«
CFllVE
FISHERIES AND MARINE SERVICE
Translation Series No. 3432
Fish Lipids. II. Seasonal dynamics of phosphatides and
triglycerides of large cisco Coregonus albula L.
By E.I. Lizenko, V.S. Sidorov, and 0.1. Potapova
Original title: Lipidy Ryb. II. Sezonnaya Dinamika Fosfatidov
Triglitseridov Krupnoi Ryapushki Coregonus Albula L
From: Lososevye (Salmonidae) Karelii (1): 164-169, 1972
Translated by the Translation Bureau(GAD)
Multilingual Services Division
Department of the Secretary of State of Canada
Department of the Environment
Fisheries and Marine Service
Halifax Laboratory
Halifax, N.S.
1975
14 pages typescript
SECRÉTARIAT D'ÉTAT
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E.I. Lizenko et al.
TITLE IN ENGLISH — TITRE ANGLAIS
FISH LIPIDS. II. SEASONAL DYNAMICS OF PHOSPHATIDES AND
TRIGLYCERIDES OF LARGE CISCO COREGONUS ALBULA L.
TITLE IN FOREIGN LANGUAGE (TRANSLITERATE FOREIGN CHARACTERS)
TITRE EN LANGUE ÉTRANGÉRE (TRANSCRIRE EN CARACTÈRES ROMAINS)
LIPIDY RYB. II. SEZUNNAYA DINAMIKA FOSFATIDOV I TRIGLITSERIDOV
KRUPNOI RYAPUSHKI ODREGONUS ALBULA L.
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Lososevye (Salmonidae) Karelii.
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Salmonids of Karelia.
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pp 164 - 169
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Lososevye (Salmonidae) Karelii, No. 1, 1972, pp. 164-169
FISH LIPIDS.
II.
SEASONAL DYNAMICS OF PHOSPHATIDES AND
TRIGLYCERIDES OF LARGE CISCO COREGONUS ALBULA L; T ED 7RAW;L%";MN
U,u
,ftr.),XTICII NOW ;I:C;Pf., 1117
By
•
se:uk)r-*:
E.I. Lizenko, V.S. Sidorov, 0.1. Potapova
(Biochemical Laboratory and Laboratory of the ecology of fishes and
mariné inVértebratés ef -the'InstitUte of Sioldgiof the Karelian .
of the Academy of Sciences of the USSR).
Branch
/164*
There is presently known a considerable number of studies in
which the seasonal dynamics of the fat content in some tissues of fish
have been established. It has been found that a marked accumulation
of fats in the foraging period and consumption of fats in the period
of intensive maturation of gonads and the spawning migration is
characteristic for many species of fish (Shurman, 1960).
However,
the majority of studies are devoted to the study of the seasonal
content only of total lipids, moreover, as a rule, only in two or
three organs (muscles, liver, gonads).
The number of studies in which
the dynamics of individual groups of lipids - phosphatides, triglycerides,
cholesterol and its esters, and others, would be elucidated, is small.
Thus, in particular, Vada (1955) found that the content of phospholipids,
*Numbers in the right-hand margin indicate the corresponding page numbers
in the original text. - Transi.
$03.400-.10-at
2
triglycerides,'hydrocarbons and other unsaponifiable lipids increases
in the liver in sardines during the process of maturation and preparation
for spawning, and afterwards, immediately before spawning, the content
of triglycerides begins to decrease, while the remaining lipids are
preserved approximately at their previous level.
After spawning, the
amount of triglycerides again increases rapidly (cited from the Journal
of Abstracts "Biokhimiya" (Biochemistry) 1956).
It has been established
(Musacchia and others, 1957), that the content of different lipids
varies with the seasons in 26 species of fish, and that a relatively
high content of phospholipids is usually mantained in the liver of
fish (Musacchia and others, 1957). It is. also known that considerable
seasonal fluctuations of total fat in the internal organs and the
abdomen are observed in the European cisco Coregonus albula L. (Luhman,
1.955; Morawa, 1956).
The study of K.I. Belyaeva and V.V. Pokrovskii
(1958) cites data on the content offat, protein, ash and moisture in
the body of the Lizhmozero and Chuzhmozero ciscos (four specimens each).
