- ,2o.oltt. JO03 15:oo 6 Requirements

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,2o.oltt.
JO03 15:oo
6
.
J. Appl. Ichchyol. I1 (1995). 225-230
Q 19% Blackwell Wlssemcti=-Verlag,
ISSN0175-8659
Requirements
Recewed Onober 20,1994
Acrept& June 27,1994
Be&
for carotenoids in fish diets
BY 0. J. TORNSSEN and R. GLNSTIANSEN
Institute of Matim
Restdd,
Matre Aqracnhre Station, N-J198 Matredal, Norway
l
l
Carot.enoi& arc arqong kc most widespread and important pigment classesin living organisms,and in
qu.&e a&&, ZSIJJ.J&~P is rhe mosf COInmody
oc~rriog red cuounoid. &mamds are vitamin A
~~~~~~~~md UC fund~ad
in pbotorynrhuis and Lightprorection in phuu. Increasing attention has
Eeen drawn to a possible light proucuon. cancerpreventronand immune enhancementbycarotenoids in
manmA. reported functions in fish r~gc from a general cnhancancnc of pcrforrnsncc 10 specific
fincdons in reproductioo and meubolm. In ibis paper, WC show rhat vtwvltin
is essenualfor
growth and rwivd of fish and crdish,
and discuss this fact in relation to the use of purified and
semipurified &a~ in nnvlriond sr~dies. The simihrir)r rn action of arraxanchin and caorhaxanrhb
E).d.retinol
(4umin A) su~ests that tbesc W.-Ocuorcnoids
corn ored to a-rocophcrol (vi-in
sho Is d be krccd
.
among the fat soluble vnam~~.
Introduction
l
l
I)
Carotenoids in modern aquaculture are mainly associated with astaxanthin or canthaxanrhin
pigmentation of rhe flesh of salmonids. The pigrnentarion of Adanric salmon (S&JO suhzr)
and rainbow trout (Oncurh~ch~
my&s) flesh is regarded a~ the most important quahry
criteria next to pro&r
freshness (KOTENG 1992). It is therefore of vital interest for the
salmon farmer co achieve a satisfactory pigmentation of the salmon flesh. The market
demand for asuxanrhin in A&nr.ic salmon flesh requires a concentration above 6-7 mg per
kg. Factors influencing the absorption and deposition of carotenolds are reviewed by
TORRISSEN
CC
al. (1989) and STOMBAKJLEN and No (1992).
it has been shown that carorenoids have at least four functions: (I) accessory pigment in
photosymhesis, (2) protective pigment againsr photoseniitization, (3) provitamin A source
and (4) communication in aquak ank&
Responses to physiological or pharmacological
adm.in,istration of carorenoids +re nordly
classified as actions.
Potential mechanisms
associated with carorcnoid ations include antioxidant and singlet oxygen quenching, proGamin A activity, up-regulation of DNA expression, co-oxidation, and enhancement of
immune functions associated -4th increased turnour immunity and modulation of macrophage and lymphocyte acdwion (BENDICH 1993).
Salmonids absorb and deposit aswanthin and cancbaxanthin in rhe muscle during the
grow out period. Ar the ame of sexual maturation, They mobilize the cruorenoid srore
and transport the accumulated vtuvlchin
or canthz~nthin by the Very High Density
Lipoproteins (VHDL) (virehogenin) or High Density Lipoproteins (HDL) to rhe ovaries
and finaJy the progeny. This active transfer of cuorenoids from the mother fish to the eggs
has led to the hypothesis that urotenoids are vital for egg and larval development.
