fisheries research boaro of canada technical report … · the relative nutritive value of...
TRANSCRIPT
FISHERIES RESEARCH BOARO OF CANADA
TECHNICAL REPORT NO. 334
1972
FISHERIES RESEARCH BOARD OF CANADA
Technical Reports
FRB Technical Reports are research documents that are of sufficient
importance to be preserved, but which "for some reason are not appropriate for
primary scientific publication. No restriction is placed on subject matter and the
series should reOect the broad research interests of FRS.
These Reports can be cited in publications, but care should be taken
to indicate their manuscript status. Some of the material in these Reports will
eventually appear in the primary scientific literature.
Inquiries concerning any particular Report should be directed to the
issuing FRB establishment which is indicated on the title page.
FISHERIES RESEARCH BOARO OF CANAOA
TECHNICAL REPORT NO. 334
HFECT OF UNICELLULAR ALGAL LIPIDS ON OYSTER
LIPIDS AND WEIR FATrY ACID COMPOSITIONS
BY
TAKESHI WATANABE
Tokyo University of FisheriesMinato-ku
Tokyo. Japan
AND
R.G. ACKMAN
Department of the EnvironmentFisheries Research Board of Canada
Halifax LaboratoryHali fax, Nova Scotia
FISHERIES RESEARCH BOARD OF CANADAUalifax Laboratory. Halifax, N.S.
ABsrnACf
Lipids and fatty acids of American (Crassostrea virginica) and
European (Ostrea edulis) oysters were studied to determine the effects of fceding
on the unicellular algae Dunaliella tertiolecta. Changes during storage of C.
virginica for six months were also examined. Separate experiments were carried out
with the algae Isochcl}Sisgalbana and Dicrateria inornata and both species of oyster
but without prolonged storage. In addi tion to study of the total lipids the algal
lipids were separated into polar and non-polar fractions, and oyster lipids into
polar lipid and triglycerides as well 35 stery! esters. glyceryl ethers, and free
fatty acids where practical. The detailed fatty acid compositions show that major
fatty acids in the diet mostly have recognizable but limited effects on the oyster
lipid fatty acid composition, presumably because the oysters stabilize lipids at near
optimum composi tions. The two oyster species responded differently to dietary fatty
acids and/or different algae. Hitherto unknown C20
and C22 non-methylene-interrupted
fatty acids are included in the detailed fatty acid analyses.
-2-
INTRODUCTION
The relative nutritive value of different species of marine
phytoplankton for zooplankton and larger filter feeders has been the
subject of speculation and study by many authors. Raymont (1963)
has reviewed the results of studies on the feeding of copepods. and
Walne (l970b) has reported on the relative food value of nineteen
genera of phytoplankton for oyster larvae. Bardach (1968) has also
reviewed recent developments in artificial rearing of many types of
marine organisms. Much of the earlier work has been based on total
calories or on amino acid "nitrogen" • with a tendency to regard protein,
carbohydrates and lipids as gross nutrient classes. with Ii tUe attention
being devoted to "essential" components in detail, although Iwasaki et al.,
(1971) have given consideration to an "essential amino acid" index.
The lipid components of planktonic organisms are a major
fraction of the metabolic energy resources of the sea. Various studies
which have been carried out on the biochemical composition of planktonic
organisms have yielded valuable infomation on the amount of lipid, on
total lipid fatty acid composition. and on different lipid components
for both zoo- and phytoplankton (Ackman et al., 1964; 1968; Corner and
Cowey, 1968; Culkin and Morris, 1968; Raymont et al., 1968; Chuecas and
Riley, 1969). Recently, Lee et al., (1971) have examined the importance
of wax ester formation as well as details on fatty acids in other lipids
in the marine food chain step between phytoplankton and copepods.
- 3 -
Phytoplankton are now accepted as the primary sources
of certain "essential" fatty acids of marine invertebrates and fish
(Farkas et al., 1961; Kayama and TSllchiya, 1962; Kayama et al., 1963)
but the step between plant and primative filter-feeding animals has
received little attention. In particular the economic importance of
oysters and potential for aquaculture indicate a need for infomation
on the fatty cid composition of oysters fed specific algae. We have,
therefore. c tducted feeding experiments with oysters using pure cuI tUTed
unicellular algae to investigate the influence of algal lipid on oyster
lipid fatty acid composition.
The chief experimental animal used in our investigation was
the American oyster. Crossostrea virginica. It is plentiful in the
coastal waters of the United States from Texas to Maine, and also inhabits
various warmer and sheltered bays and estuaries in the maritime provinces
of Canada (Bousfield. 1960). The introduced European oyster. Ostrea edulis.
was used in a limited comparison study.
MATERIALS AND METHODS
All the feeding tests were carried out at the biological
facilities of the Fisheries Research Board of Canada at Ellerslie.
Prince Edward Island. The axenic algal cultures were grown at 20_2S o C
and under sunliJtht. An enriched seawater medium was used. Both American
oysters. C. virginica and European oysters.o. edulis used in experiments
I. II and III had been kept in filtered seawater for more than one year
at the b ologieal facilities. In these experiments oysters were fed
- 4 -
Dunaliella tertiolecta. In other experiments in which oysters were
fed Isochrysis galbana and Dicrateria inornata the test animals had
been held in natural conditions in the sea near Ellerslie, P. E. T.
prior to feeding.
