Effect of Dietary Carbohydrate Levels and Source on the Growth, Conversion Efficiency and Carcas?

Composition of the Indian Major Carp, Labeo rohita (Ham.)

DISSERTATION SUBMJTTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS

FOR THE AWARD OF THE DEGREE OF

iHas^ter of ^{lilos^opljp IN

ZOObOGY

BY

BRFANUbbAH

FISH NUTRIT ION RSSEARCH LABORATORY DEPARTM E N T OF ZOOLOGY

ALIGARH M U S L I M UNIVERSITY ALIGARH ( INDIA)

1990

DS1740

D E P A R T M E N T O F Z O O L O G Y AUGARH MUSLIM UNIVERSITY

AUG ARM—202002, UP. (INDIA)

Sections : 1. ENTOMOLOGY 2 PARASITOLOGY 3 ICHTHYOLOGY & FISHERIES 4 AGRICULTURAL NEMATOLOGY 5. GENETICS

Ref. No.

Date

_, rUniversity: 285 ' ^ ' ' ° " " [public : 25646

6.12.1990

This i s t o c e r t i f y t ha t the d i s s e r t a t i o n

e n t i t l e d "Effect of d i e t a r y carbohydrate l e v e l s and sources

on the growth, conversion ef f ic iency and carcass composition

of the Indian major carp, Labeip rohi ta (Ham.)" has been

completed under my supervision by Mr, Erfanullah. This

work i s o r ig ina l and has been independently persued by the

candidate,

I permit the candidate t o submit the d i s se r t a t i on

in p a r t i a l fulfilment for the award of the degree of

Master of Philosophy in Zoology of the Aligarh Muslim

Universi ty, Aligarh.

A.K, J a f r i M.Sc , Ph.D., F.N.A.Sc. Professor,

A CKNOWI^DCEMENTS

I deem i t my p r iv i l ege in expressing my hea r t f e l t

indebtedness t o my supervisor. Professor A ,K, J a f r i , for

his most valuable guidance and unst in ted support during

the course of the present study and in preparat ion of

t h i s d i s s e r t a t i o n .

I am profoundly beholden to the Chairman

Department of Zoology, Prof. M.A. Khan, for encouragement

and providing the necessary laboratory f a c i l i t i e s .

I owe a great deal to Prof. John B. Halver, School

of F i sher ies , Washington, Universi ty of Washington, Sea t t l e ,

for useful suggestions and construct ive c r i t i c i s m .

I s incere ly accord spec ia l thanks to my senior

laboratory col leagues , messers, Mukhtar A, Khan, Mohd.

Farooq Anwar, and Mohd, Abul Hassan for t h e i r generous

cooperation and help.

The needful services rendered by Mr, Naseeruddin

i s deeply acknowledged.

Thanks are a lso due to Far Eastern Regional

Research o f f i ce . United S t a t e s Department of Agricul ture ,

for funding the programme.

CONTENTS

Page No.

ACKNOWLEDGEMENTS

GENERAL INTRODUCTION 1 - 8

GENERAL METHODOLOGY 9 -16

CHAPTER I 17 -24

Effect of va ry ing d i e t a r y

ca rbohydra te l e v e l s on t h e

growth and convers ion e f f i c i e n c y

of the Ind ian major carp ,Labeg r o h i t a (Ham.)

CHAPTER I I . . . . . . . . 25 -32

Growth, convers ion e f f i c i e n c y ,

and c a r c a s s composi t ion of Ind i an

major c a r p , jL.abeo r o h i t a (Ham.) fed

d i f f e r e n t ca rbohydra t e sou rce s .

References . . . . . . , . 33 -44

GENERAL INTRODUCTION

Bxperlmental studies in f ish nutrition are

generally undertaken to achieve three important objec­

t i v e s . First ly , to work out the gross nutrient require­

ments of the species incorporating ndnimiun level of a

particular nutrient in the t e s t d i e t s , leading to noaxiroum

growth, a phenomenon which i s of prime concern to a

cu l tur i s t . Secondly, to translate the information

obtained on the various nutrients required for optimum

growth into successful a r t i f i c i a l dietary formulations,

using both the plant and animal origin feedstuffs. Thir­

dly, to reduce the operating expenditure involved in the

culture system, particularly on feeding practices. The

success of commercially viable aquaculture system, the­

refore, depends, to a great extent, on a sound knowledge

of the nutrition of the concerned culturable f ish

species .

Fish raised in a re lat ive ly small confinement at

a high stocking density must receive the required nutrients

and energy in the form of protein and amino-acids, fat

and carbohydrates. In addition, the non-energy nutrients

such as minerals and vitamins are also adequately required

for optimum growth (Halver, 1976),

- 2 -

Over tb8 l a s t one decade considerable progress

has been made in fish n u t r i t i o n and data are avai lable on

the quan t i t a t i ve requirements of various n u t r i e n t s . How­

ever, compared t o s tud ies on prote ins and amino-acids

(Wilson, 1989), fa t and f a t t y ac ids ( S a r ^ n t gt^ fll.,

1989), vitamins (Halver, 1989), minerals (LaU, 1989)

and energy (Smith, 1989^), very l i t t l e work has been

done on the requirement and u t i l i z a t i o n of carbohydrate

in fish^

Carbohydrates a re considered a cheap source

of energy in animal production. Pishes exhib i t varying

a b i l i t y t o adapt and u t i l i z e carbohydrates t o obtain

energy (Ph i l l i p s e t a l , , I966, Buhler and H a l ^ r , 1961) p

s t o r i n g energy reserves as glycogen in l i v e r and muscle

(WBndt, 1964), and converting i t t o fat in the body

(NAS-NRc, 1983), Inclusion of carbohydrate improves the

binding capaci ty of compounded f i sh d i e t , besides con­

s iderably reducing i t s cos t .

The information so far accumulated on the carbo­

hydrate u t i l i z a t i o n i n many f ish groups i s considerably

var ied , and as such no species spec i f i c carbohydrate

requirements has been es tabl i shed , owing to the fact

tha t u t i l i z a t i o n and a v a i l a b i l i t y of t h i s nu t r i en t id

affected not only by i t s l eve l , source and complexity,

but a l so by inherent d i f ferences r e l a t i n g to the enzymatic

- 3 -

p a t t e m s found in d i f fe ren t groups of f ishes (NAS~NRC,

1983; Jauncey, 1982; Millilcin, 1982; New, 1986; and

Cowey, 1988) • Moreover, carbohydrates are merely used

to provide energy, and do not supply any e s s e n t i a l

n u t r i e n t s tha t cannot be obtained or synthesized from

other d ie ta ry n u t r i e n t s (prote in and fat ) i n the d i e t

(Piper fit a l . , 1989).

However, t he r e i s enough evidence to show tha t

both coldwater as wen as warmwater f ishas are able to

u t i l i z e varying l eve l s of carbohydrates. Fishes l ike

salmonids and ye l lowta i l a re known to be i n f e r i o r i n

cart>ohydrate u t i l i z a t i o n (Edwards a t Si* / 1977;

Austreng fit ai» * 1977; Hilton and Atkinson, 1982;

Shrimeno S^ §!•» 1979; and Furuichi and Yone, 1980),

although the former a r e able to derive some of the

energy for growth purposes and show b e t t e r nitrogen

r e t en t i on /p ro t e in u t i l i z a t i o n in the presence of

carbohydrates (Buhier and Halver, I96I; Bergot, 1979;

and Bergot and Berque, 1983), Channel ca t f i sh and

European e e l s u t i l i z e carbohydrate as an energy source,

as e f fec t ive ly as o ther energy yie lding components of

the f i sh d ie t (Oarling and Wilson, 1977; Robinson and

Lovell , 1984; Oegani and Levanon, 1987), without any

de le te r ious e f f e c t s . Omnivorous t i l a p i a and carps are

reported to t o l e r a t e elevated l eve l s of d ie ta ry carbo-

hydrates (Anderson e t a l , , 1984; sblmeno g t aj^., 1980,

19817 Takeuchi e^ g^.,, 19797 Puruichi and Yone, 1980;

Z i e t l e r fit a^.., 1984; Viola and A r i e l i , 1983; Ufodike

and Matty, 1983; and Jhingran and Pa l l i n , 1988). Among

the carp dispar i ty^ however, e x i s t s on the extent of

carbohydrate u t i l i za t ion . Jau i icey (1982), and Puruichi

e% a l . (1987), have shohrn tha t carp are able to grow

well on d i e t s containing low l eve l s of carbohydrates.

In mullet and cod increased energy consumption did not

a l t e r f ish growth (Alexis and Papapareskevs - Papout-

soglu, 1936 and Hemre et a l» . 1989)«

Similar ly , u t i l i z a t i o n of d i f fe ren t carbohydrate

sources a re a l so considerably varied in d i f ferent f ish

spec ie s . Salmonids are unable to u t i l i z e complex

carbohydrates such as s ta rch and dextr in as compared to

simple sugars (Buhler and Halver, 1961; Bergot, 1979;

and Pieper and Pfeffer, 1980). Whereas, channel ca t f i sh

do not make use of glucose, maltose or sucrose as energy

sources, although complex carbohydrates, such as dext r in

and com s ta rch , are exce l l en t ly u t i l i z e d by the f ish

(Dupree, 1956; and Wilson and Poe, 1987). Puruichi and

Yone (1982) have noted tha t Che a b i l i t i e s to u t i l i z e

glucose and dex t r in i n red sea bream d i e t s were i n f e r i o r

to tha t of oC-starch. The growth r a t e and feed e f f i ­

ciency of carp were highest when fed the ^ - s t a r c h d i e t .

- 5 -

followed by dex t r in and glucose d i e t s (Furuichi and

Yone, 1982). shiraeno et a l , (1979) have shown tha t firuc-

tose i s not used so e f f i c i e n t l y by yel lowtai l as compared

to s t a rch , Ti lapia ffed glucose containing d ie t tend to

grow slower than those fed dex t r in and starch d i e t s (Tung

and Shiau, 1989). Sturgeon fad e i t h e r maltose or glucose

d i e t s had the highest percent energy re ta ined , followed

by those fed e i t h e r dex t r in , raw com s tarch or sucrose

d i e t s , whereas those fed e i t h e r l a c t o s e , fructose or

ce l lu lose d i e t s had the lowest leve l of energy re ten t ion

(Hung fit, a i , . , 1989).