Absorption
3
Some carotenoids are absorbed well and rapidly metabolized, some are absorbed and poorly
metabolized aud some seems nor 10 be absorbed at alI. Both the absorption and metabolism
of carorenoids are highly dependent on its structure. The mechanism by which the carotenoids are taken up from the plasma by tissues and released back into plasma from specific
U. 5. Copyfight Cliche
3
and storage
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0.1. 7omsm1 and R Cbrisriansen
226
v
Absorbed
Carctenoid
Al
Growth
Q
s
Astaxanthin
Jp++&-
l *w
A2
100
80
Ye0
1
Al LA2
60
/
Canrhaxanthin
-0
0
/
Survival L%S/
“pa
Ye.5
Al LA2
YtX
Al
40t
I
20
LA2
Bcuodn
mT”a
lsozeuanthin
*
les
Al
Y&9
A2
YeS
A2
%
6(
-
Fgg. 1. lain urorenoi& used m fish IUJCI%O~and chcir SUUCWN; Al = minol; A2 = dehydroretinol;
3, ++a, =**T+ ind&ta m+wde
of difference in absorb&n
a
0
ciss~es, however, has not yet been clarified (OLSON 1993). A preferred absorption and
deposirion of hydroxy and kero caroeeooids is seen in fish. Astaxanthin and canrhananthin
are absorbed WC& whiJe B-carotene in some fishes seems hardly to be absorbed at all.
Growth
and survival
An &proved growth of Atlantic salmon was found by supplementing commercial star-~
(Fig. 1) (TORRISSEN 1984), and no significant
fee&g diets with astaxanth or ~ahrhur~lrhin
differences were found bcmvccn the asuw-u.hin and canthaxamhin supplemented diets.
Corresponding results are found’for the red tilapia (07eochromis dozincs) (BOONand UNPRASERT 198% and kuruma shrimp (PLMPHS jcrponicrts) (CHEN and
YARATPALIN
JENG I 992). Supplemcnurion of fl-ororene and canthaxanthin to the diets of major Indian
0~s resulted in a better swivel and growth compared KO conventional diets used arithour
CSUOXCW~~J
(GOSWAMI 1993), and N~GERE-SADARGUES et al. (1993) found a higher survivzl
rate for Pemerts juponicn§ receiving aSWan&-cagth=anIlchin
supplementation (50/50),
but no differences were observed in groMh and moulting.
CHRISTI~NSEN er 11. (1993) investigated tbc interaction between ascaxanchin and vir.am.in
A supplementation on 80 wth and survival in first feeding fry of Atlantic salmon. The
experimental diets were based on a semipurified diet based on vitamin and carotenoid free
casein and gelatine as protein sources developed by SHXARER et al. (1993). The results from
&is IX-day feeding study clearly showed a significandy improved growth and sun&al 00.
supplementation of osraxanrhin to the expetinul
diet (Figs2 and 3), and vitamin A
supplemenrarion alone did not suppo~ growth and survival. The vitamin
A r~urcc ured
was a mixture of retinal pal&ate and retinol acetate The bio-availability of rhe two forms
is nor known and I
also show a provi
expIain the effect c
GROS and BUDO’
addition co fi-cuot
plaKies (X+7phon
both A, and A2 in
highly dependent 3
(1985) reponed th;
Ain theintd*
burrdchws, l&n
(:
(dehy&or&ol)
(l
tilapia (Tikapiu nil
converted into vira
Fishes change the
responses during 4
as compromises ba
the ‘need’ to avoi
changes is comple;
Visual cues. It has
and xanthophors
(1952) suggested
pigments may hav
227
Fig. 2. Sutvival of A&n& salmon fed diets
supplemented with different IevcL of
aaxanrhin and vitamin A (CHRIFILANSEN
et al. 1993)
is not known and one or both might be poorly utilized by start-feeding salmon. The results
&O show a proviramin A function of astaxanti
(Fig.4), but this done is not able to
explain the effect of asuxnnthin supplementltion.
Functions of carotenoids
Provitamin A
9
a
GROS md B~OVSKI (1966) reponed that asuxanch;l, canrhuannrhinand isozeaxanthin in
ad&&on ~0 p-carotene were precursors for vitamin A in both guppies (Lebirtes reticdatw) and
pb&s (.+hqph~ms ursiaws). Ascam&
c~lrhulnrhin and ~eamnthin were precursors of
bo& A, and A, in rainbow UOUK (~COS~~C~US vkirs), bur the rate of incorporation was
&$-JY dependent on’ fish size and age, and the vitamin A status of the fish. SCH~EDT et al.