The results of the study of total fat content of foraging and spawning
ciscos in Munozero and Urosozero in 1964-1965 are presented in the
studies of 0.1. Potapova and V.F. Titova (1967, 1968).
The authors of these papers have shown that change in total
fatness of the body and gonads of the cisco is closely connected with
the maturation of the gonads. The maximal content of fat is observed
in August during the foraging period (III stage of maturity of gonads),
and minimal fat content is observed in October, in the prespawning
period (IV and V stages of maturity of gonads).
3
The present article is a result of a further deepening of
investigations of the seasonal dynamics of total lipids, phospholipids,
and triglycerides (Lizenko and others, 1969).
Material and Methods
For the comparative characteristics of the lipid composition
of the cisco, material was gathered in the foraging period (June,
August), prespawning period (October) in 1968 and at the end of the
wintering period (March-April) 1969.
Females maturing for the first
time and females spawning repeatedly with an identical stage of
maturity of sexual products were analyzed.
The size, weight and
inecators_of . fish analyzed for lipid
_ composition are .given
in Table 1.
The samples for biochemical analysis were taken from
net catches, and the freshest specimens were selected. Each cisco
was prepared for 13 organs (the liver, gonads, kidneys, heart, spleen,
brain, spinal cord, lungs, air bladder, skin, scales, adipose fin, and
internal fat). The meàian samples from eight specimens, each weighing
1.0-1.5 grams, were fixed in a 15-fold volume of a mixture of
chloroform and methanol (2:1) and were stored in tightly covered
test tubes at 3 - 4 ° C. The further isolation of lipids and their
qualitative and quantitative analysis were done following the
prescriptions given in another article in this same collection
(Sidirov, Lizenko, Nefedova, Bolgova).
4
Results and their Discussion
Given in Table 2 are data on the content of total lipids in
the organs and some of the tissues of the Urosozero cisco in
different seasons of the year.
It is clearly visible that the content
of total lipids reaches its highest values in the summer (August) in
the foraging period of the fish, in practically all organs, with
the exception of the liver and skin. The least amount of lipids
'
in these organs is observed during the spawning period (October),
and
in some cases, their reduction becomes substantial. For example,
the amount of total lipids in the muscles in October is reduced by
comparison with August by 33%, by 22% in the heart, by 28% in the
..., ...kidneys, by 19%.in the spleen, by 35% in the lungs, by 40% in the
air bladder, and by 40% in the gonads. By March and beginning of
June, the content of total fat in different organs off the cisco again
increases, reaching in many cases values which are Intermediate between
values in August and those in October.
The most likely explanation
of the marked decrease in the content of total lipids in the cisco in
the prespawning period is, in our opinion, the consumption of lipids
for the intensive maturation of the gonads which takes place in this
period and which is accompanied by considerable increase in their total
weight. Thus the coefficient of maturity (the ratio of the weight
of the gonads to the total weight of the fish, expressed in %) increases
in the period from August to October by 5 - 6 times (from 3.0 to 17.0 %).
Reduction in the content of total lipids in gonads by October can be
explained by the fact that during the period of transition of fish from
the III to IV stages of maturity, the intensiveness of biosynthesis
5
Table 1
Biological indicators of Urosozero cisco females
2 '.
1,
1101:33.U0.111
1-5
•
• •
9-23
E13111111
anrycra
19E8 r4..8 aRa.
3
•
,
.T.(nlitia Teaa no Cmitrry. (ac) , cm
.
Bec, r
.
.-
21,2
95,5
55,0 -130,0
'71
0,01-0,110
ioJ 0.06
0.01 -0,104
0,07
0.170 -1,350
, 0,705
0,-7 1,12
0,84 ..
0,81 0,93
to7
,
.. ' 1.7
Bec' nomi, r
1.
aipa . r
Bec nnyrpentiero z,
-
18.3-2o,1
--
11. Ko3(pcpututetir •;«,tpilocru
••
12. Bee runaa
13 ..Ko -,(InImittneur
apcabcric
'
14. KomIximinaeu r
ynItrantrocrn
Kaapc
15 .1.1oa I, craamn 3pe.locrii
20.0-23,7
9 1,9
89-150
62047
.
.
mousi
19E8 r., ii.9ua.
Bpemsi
-
nu
•
97,5
75-130
99,9 .