In experiment I. the oysters were brought from cold water
(ca. 2°C) to room temperature (20 G e) and held for 1 day before the
feeding experiment. For experiments II and III the animals were
transferred from cold water about 2 hours before commencement of
feeding. A representative group of oysters from each lot was used as
an unfed control for each lot. One experimental technique was used forunialgal
all feedings. Four litres of I culture (in seawater) were placed in
a polyethylene tank (45 x 30 x 15 em) containing 10 oysters. The
temperature of the culture and of the seawater from which the oysters
were removed was 20_22°C. The oysters began to actively filter the
seawater within 5 minutes of being covered by the liquid. The cell
densi ty of the cultures in seawater was determined every 30 minutes by the
packed volume method. When the cell density dropped to about half of the
ini tial value (after 1 hour of feeding). the oysters were transferred
to a new lot of culture. The total active filtering time was 6 hr and from
2.2 to 6.3 g of algae were incorporated by oysters in the different
experimental groups (Tables 1 and 2). After feeding tests the experimental
and control oysters and samples of the algal cultures were placed in plastic
bags and cooled to ice-water temperature for transport to the Halifax
Laboratory of the Fisheries Research Board for analyses.
- 5 -
The American oysters (C. virginica) in experiments I and II were
respectively divided into two groups. One half of each group was
kept at 2°C for 6 months.
Cultures were harvested by gentle centrifugation and the pellet
of algal cells drained to determine approximate wet cell weights (Table I).
Subsequent operations were carried out under an atmosphere of nitrogen.
Methanol and/or chloroform were used to transfer the cells to separatory
funnels with water in limited amounts to maintain a monophasic solvent
for 15-30 min. Additional water was then added to arrive at the final
ratios of solvents indicated by Bligh and Dyer (1959). The bottom layer
was filtered and stripped for lipid recovery (Tables and II).
Extractions of oysters were carried out on total wet organic
tissues with chlorofonn:methanol (2: 1). Polar lipids and non-polar
lipids were separated on and recovered from a divinylbenzene copolymer
bead gel column (Sipos and Ackman, 1968). Non-polar lipids were then
separated into sterol esters. glycerol ethers, triglycerides, free fatty
acids (in the case of o. edulis), and sterols on silica gel TLC plates
(Pre coated Adsorbosi 1-5, Applied Science Laboratories, Inc.). Procedures
of lipid fractionation are presented schematically in Figure 1. For the
algal lipids, the non-polar fraction was directly saponified and
non-saponi fiables were extracted and weighed. The recovered fatty acids
were esterified with BF3
-MeOH reagent. Removal of non-saponifiable
materials from lipids, preparation of methyl esters (by BF3-MeOH
esterifications of acids, or by transesterification of triglyceride), and
all GLe operations (open-tubular columns coated with aDS), followed
procedures described previously (Sipos and Ackman, 1968; Ackman and
Hooper, 1970). Particular care was taken to avoid confusion between branched
acids and plasmalogen artifacts (Ackman, 1972), as oysters are rich in these
lip~ds (Sampugna et al •• 1972).
- 6 -
The principal aim of this program was to investigate the fate
of "essential" polyunsaturated fatty acids as they were ingested
naturally by oysters from algal sources. While this work was in progress
it was discovered in this laboratory that the eicosenoic and docosenoic
acids. which are broken down by open-tubular gas liquid chromatography
into constituent isomers. contained anomalous peaks which were in fact
non-methylene interrupted dienoic acids. These have been included in relevant
data from this studY,but since their evaluation is at an early stage
we prefer to include them in publications with more detailed discussion
of the results.
RESULTS AND DISCUSSION
Lipid levels in each fraction of samples analysed are shown
in Tables I and II. tn experiments I, II and III the low lipid recovery
in comparison with those of oysters later fed I. galbana and D. inornata,
probably reflects the extended holding period without food for these
animals (Ackman and Cormier (1967); Ackman et al., 1971)). The
percentages of polar lipids. relative to total lipid. recovered from
oysters in experiment I. II and III were found to be lower than those of
other experimental groups and that reported by Ackman et al., (1971),
possibly for the same reason. Even the lipid recovery of oysters
collected from natural conditions (1.59\ for C. virginica) were lower
than those reported by Ackman and Cormier (1967); Shimma and Taguchi (1964).
Sampugna et al., (1970) and others, but comparisons are difficult due to the
imponderables such as extraction technique, size, species, environment
and habitats (Ackman and Hingley, 1968). and the effects of seasonal
variations (Ackman and Cormier, 1967; Cerma et al., 1970; Stancher et al.,
1971). However. the substantial residue of triglyceride indicates that
- 7 -
the depletion of lipids is not especially abnonnal in view of seasonal
limitation on food supply and the extreme lowering of water temperature
in the winter in the original habitat in Prince Edward Island (which
is the northern limit for this species for this reason (Bousfield, 1960)).
The fatty acid patterns for lipids of D. tertiolecta (Table III) were found
to be very similar to the result reported by Ackman et al., (1968).
whereas the fatty acid composition of I. galbana (Table IV) differed
significantly in detail from the result obtained by DeMort (1970).
especially in the level of 22:6w3 relative to 20:Sw3. This presumably is
due to the difference of culture conditions such as nitrogen concentration,
light J temperature. culture age and particular strain (Otsuka, 1961;
Otsuka and Morimura, 1966; Ackman et al., 1964; 1968; Lee et al., 1971).