I t has been shown by several woirkers tha t techno­

log ica l treatment of complex carbohydrate sources can

considerably improve carbohydrate u t i l i z a t i o n in terms

of b e t t e r protein and energy re ten t ion in rainbow t rout

(Bergot, 1979; and Bergot and Berque, 1983), In sturgeon,

the u t i l i z a t i o n of raw s ta rch was observed to be very poor

compared t o tha t of the pre- t rea ted s tarch and com

(Kaushik ^ aO,, 1989). Spanhof and Kunhe (1977) found

a lower d i g e s t i b i l i t y values for na tura l s ta rch of

potato and grain in e e l , but when s ta rch was t rea ted

hydrothermally and ge la t in ized , the d i g e s t i b i l i t y of

carbohydrate increased considerably, tfewever^ in the

same f i sh spec ies , o ther complex carbohydrates l ike

ce l lu lose and soybean meal were poorly digested even

- 6 -

a f t e r heat treatment (Schiirdtz et a_l. / 1984). Degaiii

fit flL. (1986) have shown tha t eels grew well when fed

d i e t s containing equal amounts of bread and wheat meal

followed by soluble s t a r ch , sorghum and potato s ta rch .

Inclusion of adequate l eve l s of carbohydrate in

the d i e t has been reported to spare the protein component

for prote in purposes. The fact has been establ ished in

salmonids (Delong s i aL'* 1958; Buhler and Halver 1961;

Ringrose, 1971; Lee and Putnam, 1973; RLeper and

Pfefffer, 1979; and Bergot 1979), plaice (Cowey ^ t al* /

1975), turbot (Adron fit a l . / 1976), sea bass (All iot

£ i SLL» * 1978/ and Tandler and Beamish, 1980) i, channel

ca t f i sh (Tiemeier fit, fli.,, 1965; Page and Andrews, 1973;

Garling and Wilson, 1977; Likiraani and Wtlson, 1982; and

Strange, 1984), Buropean ee l (Degani, 1987; and Degani

and Viola, 1987), t i l a p i a (jauncey and Ross, 1982;

Anderson gt a^, , 1984; and Cisneros fit ^ , , 1984) and

carp (Chiou and Ogino, 1975; Ogino et. aj^,, 1976; and

Shimeno fit ^ , , 1981).

Information on the carbohydrate metabolism and

energy k ine t i c s in f ish i s a lso l imi ted . Although

necessary enzymes for digest ion and u t i l i z a t i o n of

carbohydrates occur i n f i sh , the ro l e of d i e t a ry carbo­

hydrate and energy contr ibut ion of glucose to the energy

requirements of f i sh remain unclear (Piper sk £i» # 1989).

- 7 -

Several studies have, however, indicatecl t h a t hormonal

and metabolic regula t ions of carbohydrate and energy

metabolism vary among the various f i sh species (Tarr,

1972; Shimeno, 1978; Cowey and Sargent, 1972,1979;Wilson

and fbe, 1987; Hung fit a l . . , 1989; Cowey, 1988; Likimani

and Wilson, 1982; and Hilton £ t ^ , , 1987).

Indian major carps , the prime cu l t i vab le fresh­

water f i shes , are ra ised in India on simple supplementary

d i e t a ry combinations of plant and animal or ig in feedstuffs

(Jhingran, 1983). Intensive c u l t i v a t i o n of these f ishes

i s hampered mainly because of the non-ava i l ab i l i ty of

balanced r a t ions formulated on the bas i s of t h e i r n u t r i ­

t i ona l requirements. The importance of studies on the

n u t r i t i o n of these f ishes has been emphasized by several

workers CDehadrai, 1988; Sinha, 1988; T r ipa th i , 1988;

and Algarswaml, 1988). Although some information has

been made ava i lab le in the recent past on the gross

pro te in requirement of these f ishes (Jayaram, 1978;

Renukaradhya and Varghese, 1986; and Singh and

Bhanot, 1988), r e l a t i v e l y l i t t l e i s known on t h e i r

d i e t a r y carbohydrate requirement and u t i l i z a t i o n (Sen

fit. a i . . , 1978; DARE, 1984; and Swamy^t ^ . , 1990).

The present study, which forms part of the

continuing program on f i sh nu t r i t i on and d ie t development,

deals with the effect of varying l e v e l s and sources of

- 8 -

dietary carbohydrate on the growth, conversion ef f i ­

ciency and carcass composition of Indian major carp,

Labeo rohita (Ham), The information obtained i s con­

sidered important for the development of cost-effective

practical rations for the culture of major carps on

sc ien t i f ic l ines .

GENERAL METHODOLOGY

- 9 -

I , Source and acc l imat iza t ion of experimental fish

Flngerl lngs of h^heq rohlta (Ham) were obtained

from the Uttar Pradesh Fish Cooperative Seed Production

Centre, Kolahar, Mathura, These were t ransported to the

laboratory In oxygen-fil led polythene bags and stocked

in ear thern pond (12 x 6 x 1.5 m). Supplementary feeding

a t 5% leve l was car r ied out using the conventional r i ce

bran: mustard o i l cake ( I j l ) feed mixture dai ly at OSOOh.

Prior t o the experiment, required number of f ish were

t ransfer red to the wet laboratory and prophylactic

treatment with 5 ppm KMno4 solut ion given. The fish were

then accustomed t o case in -ge la t in moist d ie t for a week.

I I . Experimental d i e t s

Experimental d i e t s (Table I I I & IV) were formulated

s l i g h t l y modifying the procedure used by Buhier and

Haiver (1961), Casein and ge la t in were used as protein

sources. Protein content in the d i e t s was fixed to 40%

crude prote in level according to the requirement for tiils

species worked out e a r l i e r in th i s laboratory (Ann, Tech.

Rep. 1987). Diets for studying the effect of varying

l eve l s of carbohydrate on the growth and conversion

eff ic iency of f ish were formulated by varying the dextr in

component from 20-35% a t the cost of cK^-cellulose. The

- 1 0 -

d iges t i b l e energy i n the t e s t d i e t s ranged from 308 to

353 kcal/100 g dry d i e t . The energy to prote in r a t i o in

the experimental d i e t s ranged between 7,70-8.82 kca l /g

pro te in .

In the experiment designed for evaluat ing the

effect of d i f fe ren t carbohydrate sources on the growth,

conversion ef f ic iency and carcass composition of f i sh ,

equal q u a n t i t i e s of the sources (glucose, sucrose,

f ructose , potato s t a rch and dextr in) were incorporated in

the experimental d i e t s . The gross energy i n the t e s t d i e t s

ranged from 413.70 t o 428.10 kcal/100 g dry d i e t . Dietary

fat used was a mixture (2$1) of corn and cod l i v e r o i l .

Vitamin and mineral premixes (Table I&II)were prepared

according to Halver (1976). Carboxymethyl ce l lu lose was

used as binder.

Calculated q u a n t i t i e s of the d ie t a ry ingredients

were weighed on a s ingle pan e l ec t ron ic balance. A known

amount of water was heated to 80"C in a s t a i n l e s s s t e e l

attachment of Hobart e l e c t r i c mixer. Gelatin was dissolved

i n t o i t with slow s t i r r i n g and heating the mixture. Dextrin

and mineral mix were added a f t e r removing the mixture bowl

from heat ing. Casein was dissolved and blended i n the

mixture while the contents were s t i l l i n lukewarm s t a t e .

This was followed by the addi t ion of remaining d ie t a ry

i ng red i en t s , excepting the carboxymethyl ce l lu lose , which

-11-

was added in the l a s t . As the d i e t began t o solidify^

the speed of s t i r r i n g was increased gradually t o incor ­

porate a i r i n t o the mixture. The f inal d i e t , about the

consistency of bread dough, was poured i n t o a te f lon -

coated pan, and f i na l ly placed i n t o a r e f r i g e r a t o r to

j e l l . The moist d i e t (approx, 50% moisture) was cut

i n to cubes and kept i n p l a s t i c sealed boxes in a freezer

u n t i l used,

I I I . Fseding t r i a l

Elngerlings of 4,0-6,0 era ( t o t a l length) were

sorted out from the acclimatized lo t maintained i n the

wet laboratory and stocked i n t r i p l i c a t e groups of 30

f i sh each in 55 1 capaci ty high densi ty polyvinyl c i r c u l a r

p l a s t i c troughs f i t t ed with a continuous water flow-through

system. The flow r a t e in each trough was adjusted to

1-1.5 l i t / m i n .

Pish were fed d i e t s in the form of moist cake

d a i l y a t 0800 and 1600h, except on the day when weekly

measurements were taken. The feeding l eve l and schedule

were chosen a f t e r carefu l ly observing the feeding beha­

viour of the f i sh and t h e i r d i e t a ry in take . Accumulation

of the d i e t a t the bottom was avoided t o prevent any

12-

p o s s i b l e l e a c h i n g of t h e n u t r i e n t s . I n i t i a l and s u b s e ­

quent weekly weight gains were c a r e f u l l y r eco rded , a f t e r

a n a e s t h e t i z i n g t h e f i s h wi th MS-222, A thorough weekly

scrubbing and c l e a n i n g of each t rough was a l s o c a r r i e d

o u t . The feeding t r i a l s l a s t e d for a pe r iod of s i x

weeks. The average water t empera tu re and d i s s o l v e d

oxygen over the expe r imen ta l p e r i o d , baged on d a i l y

measurements, ranged between 22-25*C and 6 , 8 - 6 , 4 ppm,

r e s p e c t i v e l y . Na tu ra l photoper iod ( l i g h t t dark

cyc le ) was mainta ined du r ing the exper iment .

IV. B i o l o g i c a l e v a l u a t i o n of feed and growth parameters

Growth parameters ove r t h e e-week feeding t r i a l

were a s se s sed a s f o l l ows :

Live weight ga in {%}

- I L - J S L - 2 X 100 ! « W n o

S p e c i f i c growth r a t e (%)

log^ i w^ - Log i„w^ X 100

Gross growth e f f i c i e n c y

Ir,W^ - I„W « n t n o Food I n t a k e , g

- 1 3 -

Pood Conversion r a t i o

Pood Intake (dry weighty g) weight gain (wet weight ,g)

Protein e f f i c i e n c y r a t i o

Wfeight gain (wet weightyg) Protein in take , g

percent prote in re tent ion

BCP - BCP Z O

Protein in take , g X 100

percent energy re ten t ion

BB - BE t O

Energy in take , kcal X 100

where, I W_ " I n i t i a l average weight, g n o

^n\ " ^"^^ average weight, g

D = Duration

BCP = I n i t i a l body crude prote in , g

BCP = Pinal body crude prote in , g

BE = I n i t i a l body energy, k c a l .

BE^ = Pinal body energy, kca l .

