lity , e ro s io n o f fin s a n d s k in , lo w Z n a n d M n c o n te n t in b o n e o f c o m m o n c a rp a n d

ra in b o w tro u t; D w a rfis m in ra in b o w tro u t a n d J a p a n e s e e e l; C a ta ra c t in ra in b o w tro u t; L o w C a , M g a n d P a n d
h ig h m o r ta lity in c o m m o n c a rp ; L o w Z n a n d M n in c o m m o n c a rp a n d ra in b o w tro u t

Source: Watanabe et al. 1988

 CHAPTER 2 Minerals 63
Guide Questions

1. W hat are m inerals?

2. W hat are the 3 groups of minerals? Give examples of each group.
3. W hat are the general functions o f minerals?
4. W hat mineral is needed for normal formation of bones, scales and
teeth in fish?
5. W h a t e le m e n t is a com pon en t of th y ro id h o rm o n e s ? of
m etalloenzym es?
6. W hat mineral is a component of vitamin B 12?
7. W hat mineral protects biological membranes against lipid oxidation?
8. W hat minerals are im portant to prevent soft-shelling in shrimps?
9. W hy is the need for a dietary source o f phosphorus more critical
than for calcium?
10. W hat mineral is a component of glutathione peroxidase?
11. W hat minerals are needed for blood formation in fish? hemocyanin
form ation in m ollusks and crustaceans?
12. W hat is the consequence of iron deficiency in fish?
13. W hy is mineral nutrition o f fish one o f the least known areas of fish
nutrition?

Summary

Proteins and amino acids

Am ino acids are the basic unit o f proteins and have a general structure
with an amino group (-NH2) and a carboxyl group (-COOH) bonded to the
alpha-carbon atom. The nature o f the side chain, referred to as the R groups,
are the b asic d ifferen ces am ong am ino acids. The am ino acids are
divided into two groups: the dispensable or non-essential amino acids
and the indispensable or essential am ino acids. O f the 20 naturally
occurring amino acids, ten are essential because they cannot be made
by the fish or cannot be made in amounts that satisfy the requirement.
Th ese are: arginine, histidine, isoleucine, leucine, lysine, methionine,
phenylalanine, threonine, tryptophan, and valine. The non-essential
am ino acids can be m ade by the anim al from carbon and nitrogen
precursor compounds. Cysteine (which can replace part of methionine)
and tyrosine (which can replace part o f phenylalanine), glycine, glutamic
acid, glutamine, aspartic acid, asparagine, alanine, proline, and serine
are non-essential amino acids.
Proteins are polym ers o f am ino acids join ed by peptide bonds. The
structure o f proteins is complex, with no obvious regular structure. To
facilitate structure determination, it is customary to define four levels of
organization. Prim ary structure is the order in which the amino acids
are covalen tly linked together. Secondary structure is the hydrogen
bonded arrangement of the polypeptide chain. Tertiary structure includes
the three dim en sion al arrangem ent o f all the atom s in the protein.