The two species of Haptophyceae, I. galbana and D. inornata, were also
found to be quite different in lipid composition. The influence of the unusually
higher cuI ture temperature and natural sunlight may be profound. Thin-
layer chromatography on silica gel revealed that the non-polar lipid
of I. galbana consisted mainly of sterols, a small amount of
diglyceride and several kinds of pigments. The latter apparently influenced
the high proportion of non-saponifiable material (73.96\), whereas TLC
of D. inornata total non-polar material (only 7.3\) showed sterol
esters, glycerol ethers, triglyceride and sterols as principal components.
One difference which we had hoped to exploit in fatty acid composition
between the two species was the higher proportion of 18:4w3, but not of
18:3w3, and of 20:Sw3, fatty acids in I. galbana. However, there was no
important quantitative difference between control and experimental oysters
for these acids as indicated by eLC of total lipids from oysters fed on
these two species of algae.
- 8-
The fatty acid composition obtained fOT each lipid class
from the oysters is presented in Tables V to VIn. The fatty acids
of polar fractions (presumably mostly phospholipids from chromatographic
behaviour; the actual proportion of phosphorus was not determined) from
experimental groups fed 3 species of algae showed no real differences
between experimental and control groups. although the polar lipid
fatty acids of C. virginica in experiment I. and of O. edulis, both
fed O. tertiolecta. showed an increase of 16:0 and 17:0 acids possibly
related to a decrease of polyunsaturated C20
and e22 acids. The latter are
slightly suspect as delays in analysis of lipids may have led to some
autoxidation. This result for phospholipids seems to be in agreement
with reports that the fatty acids are highly specific and do not vary
with dietary changes because of their role as important membrane
consti tuents (Carroll, 1965; Wolfe et al., 1965; Benson J 1966; Jezyk
and Penicnak, 1966; Keenan and Morrt~J 1970; Lee et al., 1971).
The fatty acids of triglyceride from oysters in experiment
showed di fferences in the proportions of C16
, C18
and C20 monoethy1enic
fatty acids between control and experimental groups. However, after 6
months storage the high proportion of these fatty acids in c. virginica
(reflecting the low amount of 16:0 acid in the initial sample of the
experimental group) tends to show a close parallel to those of control
oys ters and an increase of 20: 5003 and 22: 6003 acids corresponding to an
intake of high amounts of dietary 18: 3003 from D. tertiolecta. In
- 9-
experiment II, the proportion of these polyunsaturated acids does
not di ffer a great deal between the experimental and control groups
both in ioi tial and after experimental 6 months. However. a higher
amount of 20:5w3 and 22:6w3 than in experiment I was recognized in
ioi tial samples and increased after 6 months. These differences in
results obtained for experiments I and II are probably due to the
di fferences in experimental design as the test animals for I had been
allowed to adapt to feeding conditions before the start of the
experiment as mentioned above, while II test animals were warmed up
only an hour before feeding. In the initial analysis (not stored)
of c. virginiclJ of the other experimental groups respectively fed
I. galbana and D. inornata. the proportion of C16
, CIS' C20
and C22
polyunsaturated acids are relatively close to those of controls. In the
fatty acid composition of c. virginica a larger amount of 16:0 (34.8\
in control, 43.5\ in experimental fed I. galbana and 42.1\ in
experimental fed D. inornata) reflects lower levels of 20: 5w3 and
22:6w3 acids. However, in o. edulis somewhat lower levels of 16:0 acid.
accompanied by higher proportions of 20:5w3 and 22:6w3 acids. were found
in the triglycerides of oysters fed I. galbana. The accumulation of normal (20
and C22 polyunsaturated acids in the group fed I. galbana seems to
reflect directly incorporation of fatty acids from I. galbana, which
contain much more 20:5w3 and 22:6w3 than D. 1nornata. The same kind of
differences between C. virginica and o. edulis were also observed in
experiment III. In experiment III, a high proportion of 16:0 acid in
-10 -
O. edulis related to lower levels of polyunsaturated C20
and e22
acids
appears to correspond to low levels of normal dietary polyunsaturated
C20 and e22 3cids from D. tertiolecta. These results probably indicate
that the mode of fatty acid metabolism I especially in relation to the
incorporation of dietary fatty acid, is different between the two species,
C. virginica and o. edulis.
The striking features of the analyses of sterol esters in experiments
I. II and III. in which oysters were fed D. tertiolecta. are: an accumulation
of 14: 0 acid and the accompanying 4.8.) 2-trimethyl tridecanoic acid (cf. Ackman
et al. ,1971) and, reduction in levels of 18:0 acid. In the American oyster
c. virginica, fed I. galbana and D. inornata, increases of monoethylenic fatty
acids along with decreases of 15:0 and 16:0 acids are noticeable, especially
in the oysters fed D. inornata. A similar tendency was observed in o. edulis,
but the proportion of 14: 0 acid was found to be low in the group fed I. galbana.
Polyunsaturated C22 acids were absent in both groups, whether control or
experimental.
Our results show that the variations in fatty acid composition between
the major lipid classes (e.g. phospholipids and triglycerides) are to some
extent independent of food intake. The relative proportions of the important
classes of I ipids may therefore be partially responsible for the seasonal
variations in total fatty acids observed in many molluscs such as those
studied by Calzolari et al., (1971). The fatty acids in total lipid of the
brine shrimp Artemia salina also show some independence of specific phytoplankters
in the diet (Hinchcliffe and Riley, 1972). but different lipid classes were not
studied.