-14-

Crude prote in

The est imat ion of crude prote in was made by

s l i g h t l y modifying the Wong»s micro-Kjeldhal method^ as

adapted by J a f r i ^ t §1» (1964). 0.1 gm of sample was

digested in 5 ml of l i l sulphuric acid and heated t i l l

fumes appeared. 0,5 ml of saturated potassium persulphate

was then added t o oxidize the d iges t ing mixture. The

d iges t ion was car r ied out for about 12-16 hours t i l l the

so lu t ion in the Kjeldhal flask became water c l ea r , i n d i ­

cat ing tha t a l l the nitrogenous mater ia ls present in the

sample has been converted i n t o ammonium sulphate . The

so lu t ion was t ransfer red t o a volumetric flask and di luted

t o 50 ml. An a l iquot (0,5 ml) of t h i s solut ion was then

nesae l i r ized with Bock and Bendict Nesslers reagent (Oser,

1965), The solut ion was kept a t room temperature for 10

mts for complete colour development. The colour in ten­

s i t y was read on a micro-processor control led splitbeara

spect ronic 1001 spectrophotometer a t 480 m^ wave length,

a f t e r adjust ing the instrument with proper blank. The

o p t i c a l densi ty obtained was r^ad off against a standard

c a l i b r a t i o n curve for ca lcu la t ing the amount of nitrogen

(Fig, 1 ) . The quan t i ty of t o t a l nitrogen was multiplied

with conventional pro te in factor (6,25) t o obtain the crude

prote in content of the sample. The values were calculated

- 1 5 -

as percent on dry weight b a s i s .

The data were s t a t i s t i c a l l y analysed using the

standard methods (Snedecor and Cochran, I968),

V, Proximate composition of feed and carcass

Proximate feed and carcass composition was

estimated according to standard procedures (Hardy, 1989).

Moisture

Moisture content was determined, drying a known

amount of sample t o constant i n hot a i r (80^C) oven.

Dried powdered sample (1-2 g) was incinerated in

a muffle furnace a t 600*0 t i l l i t became completely white

and free of carbon. After cooling in a dess ica to r , the

content was weighed to est imate the amount of ash as

percent of dried sample.

Crude fat

crude fat was estimated by soxhelet exhaustive

ex t rac t ion technique using petroleum e ther (40-60* B. P.)

as solvent .

-16-

Nltrogen-free ext rac t (NFS)

NPE was determined I n d i r e c t l y , subtract ing the

amounts of moisture, crude pro te in , e t he r ex t r ac t (fat)

and ash from 100,

Gross energy

The es t imat ion of gross energy content was made

through d i r ec t bomb caiorimetry. Pr ior to es t imat ion,

the sample was dr ied a t 100*C for 24 hours. A known amount

of sample was taken in to a meta l l i c c ruc ib le , compacted

carefu l ly , and allowed combustion in presence of 25 lb

oxygen pressure . The heat generated upon combustion was

read on the modulated galvanometer sca le , and converted

t o energy equivalent , worked out e a r l i e r using thermo-

chemical grade benzoic acid as standard. The gross energy

i n the sample was expressed as c a l / g .

Table I . Vitamin Premix

Vitamin g/100 g d i e t

Alpha c e l l u l o s e (carr ier )

Choline ch lor ide

I n o s i t o l

Ascoxt>lc acid

Niacin

Ca pantothenate

Riboflavin

Menadione

Pyrldoxine HCl

Thiamin

Fo l i c acid

Bio t in

Alpha tocopherol acetate

Vitamin B

* *

8.000

0.500

0.200

0.100

0.075

0.050

0.020

0.004

0.005

0.005

D.0015

0.0005

0.040

10 mg/500 ml H O

* Halver (1976)

**Incorporated with o i l .

* Table I I , Mineral premix

Mineral g/100 g d ie t

Calcium biptosphate 13.580

Calcium l a c t a t e 3 2.580

Fer r ic c i t r a t e 2,970

Magnesium s u l f a t e 13.700

Potassium phosphate (dibasic) 23.980

Sodium biphosphate 8.720

Sodium chlor ide 4.3 50

Alluniura ch lo r ide , 6H2O 0.015

Potassium iodide 0.015

Copper ch lor ide 0.010

Manganous su lphate , H O 0*080

Cobalt ch lor ide , SH-O 0.100

Zinc sulphate , 7H2O 0.300

M i l f c i ^ l I ' l l l i iJIII

*Halver (19 76)

0.01 0.02 0.03 0.04 0.05 0.06

NITROGtM ( mg 1

0.07 0.08 0.09

Fig. 1, Calibration curve of nitrogen.

CHAPTBR I

EFFECT OF VARYING DIETARY CARBOHYDRATE LEVELS ON THE

GROWTH AND CONVERSION EFFICIENCY OF THE INDIAN MAJOR

CARP^ LABEO RQHITA (HAM.)

INTRODUCTION

A l t h o u g h d i g e s t i v e and m e t a b o l i c s y s t e m s of f i s h e s

a r e known t o be b e t t e r a d a p t e d t o u t i l i z a t i o n of p r o t e i n

and l i p i d f o r e n e r g y t h a n c a r b o h y d r a t e s , some warmwater

s p e c i e s , p a r t i c u l a r l y h e r b i v o r o u s and o m n i v o r o u s , can

d i g e s t and m e t a b o l i z e c a r b o h y d r a t e s r e l a t i v e l y w e l l . The

u t i l i z a t i o n of d i e t a r y c a r b o h y d r a t e and i t s i n f l u e n c e on

g r o w t h have been examined i n s e v e r a l f i s h e s l i k e salmon

( B u h l e r and H a l v e r , I 9 6 I , Edwards , fit ^ . , 19 7 7 ; H i l t o n

and A t k i n s o n , 1 9 8 2 ) , cod (Kaushik 2I SX" 1 9 8 9 ) , y e l l o w -

t a i l (Furukawa, 1976; and F u r u i c h i and Yone, 1 9 8 0 ) , red

s e a bream ( F u r u i c h i and Yone, 1980 ; and M o r i t a ^ a ^ . ,

198 2), c h a n n e l c a t f i s h ( G a r l i n g and Wi l son , 1 9 7 7 ) , s t u r ­

geon and European e e l s (Wilson and Poe , 1987, and

Ande r son g t 2 l»# 1 9 8 4 ) ,

The c a r b o h y d r a t e u t i l i z a t i o n i n c a r p s i s n o t

c l e a r l y u n d e r s t o o d . However, t h e work of Shimeno ^t , . §X''

• 1 8 -

( 1 9 8 1 ) , Z i e t l e r e t ai , . (1983), , Vio la and A r i e l ! ( 1 9 3 3 ) ,

and Ufodike and Ma t ty (1983) on common c a r p p o i n t t o

t h e f a c t t h a t c a r p s a r e a b l e t o t o l e r a t e e l e v a t e d l e v e l s

o f d i e t a r y c a r b o h y d r a t e s a s an e f f i c i e n t e n e r g y s o u r c e .

I n s p i t e o f h i g h e r l e v e l o f c a r b o h y d r a t e u t i l i z a t i o n by

c a r p , t h e s e f i s h e s have been r e p o r t e d t o e x h i b i t g r e a t e r

d i s p a r i t y i n t h e l e v e l s of u t i l i z a t i o n o f t h i s n u t r i e n t

( j a u n c e y , 1 9 8 2 ) . Takeuch i e t . a^.. (1979) and P u r u i c h i

e t a i . (1987) c o n c l u d e d t h a t c a r p s can grow w e l l even

on d i e t s c o n t a i n i n g low l e v e l s o f d i e t a r y c a r b o h y d r a t e .

No d e t a i l e d i n f o r m a t i o n on t h e s e a s p e c t s i s so f a r

a v a i l a b l e i n t h e I n d i a n ma jo r c a r p s . The p r e s e n t s t u d y

d e a l s w i t h t h e e f f e c t of v a r y i n g d i e t a r y c a r b o h y d r a t e

l e v e l s on t h e g rowth and c o n v e r s i o n e f f i c i e n c y of t h e

f i n g e r l i n g L. r o h i t a .

MATERIALS AND METHODS

F i n g e r l i n g of l^. r o h i t a ( 5 , 4 t o 6 . 7 cm t o t a l

l e n g t h ) , s o r t e d o u t from t h e a c c l i m a t i z e d f i s h l o t

m a i n t a i n e d i n t h e wet l a b o r a t o r y , were s t o c k e d f o r f e e d i n g

t r i a l s i n p o l y v i n y l c i r c u l a r t r o u g h s . The c o m p o s i t i o n

of t h e . e x p e r i m e n t a l d i e t s u sed i s g i v e n i n T a b l e I I I .

The d e t a i l s of t h e f e e d i n g t r i a l s , s t o c k i n g d e n s i t y and

method of p r e p a r a t i o n of e x p e r i m e n t a l d i e t s have been

- 1 9 -

m e n t i o n e d u n d e r G e n e r a l Methodo logy s e c t i o n (page 9 - 1 2 ) .

Each d i e t a r y t r e a t m e n t was g i v e n i n t r i p l i c a t e . D i e t s

were fed i n t h e form o f s o f t / m o i s t c a k e , t w i c e d a i l y a t

0800 and I 6 0 0 h , a t t h e t o t a l i ra te of 4% body w e i g h t . No

feed was g i v e n t o t h e f i s h on t h e day t h e g r o w t h m e a s u r e ­

m e n t s were t a k e n . Weekly growth m e a s u r e m e n t s and

b i o l o g i c a l e v a l u a t i o n o f t h e d i e t were c a r r i e d o u t a c c o r d i n g

t o t h e s t a n d g r d p r o c e d u r e s d e s c r i b e d e l s e w h e r e (page 12-13) ,

RESULTS AND DISCUSSION

Data c o n c e r n i n g t h e g r o w t h and c o n v e r s i o n

e f f i c i e n c i e s o f It. r o h i t a f i n g e r l i n g fed v a r y i n g l e v e l s

of d i e t a r y c a r b o h y d r a t e have been shown i n T a b l e IV .

Dur ing t h e 6-week f e e d i n g t r i a l , a v e r a g e l i v e

w e i g h t and t h e p e r c e n t a g e o f we igh t g a i n o f t h e f i s h a t

d i f f e r e n t d i e t a r y t r e a t m e n t s i n c r e a s e d l i n e a r l y o v e r

t h e i r i n i t i a l v a l u e s ( P i g . l & 2 ) . The g a i n i n l i v e weigh t

v a r i e d s i g n i f i c a n t l y (ANOVA P < 0 . 0 1 ) w i t h t h e l e v e l s of

d i e t a r y c a r b o h y d r a t e . The f i s h r e g i s t e r e d t h e maximum

(64%) we igh t u p t o 30% d i e t a i r y c a r b o h y d r a t e i n t a k e .

Beyond t h i s l e v e l , a s i g n i f i c a n t ( P < 0 . 0 1 ) f a l l i n l i v e

w e i g h t was e v i d e n t . C a l c u l a t e d we igh t g a i n s

on e a c h d i e t , worked o u t t h r o u g h r e g r e s s i o n e q u a t i o n ,

compared w i t h t h e a c t u a l w e i g h t g a i n s ( F i g . 3 ) .