0,20-0,95
0,32
npô5
4-22 oramapn mapr-aureab
r , 14
191-4 r., 7 3K3. 19p 6
.
;
6
.
22,0-2(3,3
14,8-22,5
18,4 .
• . 23,9
32-108
112-150,5
65,3 .
167 .
28-97
85 -197
57,8
130,2
0,050 0.491
0
. -' 0,81
,.•.
0,18-0,84
.
. 0,37
0,8 -7,8
• 9.8-33.15
202
'3,1
6,01-17.4
0,9 5,0
• 2,6
-13.7
0,79-1.05
0.85--1,09
10,
(.491
IST..2: IV-V
e:, Ili,
? II, T. I1-111
•
•
• 0) 11 DI' •
- - :
. .,
0A6 .
.0.1 3,1
0 664
0.08-1.52
0.75-'1.00
0,85
2 11,.7. II-III
«
--r-.--.1
Key:
1- indicators; 2- time at which sample was taken; 3- 1 - 5 June, 1968,
8 specimens; 4- 20 - 23 August, 1968, 8 specimens; 5 - 4 - 22 October,
1968, 7 specimens; 6- March-April, 1969, 14 specimens; 7- median length
(from the anterior limit of the snout to the end of the middle rays
of the tail fin), cm; 8- weight, grams; 9- weight of kidney, grams;
10- weight of internal fat, grams; 11- coefficient of fatness;
12- weight of gonads; 13- coefficient of maturity; 14- Clark's
condition factor; 15- sex and stage of maturity.
Note: Given in the numerator are fluctuations, and average indicators
are given in the denominator.
;'.
6
Table 2
Content of total fat in organs of the Urosozero cisco
in % of dry weight
2.
1.
o p ra ltIA n
11: ■ 11111
rituttb
Brem% c5npa ta -reiztda.1
arnIcr
3.
7. 1b111.111,b1
8. fleeub
9. Cepaue
10. licenut
11. Ceae3enKa
12. )Katipbt
13. Iliaaa -reàbInefi ny34t,
14. Kom
15. roioauo mar.
16.. 1Crutiniori bt 03 1i
'17; . ibKuptisoil
18. lif1iiï«-rpetiniift }Km)?
19 • irolia,abi
20 •lu!,eillysi
•
15,3
13,5
90,0 .
22,0
•21,6
22,1
30,4
33,2
957 •
85,0
33,9
r.
27,7
959
-
99,1 .
38A
18,1'
57,0
'89,0
40,1
4959 r.
6
.5_
4.
90,0
50,2
66,0
OF:Ter b
end r.
1963 r.
:
- I
10.2
12.7
99.4
21,9 '
20A
99.5 '
99.ty
9.).8
•20A .
9a.4
188
20.n16,2 .
52,1
*
OA
24,2
3,9
.
77,7
• 3:14
.
,
9.3
Key:
1- organs and tissues; 2- time of collection of material; 3- June, 1968;
4- August, 1968; 5- October 1968; 6- March-April 1969; 7- muscles;
8- liver; 9- heart; 10- kidneys; 11- spleen; 12- lungs; 13- air bladder;
14- skin; 15- brain ; 16- spinal cord; 17- adipose fin; 18- internal
fat; 19- gonads; 20- scales.
(or accumulation) in the gonads of substances of a nonlipid nature -
proteins, nucleic acids, polysaccharides and others, markedly exceeds
the rate of *accumulation of lipids. It should be noted, that a
similar reduction in the share of lipids in gonads in the transition
from the 111 to IV stages of maturity was observed by 0.1. Potapova and
7
B.F. Titova (1968) in the Urosozero and Munozero cisco in the spawning
period of 1964. They established that, on the average, fatness of
gonads decreases from August to October in the Urosozero cisco from
15.5 to 12.0 (that is by 23%), and in the Munozero cisco from 20.0 to
18.2% (that is by 11%).
Presented in Tables 3, 4 and 5 are data on the seasonal dynamics
of phospholipids, triglycerides and the ratio of the first to the second
in different tissues and organs of the cisco. It can clearly be seen
that while the content of phospholipids in different seasons of the year
for most tissues practically remains more or less constant, the content
of triglycerides is subject to marked fluctuations. In fact, the
^ gréàtest Ariiount of `triblycérides, ^ as -welh =as -of -total lipids, is observed in all organs in August, while the least content is observed in October.