-11-
While this work was in progress it became apparent that the fatty
acid patterns for monoethylenic acids from c. virginica and other molluscs
differed significantly from those normally observed in marine oils and lipids
(for examples, see Ackman and Castell, 1966 and Addison et a1., 1972). A
partial elaboration of structures indicates that the unusual "monoethylenic"
components are eicosadienoic and docosadienoic acids, probably of 5,13 and 5, IS
unsaturation for the (20:2) acids and 7:15 and 1:17 unsaturates for the (22:2)
acids. The ( ) indicates that these do not belong to the usual methylene
interrupted polyunsaturated families of fatty acids. The CI8 unknowns have
not been characterized at the time of writing. Figures 2 and 3 show the effect
of an isolation of "normal" monoethylenic C18 and C20 acids by argentative TLC,
while Figure 4 shows unusual configuratiof6 of the (22: 2) acids.
Because of the novelty of these results we have isolated the data for
the (20:2) and (22:2) acids in separate subtables V-B, VI-B and VIII-B in
addition to the complete tables and subtotal tables of the A series. There still
remain some C20 and C22 unknowns. but these apparently occur unsystematically.
It is remarkable that the (20:2) and (22:2) acids were not found in the o. edulis
fed Isochrysis and Dicrateria (compare Tables VII and VIII). but there is no
obvious explanation for this.
ACKNOWLEDGEMENT
The authors express their thanks to Mr. R. Drinnan for arranging
for cultures of the algae and for valuable advice. T: Watanabe was the
recipient of a National Research Council of Canada Post-doctoral Fellowship.
-12-
REFERENCES
Ackman, R.C.. 1972. Tn "Pro~Tess in the Chemistry of Fats and Other Lipids
ed. R.T. Holman. PeTRamon Press, New York, Vol. !I. pp. 165-297.
Ackman, R. G. and M. G. Comier. 1967. a-Tocopherol in some Atlantic
fish and shellfish with particular reference to live-holding
without food. J. Fish. Res. Bd. Canada ~J 357-373.
Ackman, R. G. and H. J. Hingley. 1968. The occurrence and retention of
dimethyl-Bpropiothetin in some filter-feeding organisms.
J. Fish. Res. Bd. Canada ~. 267-284.
Ackman. R. G. and S. N. Hooper. 1970. Branched-chain fatty acids of
freshwater fish oil. Compo Biochem. Physiol. g. 117-125.
Ackman, R. G.• S. N. Hooper and P. J. Ke. 1971. The distribution of
saturated and isoprenoid fatty acids in the lipids of three
species of molluscs, Littorina littorea, Crassostrea virginica
and Venus mercenaria. Comp. Biochem. Physio1. ~, 579-587.
Ackman, R. G.• P. M. Jangaard, R. J. Hoyle and H. Brockerhoff. 1964.
Origin of marine fatty acids. I. Analyses of the fatty acids
produced by the Diatom Skeletonemo costatum. J. Fish. Res. Bd.
Canada ~, 747-756.
Ackman, R. G., C. S. Tocher and J. McLachlan. 1968. Marine phytop1ankter
fatty acids. J. Fish. Res. Bd. Canada ~, 1603-1620.
Addison, R.F .• R.G. Ackman and J. Hing1ey. 1972. Lipid composition of the
queen crab (Chionoecetes opi.lio). J. Fish. Res. Bd. Canada 29: 407-411.
Bardach, J. E. 1968. Aquaculture. Science .!!!., 1098-1106.
Benson, A. A. 1966. On the orientation of lipids in ch1oropast and cell
membranes. J. Am. Oil Chem. Soc. Q, 265-270.
-13-
Bligh, E. G. and W. J. Dyer. 1959. A rapid method of total lipid
extraction and purification. Can. J. Biochcm. Physiol. lZ..
911-917.
Bousfield, E. L. 1960. Canadian Atlantic Sea Shells. National Museum
of Canada. 72 pages.
Calzolari, C., E. Cerma, and B. Stancher. 1971. Gas chromatography applied
in determining fatty acids of some gastropoda and lalllcllibranchia from
Adriatic sea during an annual cycle. Riv. Ital. Sost. Grasse XLVI II,
605-616.
Carroll, K. K. 1965. Dietary fat and fatty acid composition of tissue
lipids. J. Am. Oil Chern. Soc. £. 516-552.
Cerma, E.• B. Stancher and P. Baradel. 1970. Molluscs of the upper
Adriatic Sea (Chemical composi tian of some of the Gastropods and
the Lamellibrachs). Rassegna Chimica (Chemical Review), ~.
39-43 & 3 unpaged sheets.
Chuecas. L. and J. P. Riley. 1969. Component fatty acids of the total
lipid of some marine phytoplankton. J. Mar. 8iol. Ass. U. K.
~, 97-116.
Corner, E. D. S. and C. B. Cowey. 1968. Biochemical studies on the
production of marine zooplankton. BioI. Rev. ~. 399-426.
Culkin, F. and R. J. Morris. 1969. The fatty acids of some marine
crustaceans. Deep Sea Res. .!!., 109-116.
DeMort, C. L. 1910. The culture and biochemical analysis of some estuarine
phytoplankton species. A thesis for the degree of Doctor of
Philosophy in Oregon State Universi ty. 157 pages.