- 2 0 -

The speci f ic growth rate of f i sh for various

d i e t a ry treatments have been plot ted in Fig. 4,

Specific growth ra te increased almost l i nea r ly with

increasing d i e t a ry carbohydrate level up to 30%

carbohydrate incorporat ion l eve l .

The food conversion r a t i o (as fed to wet weight

bas is) and ganoss growth ef f ic iency calcula ted for

d i f ferent l eve l s of d i e t a ry carbohydrate have been plotted

in Fig. 5, The best food conversion r a t i o ( 2 . 8 J 1 )

as well as gross growth eff ic iency (0.35) were obtained

a t 30% d ie t a ry carbohydrate. The conversion eff ic iency

got adversely affected a t elevated (35%) leve l of

carbohydrate in take .

The prote in ef f ic iency r a t i o (PER) was a lso found

t o increase progressively up to 30% carbohydrate inclusion

in the d i e t . A d i s t i n c t f a l l in PER was observed in

f i sh fed 35% carbohydrate d ie t (Fig. 6 ) .

The optimum levels of d ie ta ry carbohydrate for most

f ishes are reported to be within the range of 10-30%

(Fuiukaw^, 1976; Furuichl and Yone, 1980; Morita e^ a^. ,

1982; Buhler and Halver, I96I; Edwards et, a^, , 1977;

Cowey and Sargent, 1972; Bergot, 1979; Garling and Wilson,

1977; Metai ier e t aj^. , 1980; Degani, 1987; and Furuichi

- 2 1 -

e t a l . , 1987), with t i l a p l a perhaps being the only-

exception where leve l of carkohydrate in the d i e t as

high as 40% did not depress growth (Anderson e^ a l . ,

1984).

Although carps are a lso known to t o l e r a t e and

u t i l i z e r e l a t i v e l y high l e v e l s of carbohydrate incor­

poration in the d ie t (Shimeno e t a l . , 1981; Z i e t l e r

et. ajL., 1983; Viola and A r i e l i ; 1983; Ufodike and Matty,

1983; Jhingran and Pul l in , 1988; and Puruichi and Yone,

1980), theire i s s t i l l a considerable d i s p a r i t y as to

the ef f ic iency of t h e i r carbohydrate u t i l i z a t i o n .

Takeuchi et, aj^. (1979), and more recent ly Furuichi et, a^,.

(1987), have indicated tha t carps grew well even on d i e t s

containing low l eve l s of d ie ta ry carbohydrate.

In general , warmwater f ish have no t rue carbohyd­

ra te requirement (NAS-NRC, 1983) but incorporation of

c e r t a in level of carbohydrate in the d ie t influences

conversion e f f i c i e n c i e s and overa l l growth of the f ish

as i s evident from the r e s u l t s obtained during the present

study on L. rohi ta finger l ing. An increase in carbohydrate

from 20-30% in d ie t improved the growth and the conversion

e f f i c i enc i e s of the f ish . These findings seemed to

conform with the observations of Degani and Levanon (1987)

and Degani (1987) on European e e l , and Sen e t . a ^ . , (19 78) on

- 2 2 -

spawn, fry and f inger l inga of common carp and It. rohlta

fry. More or l e s s s imi lar r e s u l t s were reported on

£. mrlqala (DARE. 1984) and C. ca t la (Swaroy e_t a^.. , 1990).

The fa l l in d ie ta ry performance beyond a fixed (30%) level

of carbohydrate intake as observed in j ^ . rohita was

a lso evident in rainbow t rou t (Ek3wards et, a i . , 1977;

Hilton and Atkisson, 1982), Common carp (Shlmeno, 1982),

and C. ca t la (Swamy e t a^, , 1990). In cod, however,

increas ing the amount of carbohydrate in the d ie t from

0 to 30% was not found to influence weight gain and

re ten t ion values of protein arKS fa t , ind ica t ing tha t the

f ish was unable to u t i l i z e carbohydrate as a source of

energy (Hemre et_ aj^., 1989).

Bet ter performance of d i e t a t 30% carbohydrate

level in j ^ . rohi ta f inger l ing could be the r e s u l t of

grea ter use of prote in for growth purposes. The sparing

ac t ion of carbohydrate on prote in has a l so been indicated

in several o ther f i sh species (NAS-NRC, 1981, 1983).

The pa t te rn of changes in PER of i^, rohita

f inger l ing with varying l eve l s of d i e t a ry carbohydrate

a l so indica te b e t t e r prote in u t i l i z a t i o n a t the above

leve l of inclus ion in the d i e t . A coro l la ry to t h i s

finding was apparent in the work of Buhler and Halver

(1961), Bergot (1979), Lee and Putnam (1973), Pie per and

- 2 3 -

Pfeffer (1979,1980) in salmonids, Adron, £ t . a^. (1976) in

tu rbo t , Cowey, e t a^.. (1975) in p l a i ce , Garling and

Wilson (19 77) in channel ca t f i sh , A l l i o t fit a^. (19 78)

in seabass, and Ufodike and Matty (1983) in niirror

carp. In these f i shes increased PER occurred with

increas ing d ie t a ry carbohydrate up t o the requirement

l eve l . In European eel and cod, however, the PER was

reported to decrease with increased carbohydrate intake

(Degani and Viola, 1987; and Hamre e_t aj^., 1989). The

present study strengthens the view that, ' i n fishes the ut i l i ­

zation of d i e t a ry carbohydrate I s species spec i f ic , and

tha t inclus ion of carbohydrate up to a fixed level in

the d ie t may improve growth and feed eff ic iency, besides

making the d ie t cost effectives.

The depressed growth and poor feed eff ic iency

with d i e t s containing l eve l s of carbohydrate higher than

requirement may point to the fact tha t t h i s f ish i s

unable to handle excessive carbohydrate component in

the d i e t . The reduction of growth a t 35% d ie t a ry

carbohydrate incorporat ion may a l so be a t t r i b u t e d to

the a l t e r ed energy to prote in r a t i o , from 8,45 kcai /g

prote in in 30% carbohydrate d i e t to 8.82 kca l /g protein

in 35% carbohydrate d i e t . In an e a r l i e r experiment

conducted a t t h i s laboratory on the energy requirements

of if rohi ta fingerling^ optimum growth was s imi lar ly

noted a t an energy to protein r a t i o of 8.33 kcal /g

protein (Ann. Tech. Rep, 1988).

- 2 4 -

The g r o s s c a r b o h y d r a t e r e q u i r e m e n t o f t h i s f i s h

a t f i n g e r l i n g s t a g e a p p e a r t o be 30X o f d i e t a r y i n c l u s i o n ,

c o r r e s p o n d i n g t o 338 k c a i / 1 0 0 g d r y d i e t , a t 23 + 2 ' C .

I t may, however , b e p o i n t e d o u t t h a t t h e o v e r a l l low

g r o w t h t r e n d o b s e r v e d i n t h e p r e s e n t s t u d y may be t h e

r e s u l t o f low w a t e r t e m p e r a t u r e , s i n c e I n d i a n majo r

c a r p s p r e f e r s l i g h t l y h i g h e r t e m p e r a t u r e , c l o s e t o 30*0

( J h i n g r a n and P u l l i n , 1 9 8 8 ) . However, low t e m p e r a t u r e

d u r i n g t h e e x p e r i m e n t d i d no t show a n y d e l e t e r i o u s

e f f e c t on t h e s u r v i v a l o f t h e f i s h .

SUMMARY

E f f e c t o f f e e d i n g v a r y i n g l e v e l s o f d i e t a r y

c a r b o h y d r a t e i n p u r i f i e d t e s t d i e t s on t h e g rowth and

c o n v e r s i o n e f f i c i e n c y o f J^. r p h i t a ( Ham,) f i n g e r l i n g

have been r e p o r t e d . Dur ing t h e 6-week f e e d i n g t r i a l

a l i n e a r g rowth p a t t e r n (ANOVA, P ^ O . O l ) was n o t e d .

The l e v e l o f c a r b o h y d r a t e t h a t p r o d u c e d t h e maximum

g rowth and b e s t ffeed e f f i c i e n c y was found t o be 30?i,

c o r r e s p o n d i n g t o a n e n e r g y t o p r o t e i n r a t i o o f 8 .45

k c a l / g p r o t e i n . v a r i o u s g rowth p a r a m e t e r s were found

n e g a t i v e l y a f f e c t e d ( P < 0 . 0 1 ) beyond t h e above l e v e l

o f c a r b o h y d r a t e i n c o r p o r a t i o n i n t h e d i e t .

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DURATION ; V,';ok )

P i g . 1, Average weekly changes i n f i s h weight (g) a t va ry ing (X — X 20%; o- <, 25%; • • 30%; and * —4 35%) carbohydra te l e v e l s .

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D U R A T I O N ( W t e k )

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20 25 30 35

DIETARY CARBOHYDRATE (V.)

Fig. 4. Specif ic growth rate (%) of i . rohita at varying carbohydrate l e v e l s .

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DIETARY CARBOHYDRATE ('/.)

35

Fig. 5, Food conversion r a t i o (• • ) and gross growth ef f ic iency (o_ o) of L. rohi ta fed varying carbohydrate l e v e l s .

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DIETARY CARBOHYDRA! t IV . )

Pig. 6, Protein efficiency rat io of i . rohita fed varying carbohydrate levels .

CHAPTER I I

GROWTH, CONVERSION EFFICIENCY, AND CARCASS COMPOSITION

OF INDIAN MAJOR CARP, lABEQ ROHITA (HAM.) FED DIFFERENT

CARBOHYDRATE SOURCES

INTRODUCTION

The source and complexi ty of ca rbohydra t e s and

the presence of ca rbohydra te me tabo l i z ing enzymes a r e known

to in f luence the u t i l i z a t i o n of ca rbohydra te i n f i s h

(NAS-NRC, 1983).