(1985) rep0J-t.d &at utaxanchin, c~rthwanthin and zuxanthin were transformed into vitamin
and CL&s
A in he tips&al wall and the liver. In the freshwater fishes,hxubrancbwfosdri
brhnrhus, lutein ($3’ Dihydroxy a-carotene) is reported to be the precursor of vitamin A2
(dehydrorerinol) (Bmoa and &SWAMI 1977; Gosw~
and BHAITACHARJEE 1982), and in
zawxhin,
k.h
and ~naxanrhin were directly bio&pi4 (7&p
7dmif) astaxatin,
convened into vitamin A(J) (UTmN
ad ~TSUNO 1988).
Communication
Fishes change rhcir hues in response to background colouration and also display colour
responses during excitement and courmhip (FUJI1 1969). The colour pattern can be viewed
1s compromises between the ‘need’ to communicate 4th other members of the species and
the ‘need’ to avoid being eaten (MOYLE and CEW 1982). The internal control of colour
changes is complex and involves both hormones and nerves where the initiation comes from
visual cues. It has been shown rhat carorenoids are intcgrll constituents of chromacophores
and xanthophores, and, as such,. are functional in the photo-responses of fish. GOODWIN
(1952) suggested this to be the major role of wotcnoids in fish, and lack of suticienr:
pigments may have a negative effect on their general performmce.
1
Nr.Yd4Y
S.
db/4b
O.J. Tomsen and R. Cbrimanscn
228
-
B
I
w
5
p
0 N vir A
4
-0IuvirA
3
-200OlUVi~A
2
-4OOONriLA
1
0
Table 1. For-mu
and vitamin A
lngcclicnrs
C~Citl
Fi boil (San
G&ill
Dcxuin
0
10
20
30
40
50
60
70
SO
90
100 110
17~3 130 I&I
Cuboxymc
.-4JUlOSe
a
F&g.3. &JW&
of Arlanric
S&IOII fry fed die= supplemented with different conceonrmtionsof vipmk
A m d ur-&n.
TU=titemational
tits; (a) Fish fed set&purified dlea free of Aaaan&n;
(b)
Semi-purified diets supplemented wirh 20 ppm Aswrurthio; (c) Semi-purified diers supplementidwi&
4opp
As~thin(CHP~~NS~e~~l.1992)
Actions of carotcnoids
3)
DANGER
(1984) did not detect any effect of a~taxanthin supplementation on f-&v
of r&bow
FLOUT,however, improved egg buoyancy was observed from red sea bream
(chvsoph,-p mjor) broodstock fed diers containing jkarorene, canthanadin and as-an&in rhc night before spawning. Hatching was not affected, but the number of oil globules
was reduced. C~TLAMEN
and ~OIWS.EN
(unpublithed dara) .did not find any effect of
salmon eggs.
vwanch,in content on the hatching SUCCESS of Athic
Herercpne*srez fossh showed auophied gonads with damaged germinal cpithclium when
fed a carorenoid-free diet (GO~AMI 1988). SINGER et al. (1989) reported an improved liver
histology ~JJoreochromis nilortir
and Co&a lrrbioJa fed high astaxanthin 1~~1s (71132 g/kg) and IOW level (32 mgikg) in the diet. Pax-&ularly rhe parenchymal and intracellular
organization was better developed. III rilapk the glycogen storage was enhanced and &e
cell volume s~@~Itly inaalthough the biochemical mtihanism is unknown.
Marine pelagic cold water fish spawn large numbers of small eggs without visible caretend
depositions, while demersal fish and viviparous fish often have eggs con&fig
high
~CVC~S
of carotenoids. Pelagic eggs have, in general, a shorr period of development from
Amino acids
L-Ug
L&S
L-lyr
L-met
k:;b
‘The iamin
vitamin D,; 1t
riboflavin; 1s
acid; 40 mg bi
mVkg dry ing
NaHSeO,;4.c
spawning unt
satisfies their
may be an ad
halibur and tu
predominant 1
supplies of vit
that vitamin A
pnmatns). Tb
.