Farkas, T.• S. Herodek, L. CS'aki and G. Toth. 1961. Incorpor3tion of
acetate_I_C I4 into the liver fatty acids of the fish Amiurus
nebulosus. Acta BioI. Acad. Sci. Hung. g. 83-86.
Hinchcliffe, P.R., and J.P. Riley. 1972. The effect of diet on the component
fatty acid composition of Artemla salina . .J. mar. biol. Ass. U.K.
-14-
Iwasaki J H., S. Tanaka and T. Fujiyama. 1971. On Rhodomonas ovalis
Nygaard as food for clam larvae. Bull. Jap. Soc. Sci. Fish.
Vol. 37. 1044-1048.
Jezyk. P. F. and A. J. Penicnak. 1966. Fatty acid relationships in an
aquatic food chain. Lipids!.. 427-429.
Kayama. M. and Y. Tsuchiya. 1962. Possible conversion pathway of
polyunsaturated acid in fish. Tohoku J. Agriculture Res. !l.
229-235.
Kayama, M.• Y. Tsuchiya and J. F. Mead. 1963. A model experiment of
aquatic food chain with special significance in fatty acid
conversion. Bull. Jap. Soc. Sci. Fish. ~J 452-458.
Keenan, T. W. and O. J. MarrEL 1970. Phospholipid class and fatty acid
composi tian of golgi apparatus isolated from rat Iiver and
comparison wi th other cell fractions. Biochemistry U.S. A.
~. 19-25.
Lee J R. F .• J. C. Nevenzel and G. A. Paffenhttfer. 1971. Importance of
wax esters and other lipids in the marine food chain:
phytoplankton and copepods. Marine Biol. ~,99-108.
Otsuka. H. 1961. Changes of lipid and carbohydrate contents in Chlorella
cells during the sulfur starvation as studied by the technique
of synchronous culture. J. Gen. Appl. Microbiol. Tokyo 2.. 72-77.
Otsuka. H. and Y. Morimura. 1966. Changes of fatty acid composition of
Chlorella ellipsoidea during its cell cycle. PI. Cell Physio!.
Tokyo 7... 663-670.
Raymont, J. 1963. Plankton and productivity in the oceans. New York,
Pergamon. 660 p.
-15-
Raymont, J.E.G., R.T. Srinivasagam and J.K.B. Raymont. 1969. Biochemical
studi es on marine zooplankton. VI. Investigation of Heganyctiphanes
norvegica (~l. Sars). Deep Sea Res. ~. 141-156.
Sampugna, J .• L. Johnson, K. Bachman and M. Kenney. 1972. Lipids of
Crassostrea virginica. 1. Prel iminary investigations of aldehyde and
phosphorous containing lipids in oyster tissue. Lipids 2.. 339-343.
Sampugna. J., M. Keeney, K. Bachmann and L. Johnson. 1970. Lipids from
the oyster Crassostrea virginica. Unpublished data. Am.
Oil Che•. Soc. (Abstract froID. 2nd Int. Soc. World Fat Res.
Congress; Chicago. Sept. 27 - Oct. I, 1970).
Shimma. Y. and H. Taguchi. 1964. A comparative study of fatty acid
composition of shellfish. Bull. Jap. Soc. Sci. Fish. ~' 153-160.
Sipos, J. C. and R. G. Ackman. 1968. Jellyfish (Cyanea capillata) lipids:
Fatty acid composition. J. Fish. Res. Bd. Canada ~' 1561-1569.
Stancher J B., E. Cerma and P. Baradel. 1971. Mollusks of the upper
Adriatic. Variations of the chemical composi tion of some
gastropods and lamellibranchs during an annual cycle. Rassegna
chimica (Chemical Review), ~' 39-42.
Wa1ne, P. R. 1970a. The seasonal variation of meat and glycogen content
of seven populations of oysters Ostrea edulis L. and a review
of the Ii terature. Fishery Investigations. Series II,
Volume XXVI, No.3, 3S pages. Her Majesty's Stationery Office,
London.
Wa1ne, P. R. 1970b. Studies on the food value of nineteen genera of algae
to juvenile bivalves of the genera Ostrea, Crassostrea, Hercenaria
and Hytilus. Fishery Investigations. Series II, Volume XXVI,
No. S, 2 pages. Her Majesty's Stationery Office, London.
-16-
Wolfe. D. A., P. V. Rao and D. G. Cornwell. 19-5. The fatty acid
composition of crayfish lipids. J. Am. Oil Chern. Soc. £.
633-637.
Gel column
Total lipid
Saponi fication
IPolar lipid
INon polar lipid
ITotal fatty acids
INon saponi fiable
Fatty acids Non sap.
Methyl esters uu.. GLC
Hydrogenated methyl esters uu~ GLe
TLC
Methyl esters uu. GLC
HYdrogenated methyl esters uu" GLC
SterylEsters
Glycerylethers
Triglyceride Freefattyacids
Free sterol
___ ~ GLC Hydrogenatedmethyl esters
r
Saponify
Fatty acids
Methyl ~ GLCesters
Hydrogenatedmethyl esters
JGLC
Saponify
Fatty acids
Methylesters
-IHydrogenated
me:tl esters
Transesterify
Methyl • GLCesters
GLC
Methyl • GLC
~
Hydrogenatedmethyl esters
1GLC
Figure 1. Schematic representation of experimental procedure for lipid analyses.