Complex p o l y s a c c h a r i d e s and d i s a c c h a r i d e s a r e

r e p o r t e d l y l e s s u t i l i z e d by f i s h e s l i k e salmonids (Buhler

and Halver , 1961; Edwards e t a^.. , 1977; Austreng e t a 1. ,

1977; Bergot , 1979; and Hi l ton and Atkinson, 1982), Yellow-

t a i l (shimeno, 1982; Shimeno et_ aj^,, 1979; and Furuichi

and Yone, l9 7 l ) , s turgeon (Hung et^ a_l., 1989) and European

e e l (Degani and Levanon, 1987). On t h e o t h e r hand, channel

c a t f i s h (Dupree I966; Gar l ing and Wilson, 1977; and Wilson

and Poe, 1987), red sea bream (Furuichi and Yone, 1971,

1980), t i l a p i a (Tung and Shiau, 1989) and common c a r p

(Shimeno et . a_i,., 198 l i and Furuichi and Yone, 1980, 1982)

a r e a b l e t o u t i l i z e such complej: po ly saccha r ide s more

e f f i c i e n t l y compared t o simple sugar s . P r a c t i c a l l y nothing

i s known about u t i l i z a t i o n of d i f f e r e n t ca rbohydra tes

i n Ind ian major c a r p s . The p resen t study was conducted

with a view t o e v a l u a t e the u t i l i z a t i o n of d i f f e r e n t

- 2 6 -

c a r b o h y d r a t e s o u r c e s , a s r e f l e c t e d by c h a n g e s i n g rowth ,

c o n v e r s i o n e f f i c i e n c y and c a r c a s s c o m p o s i t i o n o f t h e

I n d i a n majo r c a r p , Labeo r o h i t a (Ham, ) .

MATERIALS AND METHODS

F l n g e r l i n g of L, r o h i t a ( 4 . 6 t o 6 .0 cm t o t a l

l e n g t h ) , a c c l i m a t i z e d i n wet l a b o r a t o r y , were used f o r

t h e t r i a l s . D e t a i l s o f t h e a c c l i m a t i z a t i o n o f f i s h ,

f e e d i n g t r i a l , s t o c k i n g d e n s i t y and p r e p a r a t i o n of

e x p e r i m e n t a l d i e t s have been g i v e n u n d e r G e n e r a l Metho­

d o l o g y s e c t i o n (page 9-12). The c o m p o s i t i o n of t h e

e x p e r i m e n t a l d i e t s used has been shown i n T a b l e V .

D i e t s were fed i n t h e form of s o f t / m o i s t cake^ t w i c e

d a i l y a t 0800 and ISOOh, a t a t o t a l r a t e o f 4% body

w e i g h t , e x c e p t on t h e day when week ly measuremen t s were

t a k e n .

At t h e end o f t h e 6-week f e e d i n g t r i a l , 10 f i s h

were t a k e n o u t from e a c h l o t f o r e s t i m a t i n g t h e c a r c a s s

c o m p o s i t i o n . The s a m p l e s were f r o z e n , l a t e r homogenized

and a n a l y s e d . S i m i l a r a n a l y s e s o f c a r c a s s c o m p o s i t i o n

were made on an sample o f 10 f i s h removed a t

t h e b e g i n i n g of t h e f e e d i n g t r i a l . B i o l o g i c a l e v a l u a t i o n

of feed and c a r c a s s c o m p o s i t i o n o f f i s h were made

a c c o r d i n g l y t o t h e p r o c e d u r e s d e s c r i b e d e l s e w h e r e (page 12-16).

- 2 7 -

RESULTS AND DISCUSSION

Resu l t s concerning the growth and convers ion

e f f i c i e n c y of Jg, r o h i t a f i n g e r l i n g fed d i f f e r e n t carbo­

h y d r a t e sources ove r a 6-week feeding t r i a l have been

shown i n Table VI .

The average weight of f i s h for t h e t e s t d i e t s

used i n c r e a s e d l i n e a r l y on t h e i r i n i t i a l weights

(F ig . 1 ) . S i g n i f i c a n t (ANOVft P <0.05) d i f f e r e n c e s

were n o t i c e a b l e i n t h e growth and feed e f f i c i e n c y of

the f i s h with d i f f e r e n t d i e t a r y carbohydra te sources .

Maximum (100%) growth, i n terms of l i v e weight ga in ,

was observed wi th s u c r o s e , followed by f ruc tose (85%),

g lucose (78%) and d e x t r i n (71%) d i e t s (F ig . 2 ) • The

p o t a t o s t a r c h con ta ined d i e t produced the minimum

(52.4%) growth.

The average s p e c i f i c growth r a t e of f i s h with

v a r i o u s d i e t a r y t r e a t m e n t s have been p l o t t e d i n Fig. 3.

The h ighes t (1,65%) s p e c i f i c growth r a t e was noted i n

f i s n fed sucrose conta ined d i e t and the lowest (1,0%)

i n i n those fed the po t a to s t a r c h based d i e t .

F ig . 4 d e p i c t s the food convers ion r a t i o and

g ros s growth e f f i c i e n c y of f i s h fed the d i f f e r e n t

ca rbohydra t e s o u r c e s . The bes t ( 2.28 t 1 ) food

- 2 8 -

conversion r a t i o (as fed to wet weight basis) was observed

with sucrose d i e t , followed by fructose d i e t . Fbod

conversion r a t i o s were r e l a t i v e l y low in f ish fed the

glucose and dex t r in incorporated d i e t s . Potato s tarch

d i e t produced the lowest conversion r a t i o . Almost

s imi lar pa t te rn was observed for the gross growth

ef f ic iency of f i sh vd.th d i f ferent carbohydrate sources.

Gross growth e f f i c i enc ie s were comparable in f ish fed

glucose and dextr in source d i e t s .

The PER of f ish for various d ie ta ry treatments

have been depicted i n Fig. 5, The highest ( 1.10 )

PER was noticeable i n f i sh fed the sucrose d i e t . PER

values in f ish tha t received the fructose, dext r in or

glucose contained d i e t s were not found to vary d i s t i n c t l y

from each other . The lowest PER was noted with the

potato s ta rch d i e t .

The i n i t i a l and f inal carcass composition of

f i sh fed d i f fe ren t carbohydrate d i e t s have been given

in Table v i l • Crude prote in and fat were found to

increase s ign i f i can t ly ( P < 0 . 0 5 ) , whereas moisture, ash

and NFE declined over the period of growth t r i a l , A

comparision of the carcass composition among f ish fed

d i f fe ren t carbohydrate sources ind ica te tha t changes in

body cons t i tuen ts were r e l a t i v e l y more d i s t i n c t in

f i sh maintained on the sucrose d i e t .

- 2 9 -

Protein and energy re ten t ion values of f ish fed

d i f fe ren t carbohydrate sources have been shown in Table VTII,

Protein and energy re ten t ion were the highest with sucrose

and lowest with potato s tarch d i e t s .

Results of the present study c l e a r l y indicate

tha t growth, feed eff iciency and carcass composition in

f ish are markedly influenced by the nature of carbohydrate

sources in the d i e t , and t h a t , not a l l carbohydrates are

u t i l i z e d with the same eff ic iency. Similar phenomena were

seen in salmonids (Buhler and Halver, I96 I ) , ye l lowta i l

(Shimeno £t, a l , , 1979; and Furuichi and Yone, 1971),

channel ca t f i sh (Dupree, I966; Garling and Wilson, 1977;

and Wilson and Poe, 1987), t i l a p i a (Anderson £ t £ l , , 1984;

and Tung and Shiau, 1989), red sea bream (Furuichi and

Yone, 1971, 1980), European eel (Degani fit ^ . , 1986),

sturgeon (Hung gt §X», 1989; and Kaushik e t ^ , , 1989),

cod and common carp (Hemre £t_ a i . . 1989; Shimeno e_t a l . ,

1977, 1981; Takeuchi fit ^ l* # 1979; Furuichi and Yone,

1980, 1982).

Although dif ferences in the growth parameters

were not s ign i f i can t in fructose and glucose fed f i sh ,

sucrose , a disaccharide containing fructose and glucose

molecules in i t s s t r u c t u r e , proved an excel lent carbo­

hydrate source for L, rohl ta f inger l ing . The low per-

- 3 0 -

formance of d i e t s containing dex t r in or potato s tarch

was r a t h e r an unexpected phenomenon in view of the fact

t h a t i n several of the omnivorous and herbivorous f ishes

the u t i l i z a t i o n of cotnplex carbohydrates, compared to the

disaccharides and simple sugars, has been reported t o be

much higher (Wilson and Poe, 1987; Shiroeno fit aX», 1981;

Shimeno, 1982; Puruichi and Yone, 1980, 1982; and Tung

and Shiiau, 1989.

The i n f e r i o r performance of potato s tarch and

dext r in containing d i e t s , and the u t i l i z a t i o n of mono- and

d isacchar ides , could be re la ted to the presence of specif ic

enzymes, and to the ove ra l l d i g e s t i b i l i t y of these carbo­

hydrates in carps . Kawai and Ikeda (19 71) have c l ea r ly

pointed out tha t i n common carp, carbohydrate digest ing

enzymes exis t i n varying l e v e l s . High l eve l s of carbo­

hydrate metabolizing enzymes and t h e i r s ignif icance have

been reported in f i sh i n t e s t i n e s , p a r t i c u l a r l y in herb i ­

vores (smith, 1989^), The d i g e s t i b i l i t y of mono- and

disacchar ides i s repor tedly superior (over 98.OX) to tha t

of the complex saccharides in a numbur of f ish species

(NAS-NRC, 1981). In f ishes l ike t i l a p i a (Anderson e t a^ , ,

1984; and Tung and Shiau, 1989), channel ca t f i sh (WLlson

and Fbe, 1987) and common carp (Furuichi and Yone, 1982),

the most d iges t ib le ( > 99.OX) carbohydrate, such as

glucose, was not found to r e s u l t i n increased growth or

- 3 1 -

produce b e t t e r feed ef f ic iency. This was explained in the

l i gh t of the fact t ha t a la rge port ion of the absorbed

glucose in these f i shes probably gets excreted out before

adequate in su l in i s avai lable to f a c i l i t a t e the t i s sue

u t i l i z a t i o n of glucose.

Results on prote in and energy re ten t ion of

i . rohi ta f inger l ing , as re f lec ted by changes in carcass

composition, grovrt:h and conversion eff ic iency, also i n d i ­

cates t h a t , although the f i sh i s able t o u t i l i z e d i f ferent

carbohydrate sources, i t can more e f fec t ive ly use sucrose

as a d ie ta ry component tha t seems t o f a c i l i t a t e growth by

sparing protein for prote in purposes. S ign i f ican t ly ,

compared t o the o ther d i e t s , the sucrose d ie t resul ted in

increased body crude protein content , followed by a sharp

decrease in NPE value. Higher fBR values observed in f ish

fed the above d i e t a l so strengthens the fact that the

ingested sucrose was r ead i ly u t i l i z e d for energy purposes.

However, a r e l a t i v e increase in the deposit ion of fat in

the body a l so occurred in such f ish (Table VI I ) , i nd ica t ing

an e f fec t ive l ipogenesis from t h i s d i e t a ry carbohydrate

source. The findings of the present study seem i n agreement

with the r e s u l t s reported on channel c a t f i sh (Wilson and

Poe, 1987), sturgeon (Hung et. SLL» i 1989; and Kaushik e t a i . ,

1989), European eel (Degani e t a^. , 1986), and Chinook

salmon (Buhlar and Halver, 1961).