Thus, reduction in the content of total lipids from August to
October is explained primarily by the consumption of triglycerides
which are reserve lipids. Very often the numerical indicators of the
reduction of total lipids and triglycerides are close in value. For
example, in muscles, the content of total lipids from August to September
fell by 5.1% (% of dry weight), and that of triglycerides by 4.1%,
while the corresponding indicators for kidneys were equal to 7.7 and
7.0, the spleen - 4.8 and 6.1, the lungs - 10.3 and 10.9, the air
bladder - 5.0 and 6.4, the brain - 4.9 and 3.3, the spinal cord 5.3 and 4.3%. The insignificant amount of triglycerides (3 - 5% of dry
weight) in the brain and spinal cord attract attention. As is known,
in mammals, these tissues contain practically no triglycerides (Shtraub, 1963;
8
Table 3
The content of phospholipids and other complex
lipids in the organs of the Urosozero cisco,
in % of dry weight
Ftpe—mli
• 1
.. Oriitu n Tvamr
7.; Mietuubi
8. flown
9 .; cepAge
lu notiNit
11.1 Certe3emm
12.
13.
Wa6pie •
11.-umaTe.1btima
y3b1 p b
14.; Koy-M
iu m n . U63 r.
10,1
15,4
17,1
17,5
14,5
19,1
22.4
••••■•
•
r000snon. mo3r•
Crojuttiet m or
17. >Kupbooil naaoolo:
Burpeunitii ;Tow
19,2 rmmAm
20-;
41,0
onr...cr
W18 r.
9,0
15,9
19.6
11,5
16,1
15.4
24,7
9.5
43,3
57,5.
36 0
22,0
37,3
25.0
starernaaa
CItirierf.
Ih r.
9,2
1:3,3
13.5 •
13,0
13.3
15,2
13,6
15,3
47,0
, 57,5 .
•
11.7
0.0
14,7
3,5
9,2
16,7
12,6 :
16.4
/1,9
15.0
•
21,4
7.3
3'3,2
47.0
34.6
21,1
1,4
Key:
1- organs and tissues; 2- time of collection of material; 3- June, 1968;
4- August, 1968; 5- October 1968; 6- March.-April 1969; 7- muscles;
8- liver; 9- heart; 10- kidneys; 11- spleen; 12- lungs; 13- air bladder;
14- skin; 15- brains; 16- spinal cord; 17- adipose fin; 18- internal
fat; 19- gonads; 20- scales.
9
Table 4
The content of triglycerides in the organs of the
Urosozero cisco in different seasons of the year,
in % of total weight
.
1.
•
.
.
19:a r.
•
7.1quwum
8 .11enes4b
•
9 •Cep;ute
10.110,115,1
•.
••
11 .Cc.u.s3mirt
•
.•
12. .;Ka6pbt
13 .11:1a s3a.reablibirt n.9bipt, • '
14. Rdzw
r
18 1. Bitp pelinstit
1
191. 1-0sta:1m
_
20 .1. 1 eunt si
-
np
•
siseycr
•
'3.
••.- •
2J6
34
3 j ,
•
32
3.3
8,9
7,0. '
•:
J...1'0.1outiors mo3r .. •
I
-Lo:•Cissusitors. .r om'. .
„1
/ i•?1,*siponoft naatimiK
Bposs can •a
• 2:
••
'
• 1.fit. •
O vra si m i s 1• 19!
.. 4.0
•
•
.• osmsapt,
19E8 r,
4.
• 0,6•1
.4,7
• 3,9
9,3
3,5
' 1,8
8,9
97,
1,2
, 13,2
10,6
7.5
91,3
. 8.0
4,8
•
••
• - 44,0 '
•
10.0
48.0
15,3 •
6. _
2,1
2,8 3,5
4,6 .
4.0
.
47
tO
4,7
5,9
6,9
7,5
5.3
0,02
7,8
-
. 193i r.
0,0 .
10,6 •
39„:1 PA •
•
Isfn
1— organs and tissues; 2— time of collection of material; 3— June, 1968;
4— August, 1968; 5— October 1968; 6— March—April 1969; 7— muscles;
8 — liver; 9— heart; 10— kidneys; 11— spleen; 12— lungs; 13— air bladder;
14— skin; 15— brains; 16— spinal cord; 17— adipose fin; 18— internal
fat; 19 — gonads; 20— scales.