OYSTE R (c. virginico)
T
PART OFTOTAL ESTERS 18:2""6
18: 0+
18: I
r--'---o",,9
""7
MOST MOBILEBANOS
80S
ADIENEBAND
LLJf/lZoQ.f/lLLJ0:
0:LLJC0:oULLJ0:
18:0\
18:0
""9",,7
TIME ----t
Figure 2. Comparison of totol C18 esters of C. virginico lipids (left) with nominal monoene (plus saNrated)ester bond (center) and a diene bond (right). 80S column, slightly diff.rent temperature on right-handgroup af peaks.
TTOTALESTERS
w(f)
ZoQ.(f)
Wa:a:woa:ouwa:
20:lw7
(20: 2)
20:0 +20: I
w7
OYSTE R( C. virginico )
BDS~
( 20:2 = NMI DIENES I
(20: 2)5,13 ?
TIME --.
Figure 3. Comparison of total C20 .sters of C. virginico lipids (left) with 0 monoene bond (center) and thediene bond (right). 80S open-tubular column, slightly c1ifferent temperature on right-hand group of peaks.
(7,17? )I .
(7,15 jl)"
(22:2=NMI DIENES)
OYSTER(C. virginico)
22:1",7
{"'13 11\22:1 ,./ir .
I 22:1(0)9
I
+- ex I 4X +
80S leOleo
~
20:5",3
PART OFTOTAL LIPIDSw
lI)
zoQ.lI)
Wa::a::woa::ouwa::
TIME~
Figure 4. GLC chart to show portion of non-methylene-interrupted C22 diene! relative to normal dienes. Normal22: 1 w13 + 22: 1 w 11 and 22: 1II) 9 were isolated from this oyster lipid, bYt shoulder thought to be 22: 1w7
was not OccOUflted for.
" :::::::::::::::::;t~<l~ClCl~ ;;.;;;;;;;;:;f I[nt:tr£tn~ tu:ut~n~. ~
oeeee!!ll::::;:;: ;,::S:::;:,;t 1~2 l:ol$:'t
Cl~;tll:;;;;;;
;;~ i:!:~~:i:;: ~~:~:::n= ,;:
,. oj
.. " ~ .. _e ~ ~ .... ~ .......
;;.,t ;;s ~ ~~~ r~;!~,
, :' e ........ e:-i: .. :-~ .. :- ...;:;;: ~ ;;:;~~;;:::::O:l::::~:::~
~ .. eeeee .. e:-::!::i~:Oi;:;::i:l
. e~eeeeee2' :=:::2'U:lIl: :::::! e;::~
i:!:'
,ee,;·u a ~~'
...... ,.~:!:e r e:
e~ .. , ......:;::::::; ;:;
........... e ... :: .. ~ .. ~:-~:;;;Ii:.l;:t,.;:;:; ';:~:;:::::8;:
e., .... _... t: .. _ .... __ •~;;.':;:;::!:!:;:;::~s::::se.::::
....... _e_ ..;:;I:!::~ee;
e ... e ... _ .. e:;:;;;::a:::: :!:::l::Cli:!
_eo e_ ... ll:~'=~ ;;;:;::.-:1: ~~:il:il <!l
._e.ee .. __ :::
;;:;:::;t,i~':"'::
.:-_::.o:- .. :-?:;:lj'~; :::::0:: :::=::~;;
....... _ -';1-
:~:r:8 :'!
"~~~?~?~ . - .. e_"=;:;;:;o~;~: , t:::::l: :;lie ~:O:~i:il~ , , • ~ r, ::: :; ~ "':::;:: ~::::
; -,; ,;; :';::l ::~ ~8::; ~2': ~ ~ ~ ~ ~~,;z =-;;::::(::!: :::::;:::l'l
, ,
, .,
o :.:
" .
Table V-B. UnU'IU" I non-.cthylenc-Intcrrupted fatty aclds hoillted frOll
O. HuH. total lipid and vulous lipid fnctlons be foro and after feedln. on D. tertl('llect•.
Qrtrea edulla, Control Ind Exper!laental fed Dunallella
Total Lipid Polar Lipid Triglyceride
Control Bxperluntal Control Expert.ental Control Expert.ental
(20:211)
(22:2s)
0.38
1.29
1.11
0.70
0.54
3.70
0.21
1.94
0.19
0.54
0.42
1.26
Steryl Ellters Glyceryl Ether Free Fatty Acid
Control EllperillCntal Control Expert.ental Control Experhaental
(20:2,)
(22:25) 3.02 1.81 0.72
"' ..... 1 ...1... 1 ,.",.•"J, hal..... ,- ~"qJn."'. '0'01 I,p'" ,"j,·."o., , .,',~ -, .... ~"q""'-'. lUo,.ol .noIl.,.......... 1 r.4 _1J.lJo
e., " -,q-.,--~I~ Coo'n>1 up<'r, ......1 CoB'",1 Eqert ....'.1 Co",,,,1 £oport_IOI c.o'n>1 !:.>;pe.._.al 0>0..<11 LLpo""UI
", '" .n •. Ol l.n .." ,.~ 1,11 l." '.n l.~l l."•.•• 1. ~..,., '" D.n ,~ O.ll ,~ O.ll D.ll 0.'1 "."l •. U... '" D.S' D,SI O.SS ,~ on ,.~ O.SI .,. '.'6O.'l
U...... " ..·130 '" 0.1l " . 0.11 D.H 011 0.11 0,07I."'" '" 1., ». 1.0: l,U '.1' I.Dl '" 1.1' ". L. 1.U l.n1... ·1.. 0 oc O.ll ." , .. O.l' 01O 0,11 OM 0.0l
U0.': O.l·1 •••IO.I"~ ,~ 011 ... 0."