Requvemenrr for carormoids m fish drprr
229
Fig. 4. Vitamin A content of Aclan-
tic salmon fry fed diet.3 supplemcnced with vitamin A ud
ascexoncllin
T&& 1, ~o~u~ntion
of a semipurified test diet for Atlantic salmon. Concentrationr ot astaxanthin
2nd vitamin A mg/g dry diet AU other ingrcdicntr are given as percentage (SHEAMR et J. 1993)
0
Ingedicncs
casein
Fishoil (Sardine)
G&till
Dentin
Carboxyrymerbylcehlosc
a-cellulose
3
Amino acids
kz
L-lyr
L-met
44.8
17.1
10.0
12.0
1.0
4.6
1.0
0.2
1.0
a.4
0.5
1.0
Viwnin mis
VitaminA~
As-clun
kg/g-J
Choline Chloride
KC1
NOCI
CaHPO, y H,O
MgO
TraceminsoL2
2.0
~,600,1200
0,20, 40
1.0
1.5
0.3
12
0.3
1The &min m;r contained
the following viwnins per kg of diet: 1.188 g a~corblc acid; 4 mg
vica& D,; 100 mg uYtoc?pherol acetate;6 mg ritnmin K,; 15 mg thinti hydrochloride; 30 mg
riboflavin; IS mg pyrldome hyhodrloride; 453 mg C&WI
D-phantothenar; 150 mg nicorinic
acid; 40 mg &oh (2%); 4 m.g fdc acid; 3 mg G.amin B,, (1%); 300 mg inosirol: ‘Supplied a 100
ml/kg dry ingredients cools): 1.9 mg Kk 32.5 mg MnSO. I&O; 88.0 mg Zn!iO. 7H,O; 4.2 mg
NaHScO,; 4.0 c&l, 6I-W; Il.8 mg hS0,5H,O
spawning until the progeny start exogenous feeding (phyto- and zooplankcon) which
satisfies heir urorenoid requirement Tbe lack of visible cuotenoids in transparent eggs
may be an adaptation 10 minimize predation pressure. Abnormal skin pigmentation of
halibut and arbor is a large problem in first feeding of larvae on rotifers and utcmir The
predominant hypothesis is that unpigmenred skin is a deficiency syndrome for insufficient
supplies of vknin A and highly unsaturated fatty acids (HUFA). It has also been sbowr,
that vitamin A deficiency &uses a depigmentation of the skin of channel catfish (Zcr&r~s
punrtor~). xs deficiency syndrome seems to be reduced by enriching the Jive food with
L
,?O.Okt. 2003 15110
Nr-9349
0
I
0.1 Totisen and R Chnr~unren
233
urotenoid
conri&g
for A&,&
salmon
fish larvae require
S. 38/46
J. Appl. Ichchyol. I
Q 1995 B1.&-cU ’
algae. We have shown that vitamin A esters have a limited availability
ft-y (CHRISTIANSEN et aI. 1993). and pilot studies indicate char marine
carotcnoids 01 preferably astaxanthin.
IsSN
Diet supplementation
l
0175-8659
The applk
I[ is sbown h a series of mvesrigations on a wide varict)r of fishes and crayfish chat
supplemen&on
of cnrotenoids to the diet improyc~ grow&, reduces the mordity race and
enbcces +,e general performance of the anid.
In addition a large amount of cmpnical
dam suggests &,t a sufficient carotenoid supply is essentiai for the well-being of the animal.
Asuxanth
or ~~rhaxanchin should be regarded as a vitamin for fish and crawfish and
&&d COau fish diets at a levd above lomgkg’ dry diet.
By C. (
Depammnr of
‘Depanmcnc
of.
References
BARU~,,A.B.;
GOSWAMI,
U-C..
1977: Formaoon
of Vitamin A 10a freshwater fish. Isolation of rcdnoic
ncld. Biochem. J. 164: 133-136.
l
EE~~cH, A., 1993; Recent adyaoccsin chnical researchlnvohing
cuorenoids. 10th Int Symp. Croten&& rrondheim, 20-25 June, 1993,Book of AbsVaN, SL-9.
BooN~AIIZ(TP,~J,~, M.; UNPRASERT, N, 1989: Effect of pigments from different source6 on colour
chan m and grow& of red Oreocbromi nilonrur. Aquaculo~re 79: 375-380.