10
Table 5
Change in the ratio of phosphatides to triglycerides
in different organs of the Urosozero cisco
é l ItIt tl n
1 leg1:131
7>i Mbautcht
8L flegeth
I
g cigla
mœm 19 .3 n 1
-1,5
57
5.5
91. Cepaue
1 0 ellottbat •
11 iCe.ie3eliiia
12 Xa5pbt
13 I lbaBare.lbiluil nyealii,
14iKowa
1511.0:1013Hoil bto3r
16 :C111111110171 M 0 M'
17 j)K'sipolioet n.namini:
18 .18urrpe:iiiiiil ?imp '
_ .
19 irolia.u:b:
ître
50
5,6
•ci tw.ra
------- 6 '
4,4
59
se
10 9
15,0 .
4A
2,4
LI
r,9
9.3'
4:3
7,7
13,0
5,4
12,3
• 0.8
1,6
1,5
• 0,0
1,8
3e
—
12
• 10,3
20.Xleuval
eape—arsre.n.
rfrb
-
3:3
2.5
3.1
1.0
• 7,3 .
14,5- •
33
0,8
•
22 .
. 3A
JÇFYI;
1- organs and tissues; 2- time of collection of material; 3- June, 1968;
4- August, 1968; 5- October 1968; 6- March-April 1969; 7- muscles;
8- liver; 9- heart; 10- kidneys; 11- spleen; 12- lungs; 13- air bladder;
14- skin; 15- brains; 16- spinal cord; 17- adipose fin; 18- internal
fat; 19- gonads; 20- scales.
Ferdman, 1966).
Unfortunately, we were not able to find papers on
the study of the qualitative composition of lipids of medullary tissues
of fish. For this reason these results still require additional
verification.
/168/
11
The data on the variation of the phosphatide-triglyceride
coefficients from season to season are very significant (see Table 5).
For most tissues, in March and April and the beginning of June, they
are close to each other, in August they are reduced (due to the increase
in the proportion of triglycerides), and in October they increase markedly,
since during this period triglycerides are intensively consumed. Only
in the gonads is the ratio of phosphatides to triglycerides maintained
approximately constant in all seasons (2.0) although both the proportion
of triglycerides (from 9-10% in March-April and 15.3% in August to
8.0% in October), and the proportion of phosphatides (from 21-22.0% in
March-April and 25.0 in August to 15.0% in October) are reduced towards
October.
This fact once again confirms the conclusion that the reduction
in lipids in gonads by the IV stage of maturity is connected not so
much with their consumption (if this were the case, predominant
consumption of triglycerides, which are reserve lipids.by comparison
with phospholipids, which are structural lipids, would take place,
and consequently, the Phosphatide-triglyceride coefficient would
vary) so much as it is due tothe reduction in the total proportion
of total lipids in connection with the neogenesis and accumulation
in the gonads of substances of a different chemical nature (proteins,
nucleic acids, polysaccharides).
Conclusions
In field conditions, material was gathered and fixed for the
study of the seasonal dynamics of total lipids, phosphatides and
triglycerides in 14 organs (or tissues) of the large cisco (Coregonus
albula L.
12
It was established by the methods of thin layer chromatography
and spectrometry that seasonal dynamics in the content of total
lipids and triglycerides, with a maximum in August and a minimum in
October, which clearly correlates with the foraging and prespawning
periods in the life of the cisco, and also corresponds to the III and
IV stages of maturity cf the gonads, is characteristic for almost
all organs.
It is shown that the seasonal dynamics of total lipids correlate
quantitatively with the seasonal dynamics of triglycerides for all
organs with the exception of the gonads.
The phosphatides, which are structural lipids, are iess subject
•
' t seàsonal
It is established that the content of phosphatides in the
gonads, in distinction from other tissues, is also subject to seasonal
fluctuations, which is due, primarily, to change in the ratio of
accumulation of lipids and other nonlipid substances (proteins, polysaccharides) in the process of maturation of the gonads (ovulation).
13
References
1. Belyaeva, K.I., Pokrovskii, V.V. The large cisco of the Karelian
lakes as an object for artificial culture. In book: Karelian
fisheries.