1I.1l lS.ll :I.,," 1:.l1'.n
~.'n
n.n XI.'I", IS.O: .,.'I.ll ".n n.n1... ·170 1.11 ,~ ". 1.t>S ,. 0.1S I.n 1.15 1.U I.'l I.SI l.,'..1.....·110 ,. I.n D." 0.• ' ,.~ O.ll ,.~ .. ... 'N,', ,..., 151 .n l.l" '.Ol 1 .. 'N L' L. U
,n1 .•1 ,. ,..'.'.II.IS·"" , .. ",' 0.1. U I.ll O.IS I.ll1"..·1.... 0." 1.1' 0.10 0.1' I.'l ,n O.ll
'" .. '.fl I." ,. U '.1: '.16 ,. ,.. ".1'I' .• ." 0.1l ,n ,.. D." 0.11 U 0.0';0.1 ." .. '" II' 0."1:050....." 11.DI ., .. <>1,0"
lSI' 0.1. 1.10 ,11 ou1.1... 1.11 1.1. ,. ", 0.'1 ,n 1,0: 0.".l.·O
1.1.' .. 6.1' •. r ..•;I.S.: :..., ll,n
l,S'
".1"·0." ,~ ,~ O.CI 0.11
lilA 1111 " l.': '" .n .. '.11I.n
..L.
1.tI11,1.' ,n 10; ,.. ,. l.U 1,7l 1.1' I.~1I.I.l ,,' n n 0.:1 0.•0 '.U ... ,~ ,n 1.21 ,.O.ll
1"1' on U ,.O.I~
:0'1.11 ,n ,~ on ,. D.ll D,57 .. ,~ l.0l L. ,~ 0.11n ,~ • n " . L. D.'I 0.10 ,. 1.11 l.'l I.n l.U 0.09 'n1O:1..' .. ') S.O' ,n ,. ,~ U •. 11 I.n '.n l.H}O'IA '.M ." Ln I,n 0.03 0.11 '.S!in:f 1m '" O,IS ,.:i.:0.1. ,n ..
D." '",.
0.:',~ o.n 0,11
:.Sl '.~ o.nI," ... • n 'n O.OS ,. 'Nn .., 1'1lI IfU " .•S I'." 11,00 n.1l I'.n ".'1 11 .•1 ll.~ lO.to '.n1.0-3 on ,. ,. ". 0.11 ,.161'" '.11 ,n .n O.l: 0,11 ,. O.ll '.lS '.n,,~ , .. .., h
o.n0.11 0.61 ,. ,. o,n ,n O.ll 0." o.n D,1O1.:JwJ D.P 'n ,.
D.IS D,n ,. n O.ll 0.'11"_1 0.'1> •. n O.ll11,:..0 ,~ 1.11 '.n 1.11 OM l..S 1.1.11:1'" •. n O.OS 0.13 '.N1I,J..6 .~ ,~ ." '.M 0.10 0.1l 0.11I.:lo.l .n o.n ." LN 1.6S '.N I.Jl ,.~ I.ll I.U L~II • .,J lOS OM l.ll '.Jl '.N '.N 'N 0.'1 I.l'11,,.._. III I .•) ." 1,1S 1.01 LM 1.•0 O.lS o.n : ..:E:EI'3¥9J , .. 'N ...0.01
:.ll 0.15 D,IS ... 0.1' 001 '.n 01' '.M 0.'10.0: ,. , .. o.os lI.n '.n 'M 0,07 '.Il 0.11O.IS
D.H 0.0' O.ll '.n 0.11 D,IS 0.1. 0.11 0."1O:'.,)
1.23 • n ...'N 0.0l
.. I.n 'N 1.'1 D.1I l." ,~ l"1 1.SS.. .., 0.11 D.'S O.IS 0.11 ... 0.17 11.1' O.lS ..~lO,So.) .., ", ,. 1.1l ..0.2'
10.56 12.ID •. 31 0.10 ..16 'M 11.06 10.5.010,,"'ho,,", 0.01 1.11> I." l.~: O.ll 0." 0." 1.61 2.81 O.S' .."11:" ,. ~. II 0.2S 0.120.11
0.57 0.61 0.10 0,'1 D.lIl ..~ 0.:1>12:..... ,. 0.02 0.2S 0.11 •.n ,.. 0.01o.nn:l.6 0.15
'N..,
0.10'.m .~ o.n 0... 0.1'
0.l70." D.ll11,SooJ D.ll ." D.ll 0.1' 0 .•1 o.n 0.)9 '.n 0.1.
•. U11:1>003 S.O) D.n ~1! ,. .. IS.IS 1l.71 l .•' l.a
1.37 '.N '.nn'""b.-.. I,n ". '.n o. 1~ 0.2. 0.17 ... '.Nll'ollcno 10.15 ." 20,'5 26.0\ 16,11 SO.19 ... 01 25 " Ill.$:! lO.pS JJ.P' 31." 3325
f'"l,,,h"d " I"~ ". ..
frnllo.... l.~ 'M ."hou' r.... ,•• ""D. ,.,tJ.j_~ • VI<! .r... oh ...,~. 010<>1'.