-3 2 -

Indepth s t u d i e s i nvo lv ing the d i s t r i b u t i o n and

c h a r a c t e r i z a t i o n of ca rbohydra te metabolism enzymes in

i . r o h i t a f i n g e r l i n g a r e , however, needed t o e x p l a i n the

s i g n i f i c a n c e of t h e above r e s u l t s i n more c l e a r t e rms .

SUMMARY

F e e d i n g L. j:;^Qhita (Ham. ) f i n g e r l i n g w i t h v a r i o u s

d i e t a r y carbohydra te sources i n d i c a t e s t h a t , a l though t h e

f i s h i s a b l e t o u t i l i z e d i f f e r e n t ca rbohydra te sources , the

e f f i c i e n c y of t h e i r u t i l i z a t i o n i s afffected with t h e n a t u r e /

complexi ty of t h e c a r b o h y d r a t e s .

Maximum growth, convers ion e f f i c i e n c i e s ^ and

i n c r e a s e d energy and p r o t e i n r e t e n t i o n occurred in f i s h

fed suc rose d i e t , t h e changes being s i g n i f i c a n t (ANOVA

P < 0 , 0 5 ) , The v a l u e s for t h e s e parameters were found

n e g a t i v e l y a f f ec t ed (I'< 0.05) with the p o t a t o s t a r c h d i e t .

The v a r i a t i o n have been c o r r e l a t e d with changes i n c a r c a s s

composi t ion of f i s h . The s i g n i f i c a n c e of these f ind ings

have been d i s c u s s e d .

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Table VIII.Protein and energ r retention In l^, rohlta

fed the different carbohydrate sources

Source Percent Retention

PiTotein Energy

Glucose 13.53 4 24.800

Sucrose 20.800 31,480

Fructose

Potato s tarch

10.950

10.950

20,524

16,595

Dextrin 11.870 20.589

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8.0

5 ^-0'

fc.O

5.0

^.0 1 2 3 4 5 6

D U R A T I O N ( W e e k )

Fig. 1. Average weekly changes In weight (g) of i . rohi ta fed d i f fe ren t carbohydrate sources ( • • sucrose; ^ A dex t r in ; X X fructose; A A glucose; and o~ o potato s t a r ch ) .

l O O r

<

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Fig . 2.

1 2 3 4 5 6

DURATION ( Week )

Changes in l ive weight gain percent of i . rohi ta fed d i f fe ren t carbohydrate sources, (» m sucrose;

• A fructose; x- -X glucose; A< dex t r in ; and o- -o potato s t a r c h ) .

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<

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Q.

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DIET

IV

Fig. 3. Specific growth ra te (%) of I*, rohlta fed different carbohydrate sources, (diet I , glucose; diet I I , sucrose •; diet I I I , fructose diet IV, potato starch; and diet V, dextrin respectively).

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DIET

Fig, 4. Pood conversion r a t i o (o- —o) and gross growth eff ic iency (» •••••) of L- rohi ta fed d i f ferent carbohydrate sources, (d ie t I^ glucose; d ie t I I , sucrose . (jiet I I I , fructcis^ d ie t IV, potato s ta rch ; and d i e t V, dext r in r e spec t ive ly ) .

1.0

< a. > •

z ^ 0.8 LL

U-UJ

z I— o cr Q.

0.6 h

II III IV

DIET

Fig, 5. Protein efficiency ra t io of !<• rohita fed different carbohydrate sources (diet I^ glucose; diet i l , sucrose ; diet I I I , fructose,c3iet IV, potato starch; and diet V, dextrin respectively).

REFERENCES

Adron , J , W,, B l a i r , A , , Cowey, C,B. and S h a n k s , A.M.

A l g a r s w a m i , K.

A l e x i s , M.N. and P a p a p a r e s k e v a -P a p o u t s o g l o u e , E,

A l l i o t , B . , P a s t o u r e a u d , A . and N e d e l e c , J .

A n d e r s o n , J . , J a c k s o n , A.J .^ M a t t y , A . J , and Capper , B. s .

Annual T e c h n i c a l R e p o r t

1976 B f f e c t o f d i e t a r y e n e r g y l e v e l and d i e t a r y e n e r g y s o u r c e on g r o w t h , feed c o n v e r s i o n and body compo­s i t i o n of t u r b o t ( S c o p h t h a l -rjius maximus L . ) . A a u a c u l t u r e 2 : 1 2 5 - 1 3 2 .

1988 R e s e a r c h Needs f o r B r a c k i s h -w a t e r Aqua c u l t u r e i n I n d i a by 2000 A.D, I n t e r n a t i o n a l Symposium on A q u a c u l t u r e Needs f o r t h e Year 2000, New D e l h i , 15 -18 November, 1 9 8 8 , 26 p .

1986 A m i n o - t r a n s f e r a s e a c t i v i t y i n t h e l i v e r and w h i t e musc le o f MugJii c a p j t o fed d i e t s c o n t a i n i n g d i f f e r e n t l e v e l s o f p r o t e i n and c a r b o h y d r a t e . B4gg^ffl« P h v s i o l . . ^ s 2 4 5 - 2 4 9 .

1978 E t u d e d e I ' a p p o r t c a l o r i q u e e t du r a p p o r t c a l o r i c o - a z o t e d a n l * a l i m e n t a t i o n du B a r , D i c e n t r a c h u s l a b r a x . I n f l u ­e n c e s u r l a c r o i s s a n c e e t l a c o m p o s i t i o n c o r p o r e l l e , I n i P i n f i s h N u t r i t i o n and F i s h -feed T e c h n o l o g y , Vo l . I , Hsenemann, B e r l i n , F e d e r a l R e p u b l i c o f Germany ( e d s . H a l v e r , J . S. and K. T i e w s ) . p p . 2 4 1 - 2 5 1 .

1984 E f f e c t s o f d i e t a r y c a r b o ­h y d r a t e s and f i b r e on t h e t i l a p i a Qreochromls n i l o t i c u s ( L i n n ) . Aqua-c u l t u r e 37i 3 0 3 - 3 1 4 .

1987 I n v e s t i g a t i o n on t h e N u t r i ­t i o n of Some C u l t i v a b l e F i n -f i s h S p e c i e s and Development o f C o s t - e f f e c t i v e Formula F e e d s . ICAR-USDA R e s e a r c h P r o j e c t . F i s h e r i e s R e s e a r c h L a b o r a t o r y , A l i g a r h Muslim U n i v e r s i t y , A l i g a r h ( I n d i a ) . 38 p .

1988 I b i d . 50 p .

-3 4 -

Aust reng, E , , R i s a , S . , Edwards, D.J , and Hvidsten, H.

Bergot , F,

Bergot , F, and Berque, J .

Buhler , D, R, and Halver , J . E .

1977 Carbohydrate i n rainbow t r o u t d i e t s , I I , I n f luence of ca rbohydra te l e v e l s on chemical composi t ion and feed u t i l i z a ­t i o n of f i s h from d i f f e r e n t f a m i l i e s . Aquacul ture _lii 39-50.

1979 carbohydra te i n rainbow t r o u t d i e t s : E f f e c t s of the l e v e l and source of ca rbohydra te and t h e number of meals on growth and body composi t ion , Aqua-c u l t u r e i a s 157-167,

1983 D i g e s t i b i l i t y of s t a r c h by rainbow t r o u t j Ef fec ts of t h e phys ica l s t a t e of s t a r c h and of the i n t a k e l e v e l , Aquacul­t u r e 2 4 : 203-212,

I96I N u t r i t i o n of salmonid f i s h e s , IX, Carbohydrate requirement of Chinook salmon, j , Mutr, l±t 307-318.

Chiou, J , y , and Ogino, C,

1975 D i g e s t i b i l i t y of s t a r c h i n ca rp . £ u l l , J a p . Soc. S c i . F i sh . ^ : 465-466,

C i s n e r o s , J , A . , Toledo, J . and O r t i z , B,

Cowey, C.B. and Sa rgen t , J .R.

Cowey, C. B, , Adron, J . W. and Brown, D,A,

1984

1972

1975

N u t r i t i o n a l requi rements i n Qreochromis aui;eus ( t i l a p i a ) f i n g e r l i n g s 2, ca rbohydra t e -p r o t e i n r a t i o . Rev. Latinoam Aqua c u l t . 12: 29-33.

Fish N u t r i t i o n Adv. l i i : 383-492.

Mar. Bifll.

s t u d i e s on the n u t r i t i o n of marine f l a t f i s h . The metabolism of glucose by p l a i c e (P Ieu ro ­ne c t us p i a t e s a a ) and the e f f e c t d i e t a r y energy source on p r o t e i n u t i l i z a t i o n i n p l a i c e . B£. ^ . Nutr . i l : 219-231.

Cowey, C.B. and S a r g e n t , J .R.

1979 N u t r i t i o n . I n : Fish Physiology, B i o e n e r g e t i c s and Growth, Vol. V I I I . Academic P r e s s , New York (eds . Hoar, W,S., Randall , R .J , and J ,R, B r e t t ) , pp, I -69 ,

-3 5-

Cowey, C. B,

DARE

D e g a n i , G.

D e g a n i , G. , V i o l a , S. and Levanon, D,

Degani , G. and Levanon, D.

Degani, G, and Vio la , S.

Dehadra i , P. V.

DeLong, D.C. , Halver , J . E . and Mertz, E,T.

Dupree, H. K.

1988 The n u t r i t i o n of f ishx The deve lop ing scene . N u t r i t i o n Research Reviews. l_x 255-280.

1984 Annual Repor t , Department of A g r i c u l t u r a l Research and Educa t ion , Min i s t ry of Educa­t i o n , Govt, of I n d i a , 144 p .

1987 The i n f l u e n c e of r e l a t i v e p ropor ­t i o n s of d i e t a r y p r o t e i n and carbohydra te on body v*eight ga in , n i t r o g e n r e t e n t i o n and feed convers ion of European e e l s , Agu i l l a a n q u i l l a L. Aguacul ture and F i s h e r i e s Management Ifii 151-158.

1986 Ef fec t of d i e t a r y ca rbohydra te source on grovi^h and body com­p o s i t i o n of European e e l s

c u l t u r e ^ t 97-104.

1987 E f f e c t s of d i e t a r y carbohydra tes and t empera tu re s on slow growing Juven i l e e e l s , Anqui l la a n q u i l l a . Environ. Bi^l . F i sh , l^t 149-154.

1987 The p r o t e i n s p a r i n g e f f e c t s of ca rbohydra tes i n the d i e t of e e l s (Anquil la a n q u i l l a ) . Aqua-c u l t u r e 64X 283-291.