No. 7. Petrozavodsk, Gosizdat KASSR, 1958.
2. Lizenko, E.I., Potapova, 0.1., Sidorov, V.S. Seasonal fluctuations
in the content of lipids in various organs of the Urosozero
cisco.
In book: Eighth session of the Learned Council on the
problem: "Biological resources of the White Sea and internal
bodies of water of the European North". Theses of reports.
Petrozavodsk, 1969.
3.
Potapova, 0.1.
Titova, V.F.
Materials on the fatness of the
Munozero cisco in the foraging and spawning periods. Tr.
,;^,_,.. ... .,<^.-- , -... . >.:_: . - -, 4 ,-^r •-. . ...-_ ^r-::,.t;-^...K. °2:- ...--.^,- .. ^ ...^^ra . ., ... .,,. -- .
Karel. otd-niya GosNIORKh, 1967 (1968 edition), vol. 5,
_ - ^,>.. ..,.^ ..
no. 1.
4.
Potapova, 0.1., Titova, V.F.
Materials on the fatness of the
large cisco of some Karelian lakes. In book: The metabolism
and biochemistry of fishes.
5.
Ferdman, D.Ya.
6.
Shtraub, R.B.
Biochemistry.
Biochemistry.
Moscow, "Nauka", 1967.
Moscow; "Vysshaya shkola", 1966.
Budapesht.
Publishing House of
the Academy of Sciences of Hungary, 1963.
7.
Shul'man, G.E.
The dynamics of fat content in the body of fishes.
Uspekhi sovrem. biol., 1960, vol. 49, no. 2.
8.
The storage of lipids in Coregonus albula L.
9.
The regional distribution of fat in various freshwater fish species.
Remaining reference items listed in English on attached copy of original.
14
B e .11 Rena 11/1., onponcnn Fr B. B. Kpyrinan panyunza o3ep Kapearin Kan
o6.berrr ricKycernennoro pameenusr.- B
Pw6noe X03-B0 Kape.amr. Bbin. 7. rferpo3aBOACK. fOCH3.UaT KACCP, 1958.
L • J1 u3enno E. H., 1.1 or anona O. H. C « gopon B. C. Ce3onubie none6anna
co.a,epn:ainta muntiron n pa3auttribix opraitax panymn Ypoco3epa,- B NH.: Bocbmaa cecena
it sayTp. noa,oemors
YLICH. conera no npo6oeme: «Bno.lorntrecime pecypcm beaoro
EnponeikKoro Cenepa». Te3. Rom. IleTpo3ano,3,cn, 1969.
• 11 oTanona O. 117,-T nrona B. O. MaTepuaam no a:111 3 1mm myno3epcnoil .panymKII n 1-larva:A Ibffi n nepecroumil nepiroam.- Tp. Kape5. OTJ2(-111151 FocHHOPX, 1967 (113n.
1968). T. 5. BIAH. I.
4.1-10 ranona O. 14., Tarona B. D. Marepumm no BilipHOCTH npynnoilpanyunur
nencrropmx 03ep Kapemm.- B NH.: 06men neaten-Es u 6noxinnua pr,16. M., «Hayxa», 1967.
.(1) epam an U. 51. Bnommila. M., «Brecuran uncoaa», 1966.
6•1lirpa y6 P. B. Eatoxwmna. By.a,anetur,
A1-1 Bel-Irian!, 1963.
.7 .111 yabsau r. E. JInnamuna, coep:nansia nzupa a Te.ne pm6.-7 Yule» conpem.
6no.I., 1960, T. 49. nwn. 2.
81. fl h in a n M. Ober die Fettspeicherung bei der Kleinen Marne Coregonus albula.
.
11.-- Arch. Fischereiwiss., 1955. Bd. 6, 411/2.
9.M orawa J. Die regionale.Verteilung des Fettes bei verschiedenen Siisswasserkicharten.- Zeitschrift für Fischerei, 1956, B. 5, H. 1-2.
-LU•Musacehia X. J., Sullivan B. J., 'Wilber C. Y. A composition of liver
and mm-cle lipids in artic fishes.- Copeia, 1957, 1V,..> 1.
11. V a d a. Agile. Chem. Soc. Japan, 1955, 29, A'"? 5-6 (an) 169 L11Ir. no PA «BnoxnNun», W56, :V2 6695.