CHIEN 8! ,H., . J&NC, S.C., 1992: Pi menurion of kurum praan, Pmaru ~“pon~curBate, by wious
pi&em sourcea end levels and kecdiog regimes. Aquaculcure 102: 33-346.
C~ANSEN,
R.; LIS, 0.; TORRISSEN,O.!, 1993: Effect of vtvanthin and vitamin A on growth and
sur&d during first feeding of ArJantlc salmon Saimo saiar L. Aquaculture end Fisheries Menogement (in rcrs).
FUJII, R., 1969: e bromarophorcs and pigmenrr In: HOW W.S.; RANDALL, D.J. (eds),Fish Physiology,
Biochenustry of the Cuoceno&. Chapman and Hall, London.
diet and irJ role in re reduction of fresh water f&h. 8* ht.
Sp~p. Carounoids,
3
Boston, 27-31 July, 1?88,.Absuacts of F resentzdons 32.
Cosv.+& U.C., 1993: Meubohsm of carorcnolds in freshwar.erfish: (i) Biogencris of 3-4 dehydrorerinol
(ii) rupplemencation of carotenords Mrh fish food for better eur~l~ol and growth. 10” Ins. Symp.
Cuorcnoids, Trondhelm, 20-25 June 1993, Book of Abstracts, CL 10-7.
GoswAMI, U. C.; BHATTACHAXJEE,
S., 1982: Bios n&sir of 3-dehydroretinol: metabolism of lutein
(fi-carotene-3,~’ Dial) in Clrt;s barrachesand Ompokpabo. Biochem. Inc. 5: 545-552.
Gross, J,; p. BuDOVWJ, 1966: Cosw~sion of cvoccnoids UIIO Viramin A, and A, in rwo species of
fr&watm f;& Biochem. J. 101: 747-754.
KAI~IJYAMA, M.; T. J~~ATsUNO, 1988: Carotenoid and vitamin A, and metabolism of cerorcooids, betacsrotenc can&uanthin,
ascaxanmrhin,zcasanthin. hem
and mnaxanrhin in rilapia T&pi0 nilorica.
Camp. Biochem. Physiol. 90Bz 131-139.
KOTENG, D.F., 1992: Markedsundcrsekclse norsk k&s. FNL, Bs en.
R, 1982 Fishes; or Lnvoduction to Ich ti yology, pp. 162-168.Prcncice-Hal1
Mo&f: ~f~~~r!kds,
New Jersey.
N$GRE-SADARGUES, G.; CxnUO,, R.; PE’ITI, H.; SAXE,
5.; GOMYly R; MILICU$ J.C.; CHOUBERT,
G.; TRIUES, J.P., 1993: Udl~s~on of aynrheoc carorenoids by she pawn &ZUeJ#J japonirvs under
cx crime&
rearing condid~~~
lO* Int Symp. Carotcnoids, Trondheim 20-25 June 1993, Book
o P Abstracts, CL 10-3.
OLSON, J. A., 1993: Absorption,
tw~spor~ and metabolism of carorcnoids in humans. 10th Inr. Symp.
20-25 June 1993, Book of Abstncrs, SL 3.
SC~EDT K. L~uENBERGEX,
F.J: VECCHI, M.; GLI
E , 1985: Absorption retention and mcrebohc
~KUXG&OUS
of caroCenoLcLin rsinbo~ trout !3rn. on and ducken. Pure and Applied Chemistry
Carotenoids, Trondhdq
57: 685-692.
The measurement I
enabled the develo
of individually nun
to sranderd nut&c
experiment Analy:
consumption-grow
differences in apt
establish coosump
consumption rates
factors, such cs rbs
end protein curnow
between fish in rhc
compare rbe growr
The mcasurcmer.
graphic method (
several aspects 0
and JOBLING 199
(e.g. CARTER et
used to m-we
growth perform:
odology, accurac
(MCCARTXY er a
individual CON,,
compare the grc
chareccerinics. A
nutrition trials w
laboratory.
.
of rainbow trout. Agumhre
-effect
100: 209429.
of carorenowzls in eggs and star&e&g
dies
nutitional
st
importance chat
calculate consun
diets. For examp
may result in vari
In
U. S. Copyright Cl