1988 Problems of Aquacul ture i n India and Southern Asian Region. I n t e r n a t i o n a l Symposium on Aquacu l tu re Needs fo r t h e Year 2000, New Delh i , 15-18 November 1988, 7 p .

1958 N u t r i t i o n of salmonid f i s h e s . VI. P r o t e i n r equ i remen t s of Chinook salmon a t two water t empera tu res , j j . Nutr . e^x 589-599.

1966 Carbohydrate molecular s i z e . In» Progress i n Sport F i s h e r i e s Research 1965, Bureau of Sport F i s h e r i e s and Wi ld l i f e , Res. Pub. 38 . pp. 129-130.

- 3 6 -

Ed w a r d s , D. J . , A u s t r e n g , E , , R l s a , S. and Gjedrem, T .

F u r u i c h l , M, and Yone, y .

1977

1971

1980

1982

F u r u i c h i , M,, I t o , G. and Yone, Y,

1987

Furukawa, 19 76

G a r l l n g , D. L, J r . and W i l s o n , R. P.

H a l v e r , J , E,

1977

1976

C a r b o h y d r a t e i n r a inbow t r o u t d i e t s , I . Growth of f i s h of d i f f e r e n t f a m i l i e s fed d i e t s c o n t a i n i n g d i f f e r e n t p r o p o r t i o n s of c a r b o h y d r a t e , A a u a c u l t u r e i l : 3 1 - 3 8 .

S t u d i e s on n u t r i t i o n of r e d sea b ream, I V . N u t r i t i v e v a l u e of d i e t a r y c a r b o h y d r a t e . Rep. F i s h , fies,. Lab . Kvushu Univ . l^t 7 5 - 8 1 .

E f f e c t o f d i e t a r y d e x t r i n on the g rowth and feed e f f i c i e n c y , t h e c h e m i c a l c o m p o s i t i o n of l i v e r and d o r s a l m u s c l e , and t h e a b s o ­r p t i o n of d i e t a r y p r o t e i n and d e x t r i n i n f i s h e s . B u l l . J a p . S a S . S c i . F l a h . j[6j 2 2 5 - 2 2 9 .

A v a i l a b i l i t y o f c a r b o h y d r a t e i n n u t r i t i o n o f c a r p and r e d aea bream. B u l l . j2d£' SOSi* SS^* fA?^. j ^ : 9 45-9 4 8 .

E f f e c t o f ^ - s t a r c h on g rowth , ffeed e f f i c i e n c y and c h e m i c a l components o f l i v e r , musc le and b l o o d o f c a r p and r ed sea bream. Nippon s u i s a n Ga^kai s h i §2* 143 7 - 1 4 4 2 .

D i e t i n y e l l o w t a i l c u l t u r e , i n : P r o c e e d i n g s o f t h e F i r s t I n t e r ­n a t i o n a l Confe rence on Aqua-c u l t u r e N u t r i t i o n ( e d s . P r i c e , K. S, Shaw, W.N. and K .S . D a n b e r g ) . Newark, D e l a w a r e : U n i v e r s i t y , 14-15 O c t o b e r , 1975 , pp . 8 5 - 1 0 4 .

E f f e c t s of d i e t a r y c a r b o h y d r a t e t o l i p i d r a t i o s on g rowth and body c o m p o s i t i o n o f f i n g e r l i n g c h a n n e l c a t f i s h . P r o g . F i s h c u l t . 21* 4 3 - 4 7 .

N u t ; c i t i o n a l d e f i c i e n c y d i s e a s e s i n s a l m o n i d s . F i s h P a t h o l o g y 1 0 : 1 6 5 - 1 8 0 .

-3 7-

Hardy, R, W.

Hemre, G , - I . , L ie , 0 . , Lied, E. and Lambertsen, G,

1989 Tb2 v i t a m i n s . I n : Fish N u t r i t i o n , Second E d i t i o n , Academic P r e s s , New York (ed. Halver , J . B . ) . pp, 32-109.

1989 Diet p r e p a r a t i o n . I n : Fish N u t r i ­t i o n , Second E d i t i o n , Academic P r e s s , New York (ed. Halver , J . E . ) , pp. 475-548.

1989 S ta rch as an energy source i n feed for cod (Gadus morhua): D i g e s t i b i l i t y and r e t e n t i o n . Aquaculture 80 : 261-270.

H i l t o n , J.W. and Atkinson, J . L.

198 2 Response of rainbow t r o u t (Salmp g a i r d n e r l ) t o i nc reased l e v e l s of a v a i l a b l e carbohydra te i n p r a c t i c a l t r o u t d i e t s . Br. j r . Nutr . 47: 59 7-607.

H i l t o n , J . W., Atk inson , j . L . and S l i n g e r , S .J ,

1987 Eva lua t ion of ne t energy value of g lucose ( ce ro lose ) and maize s t a r c h i n d i e t s for rainbow t r o u t (Salmo fl^lLrc^ner^) B£. ^ Nuty. ^ t 453-461.

Hung, S . S . O . , A i k i n s , F . K . - P . , Lu te s , P.B. and xu, R.

J a f r i , A.K., Khawaja, D,K, and Qasira, S,Z,

j auncey , K.

Jauncey, K. and Ross , B.

1989 A b i l i t y of j u v e n i l e white s turgeon (AcJpenser t ransmontanus) t o u t i l i z e d i f f e r e n t ca rbohydra te s o u r c e s . ^, Nutr . 119: 727-733.

1964 S tud ies on the b ioc temica l compo­s i t i o n of some f reshwater f i s h . I . Muscles. F ish Technol. 1: 148-157.

1982 Carp (.2x2£iliii2. ca rp io) n u t r i t i o n - A Review. I n : Recent Advances i n Aquacu l tu re , Groom Helm, London ( e d s . Muir, J . P. and R . J . Robe r t s ) , pp . 215-263.

1982 A guide t o t i l a p i a feeds and feeding. I n s t i t u t e of Aquacu l tu re , Univer­s i t y of S t i r l i n g , S t i r l i n g , 111 p.

v,- 3S, c 11 ^ I %^.

^<' I :•

- 3 8 -

Jayaram, M, G. 1978 S tud i e s on a r t i f i c i a l on

J h i n g r a n , V, G,

J h i n g r a n , V,G, and P u l l i n , R.S.V.

Kaushlk, S . J , , Luquet, P . , Blanc, D. and Paba, A,

Kawai, S. and Ikeda, S.

L a l l , S.P,

Lee , D.J , and Putnam, G, B.

Lik imani , T,A, and Wilson, R. P,

the formulat ion of feeds and t h e i r e f f e c t

and Cvprinus c a r p i o ( L i n n , ) , M.F.Sc. T h e s i s , U n i v e r s i t y of A g r i c u l t u r a l s c i e n c e s . Bangalore , I n d i a ,

on the growth of Cat la ca t l i (Ham,), Labeo r o h i t a (Ham.)

1983 Fish and F i s h e r i e s of I n d i a . Hindustan Publ , Co, ( I n d i a ) , De lh i , 666 p ,

1988 A Hatchery Manual for Connraon, Chinese and I n d i a n major Carps , ICI> RM S tud ies and Reviews 1 1 , Manila, P h i l i p p i n e s , 191 p .

1989 Sttsdies on t h e n u t r i t i o n of Sit>erlan s t u rgeon , Aclpenser b a e r i , I , U t i l i z a t i o n of d i g e s t i b l e ca rbohydra tes by s tu rgeon , Aauacul ture 76i 97-107.

1971 S tud i e s on d i g e s t i v e enzymes of f i s h e s , I , Carbohydrates i n d i g e s t i v e organs of s e v e r a l f i s h e s . B u l l . i 2 £ , SQ£, S c i . Fish 3Jj 333-337.

1989 The m i n e r a l s . Inx Fish N u t r i ­t i o n , Second Ed i t ion , Academic P r e s s , New York (ed. Halver , J . E . ) pp. .219-257.

19 73 The response of rainbow t r o u t to va ry ing p r o t e i n / e n e r g y r a t i o s i n a t e s t d i e t , j j , Nutr.103 t 916-922.

1982 E f f e c t s of d i e t on l i p o g e n i c enzyme a c t i v i t i e s i n channel c a t f i s h h e p a t i c and ad ipose t i s s u e . iJ. Nutr . 112t 112-117.

-39-

Metaiier, R,, Dehapiot, T,, Huelvan, C. and Vendevilie, J.E,

Millikin, M.R.

Morita, K,, Furuichl, M, and Yone, Y.

NAS-NRC.

New, M. B,

1980 In f luence of the feeding l e v e l on growth, ffeed conver s ion , p r o t e i n e f f i c i e n c y and chemical composi t ion of Juven i l e European sea b.3ss (Dicent rachus l a b r a x ) . Proceedings of World Marl c u l t u r e S o c i e t y , i l l 436-444.

1982 Q u a l i t a t i v e and q u a n t i t a t i v e n u t r i e n t requ i rements of f i shes x A Review-. F i s h . B u l l . QQ,t 655-686.

1982 Ef fec t of carboxymethyl c e l l u l o s e supplemented t o d e x t r i n - c o n t a i n i n g d i e t on the growth and feed e f f i c i e n c y of red sea bream,Bul l . iIS£. Soc. S c i . F ish ^ t 1617-1620.

1981 N u t r i e n t Requirements of Coldwater F i s h e s . Na t iona l Academy of Sc iences Washington, D.C. 63 p .

1983 N u t r i e n t Requirements of WQrm-water Fishes and S h e l l f i s h e s , N a t i o n a l Academy of Sc iences , Washington, D.C, 102 p .

1986 Aquacul ture d i e t s of p o s t l a r v a l marine f i s h of the super - fami ly p e r c o i d a e , wi th s p e c i a l r e f e r ^ i c e t o sea b a s s , sea breams, groupers and y e l l o w t a i l : A Review. Kuwait B u l l e t i n of Marine Sc ience . J : 75-148.

Ogino, C. , Chiou, J . Y, and Takeuchi , T.

Oser, L. B.

1976 P r o t e i n n u t r i t i o n i n f i s h . VI. E f f e c t s of d i e t a r y energy sources on the u t i l i z a t i o n of p r o t e i n s by rainbow t r o u t and carp.Bu^^. i2a£. Soc. Sc i . Fish, ^ t 213-218.

1965 Inx H-awk's P h y s i o l o g i c a l Chemistry (ed. Oser, B . L . ) , 14th e d i t i o n . Tata McGraw-Hill Pub l i sh ing Co. , L t d . , New Delh i . 1472 p .

- 4 0 -

Page, J. W, a nd Andrews, J.W,

1973 In te rac t ions of d ie ta ry leve ls of prote in and energy on channel ca t f i sh ( Ic ta lurus punctatus) . J[- M t r . 102. : 1339-1346.

P h i l l i p s , A.M., J r . ,1966 Livingston, D, L. and Poston^ H.A.

Pieper,A, and Pfeffer, B.

Piper, R.G., McElwain, I . B . , Orme, L. E. , McCraren, J, P . , Fowler, L.G. and Leonard, J.R.

Renukaradhya , K.M, and varghese, T . j .

Ringrose, R, C.

Robinson, E, H. and Lovell , R.T,

The ef fec t of changes in protein q u a l i t y , ca lo r ic sources and ca lo r i c leve ls upon the growth aiv3 chemical composition of brook t r o u t . Fish-Res. Bull . NQ.. 21* State of New York Conservation Department, Albany.

19 79 carbohydrates as possible sources of d i e t a ry energy for rainbow t rou t (Salmo qa i rdner i . RicharxJson) In : Finfish Nutr i t ion and Fishfeed Technology, Vol, I , Heenemann, Ber l in , Federal Republic of Germany (eds, Halver, J ,B, and K. Tiews). pp. 209-219.

1980 Studies on the ef fec t of inc rea­sing proportions of sucrose or ge la t in ized s tarch in d i e t s for rainbow t rout {Salmo qairdneri R,) on the u t i l i z a t i o n of d ie ta ry energy t o protein. Acruaculture ^ : 333-342.

1989 Nutr i t ion and Feeding. In» Fish Hatchery Management, Department of I n t e r i o r U.S. Fish and Wild­l i f e Service, Washington D.C. pp. 208-262,

1986 Protein requirement of the carps, Catla cat la (Hamilton) and Labep rohi ta (Hamilton). Proc. Indian Acad, s c i . 2^» 103-107.

1971 Calor ie - to -pro te in r a t i o for brook t rout (Salvelinus fon t i -na l i s ) 2' Fish. Res. Bd. Can. 28 J1113-1117.

1984 Nutr i t ion and Feeding of Channel Catfish, Southern Cooperative Seriea Bul le t in No. 296, 56 p.

- 4 1 -

Sargen t , J , , Henderson, R , J , and Tocher, D. R,

Schmitz , 0 . , Gene l , E. a nd P fe f f e r , E.

Sen, P. R. , Rao, N.G. S, , Ghosh, S.R, and Rout, M,

1989 The l i p i d s . I n : Fish N u t r i t i o n , Second E d i t i o n , Academic P r e s s , New York. (ed. Halver , J . E . ) . pp. 153-218.

198 4 D i g e s t i b i l i t y of c rude p r o t e i n and o rgan ic m a t t e r of p o t e n t i a l sources of d i e t a r y p r o t e i n for e e l (Anouil la L . ) . Aquacul ture i i : 21-30.

1984 Obsejrvations on the p r o t e i n and carbohydra te requ i rements of c a r p s . Aquaculture 13i 245-255.

Shimeno, S, 1982 S t u d i e s on Carbohydrate Meta­bol ism i n F ish . Amerind P u b l i ­shing 123 p.

Co. Pvt . L t d . , New Delhi .

Shimeno, S. , Hidetsuyo, H., H i r a t a , H. and Takeda, M.

Shimeno, S , , Hosokawa, H« and Takeda, M.

Shimeno, S . , Hosokawa, H. , Takeda, M. a nd Kajiyama, H.

Shimeno, S . , Hosokaw, H, , Takeda, M., Takayama, S. , Fukui, A. , S a s a k i , H, and Kajiyama, H.

1977 Comparative s t u d i e s on caj±>o-hydra te metabolism of ye l low-t a i l and c a r p . B u l l , j a p . Soc. jSSi. F i sh . ^ 1 213-217.

1979 The importance of carbohydra te i n the d i e t of a ca rn ivorous f i s h . I n : F i n f i s h N u t r i t i o n and Fishfeed Technology, Vol. 1/ Heenemann, B e r l i n , Federa l Republ ic of Germany ( eds . Halver , J.E^ and K. Tiews) . pp. 127-143.

1980 E f f e c t s of c a l o r i e t o p r o t e i n r a t i o s i n formulated d i e t on the growth, feed convers ion and body composi t ion of young y e l l o w t a i l . B u l l , j a p . Soc. S c i . F i sh . 4 6 : 1083-1087.

1981 Adapta t ion of h e p a t o p a n c r e a t i c enzymes t o d i e t a r y carbohydra tes i n c a r p . Bu l l , j a p . Soc. Scd £Lsh' 42» 71-77.

- 4 2 -

Singh , B.N. , and Bhanot , K.K.

S i n h a , V.R. P.

1988 P r o t e i n r e q u i r e m e n t o f t h e f r y c a t la c a t la (Ham.) . I n : The F i r s t I n d i a n F i s h e r i e s Forum, p r o c e e d i n g s . A s i a n F i s h e r i e s S o c i e t y , I n d i a n Branch , Manga l o r e ( ed . J o s e p h , M.M.) p p . 7 7 - 7 8 .

1988 A q u a c u l t u r e R e s e a r c h and D e v e l o p ­ment i n I n d i a . I n t e r n a t i o n a l Symposium on A q u a c u l t u r e Needs f o r t h e y e a r 2000, New D e l h i , 15 -13 November, 1988 , 20 p .

Smi th , L . S .

Smi th , R.R.

1989 D i g e s t i v e f u n c t i o n s i n t e l e o s t f i s h e s . I n J F i s h N u t r i t i o n , second E d i t i o n , Academic P r e s s , New York (ed . H a l v e r , J . E. ) p p . 3 3 1 - 4 2 1 .

1989 N u t r i t i o n a l e n e r g e t i c s , Inx F i s h N u t r i t i o n , Second E d i t i o n , Academic P r e s s , New York (ed . H a l v e r , J . E . ) . pp . 1-29.

S n e d e c o r , G. W, and 1968 Cochran , W. G.

Spanhof, L. and 1977 Kunhe, H.

S t r a n g e , R . J . 1984

Swamy, D. N. , 1990 Mohanty, M.N. and T r i p a t h i , S.D.

T a k e u c h i , T, , Watanabe, T. and Ogino , C.

19 79

S t a t i s t i c a l Methods . Iowa Univ . P r e s s , Iowa, 593 p .

S t a t e

U n t e r s u c h u n g e n z u r Verwertung V e r s c h i e d e n e r F u t t e r m i s c h u n g e n d u r c h e u r o p a i s c h e Aa le ( A n q u i l l a a n q u i l l a ) . A r c h . T i e r e m . 27i 5 1 7 - 5 3 1 .

c a r b o h y d r a t e s . In» N u t r i t i o n and Feed ing o f Channe l C a t f i s h . S o u t h e r n C o o p e r a t i v e S e r i e s B u l l e t i n No. 296. pp . 1 5 - 1 6 .

D i e t a r y c a r b o h y d r a t e r e q u i r e m e n t s o f o a t la c a t la (Ham. ) f r y and f i n g e r l i n g s . The Second I n d i a n F i s l ' i e r i e s Forum, A s i a n F i s h e r i e s S o c i e t y , I n d i a n Branch . C o l l e g e o f F i s h e r i e s . Univ . o f A g r i c u l t u r e 1 S c i e n c e s , Manga l o r e , K a m a t a k a , 27 -31 May. 1990 , p . 80 ( A b s t . ) .

A v a i l a b i l i t y o f c a r b o h y d r a t e and l i p i d a s d i e t a r y e n e r g y s o u r c e f o r c a r p . B u l l , j a p . Soc. S c i . F i s h . 4 5 : 9 7 7 - 9 8 2 .

- 4 3 -

Tand le r , A. and 1980 Spec i f i c dynamic a c t i o n and d i e t Beamish, F. W.H, i n largemouth b a s s , Micropterus

salmonides (Lacepedae) . ^, Nutr . 110; 750-764.

T a r r , H . L . A , 1972 Enzymes and systems of i n t e r ­mediary metabolism. I n j P in f i sh N u t r i t i o n . F i r s t Ed i t i on . Aca­demic P r e s s , New York (ed. Halver , J . E . ) pp. 255-3 26.

T iemeie r , O.W., 1965 E f f e c t s of growth of f i n g e r l i n g Deyoe, C.W. channel c a t f i s h on d i e t s c o n t a i -and wearden, S. ning two energy and two p r o t e i n

l e v e l s . T rans . Kans. Acad. Sc i . ig. j 180-186.

T r i p a t h i , S.D, 1988 Freshwater Aquacul ture i n I n d i a . I n t e r n a t i o n a l Symposium on Aquaculture Needs for the Year 2000, New D e l h i , 15-18 November, 1988, 60 p .

Tung, p . -H. and 1989 E f f e c t s of meal frequency on the Shiau , S.-Y. growth, fteed e f f i c i e n c y and

carbohydra te metabol ic enzyme a c t i v i t i e s of t i l a p i a (Oreo-chromis n i l o t i c u s X Q, aureus) fed d i f f e r e n t carbohydra te d i e t s . The Second Asian F i s h e r i e s Forum, Tokyo, Japan, 17-22 A p r i l , 1989. p . 63 ( A b s t . ) .

Ufodike, B,B.C. 1983 Growth responses and n u t r i e n t and Matty, A . J , d i g e s t i b i l i t y i n m i r r o r ca rp

(Cvprinus ca rp jo ) fed d i f f e r e n t l e v e l s of cassava and r i c e . Aquacul ture ^ , 41-50.

V io la , S. and 1983 Eva lua t i on of d i f f e r e n t g r a i n s A r i e l i , Y. as b a s i c i n g r e d i e n t s i n complete

feeds for ca rp and t i l a p i a c u l ­t u r e Bamidoeh 35; 38-43 .

l^tendt, C. 1964 Diet and glycogen r e s e r v e s i n ha tche ry reared A t l a n t i c salmon d u r i n g d i f f e r e n t s e a s o n s . I , Winter. Swed. salmon Res. I n s t , , RPT. LFI. MEDD.

- 4 4 -

Wilson, R.P, and Poe, W.S,

1987 Apparent i n a b i l i t y of channel c a t f i s h t o u t i l i z e d i e t a r y mono- and d i s a c c h a r i d e s as energy s o u r c e s . ^ , Nutr . 117i 280-285.

Wilson, R.P. 1989

Z i e t l e r , M.H., 1984 Kr ichgessne r , M, and Schvrarz, F . J ,

Amino a c i d s and p r o t e i n s . I n i F ish N u t r i t i o n , Second E d i t i o n . Academic P r e s s , New York (ed. Halver , J . E , ) . pp. 111-151.

E f f e c t s of d i f f e r e n t p r o t e i n and energy s u p p l i e s on c a r c a s s composi t ion of ca rp (Cvprinus ca rp io L.) Aquacul ture 36t 37 -48 .