',·.....«_.;...,10,<•. "'.'rol u6f..""I ...uI '" .......11.11.
T·'rl1<.r,~. SU.,I h .... (••h~..hl
~ ... , .... H ... ['I'. H .~. ,"",..,01 I...,..l_ul .....IE.<porl ...UI C••" ... I [''1''''''''101 ",*, ...1 Exp<t.'......1 """ ...1 Eo:porl ......1 """"'1 Eo:po.I_..l e-'.... l Er,>r""o'OI
~,U 3.Z' 'M .~ ." •. » •.•1 •. Il ..• 1.•1,Io.n •. S3 10.IS0.1. o.r. 0,70 0 .•• .~ 0.•' .~ o.n 0,'1 o.n o.s' .u Z.•'D,lt O.ll 0.•' '.M 0.'. 0.'1 D.•' W .~ 0,51 0» 0.11 •.u L"O.lf ..~ .~ .., .» ." o.u O.U 0.1' 0./1 .~ O.'S l.ll '.MI.U 1.17 ,.~ 2.10 I.n L~ I. ~S 1.63 I.ll LM ,~ Z.ll ,.~ ,.~O.ll O.lZ 0.'1 0,31 .., U O.U U 0» 0.1' ." 0 .• ' O.lS a.•'n.n n.u 51,'1 ,...•. ". 0,.0
u .• 3S.~ ~. 2..... 1'.39 n,.3 Z'.·S.." 1.13 0.'3 L~ .,.. n.t!>I.U 1.51 ,." •. n 0.11 0.S30.•1 0 .•1 0,.7 ... I.Jl •. » •. n •. n O.U O.S' o.n ... 'M.." ,.
'M ,~ 2.'J /'.1 1.11 2.05 I.U lo.ll 'M ,.OOJ • 11 •. Z• ... 0,51 O.Sf 0,1'0.15 ." 0.13
'·11 .. 1·'5 '.51 .... 0.51 •. n '.01 0» '.1'·.11 l." '.15 ,. ,. 1.Z' '.u ,." 1.51 I'.n .." 5.11." 0.10 '.M 0.11 .." ..~ 0.'7 .".~
n.OJ 0972 Sl·72 Rl. n.so '0."0» 0.11
L~..~ '.M .~
JSl '.M ..... 1,15 0.93 1.2' 1.J1 L. .. I.ll ,~'.n 1.13 lo." 10..10 '.n lus ,.. '.'0 ..11 llOU ;1.... ill... lI.n..~ ." 0.61 •. n 0." ... O.Ul,21 ,u I.n .. ... ,. l.U ..n '.11 u 7.'1 I •.•• 11.79,.~ I.U ,. '.n 'M 3.S1 '.M '.21 1.72 I." 1.J1 L..U 0.'1 0,S1 0» '.3' O.lJ 0.'0 .. O.lf ..n on 0.15 0.11 •. n'.10 o.M O.IS 0.1S .... ..,.., .. • n ,. 'M L. ... 0.11 1.1. ... ,.» I.IJ•. Tt on .. '.Sl •. 02 ,.~ •. n '.1' 1.01 L~ ... ... 0.12 1.21,.~ OM 'u ,." L. Z." 1.13 LM •. » 1.01 I.U '.M
0,11 0» '.2' •.» 1.15 '.M0,'0 •. 21 ..• O.la O.IS ...0.03
OU ..~ U L~ 1.17 •. 1' 0.1I '.21... ." .. •.U ..• 0.11 ..~ ..~ .."1'.lI 1',N n.n n.u n.l. 21.lS /1." 71.11 '.M ZI.OI 03.... .uo.n 0,31 ... 0."0.1' 0» ...
..u •.uh '.M
0.'10.21 .." 0.11
0.13 o.n0.1' 0.01 0,11 O.U ..~ O.SJ 0.15 0.31 O.U 0.33O.U 0.31 .." 0.11 '.M 0.1' ... .." .~ 0.11 0.51 U0» O.JJ
2.U 1." 1.Sl 1.•' '.M L. I.UD.51 1.510.10 0.17 ... 0.10 .." 0.01 l·.n' ......• .. 0.11 O.ZS .." 0.11 D.n 0.1l1.12 ,.» 0.31 1.21 1.'1 1.11 L~ 2.Sf 1.Sl ~.Jl o,n ..~ 1.1.I.U 0 ... I.•• '.n 1.01 1.01 3." 1.11 2.n '.M 0.61,." 5." 1.73 l.n L. ,.~ l.n '.u 1.12 ,."0.2& o.n 'M 0.11 O.lS 0.12 0.1' U 0.31 o.lT0.05 O.U 0.10 0.03 ..• .." 0.11 0.05.~ .." 0.10 0.01 .." .» 0.03 0.01 0.13 ... 0.1' ..•
0.11J.U 1.1l ." O,~' 0,71 •. u .." 1.1l 0.13 0.51 O.OJO.Sl .." O.OJ ..• O.SI O.U 0.10 0.3J 0.01 .. " ... 0» ...11." n.IJ u D.'l ... •• 11 •• 13 '.03 '.71 6.111 L"L. 0.65 •. 61 0.16 o.n 0,'0 O.lI0.01 0.25 O.OJ O. l~ 0.11 .." 0.19 o,n 0.110.01 •. n 0.11 D.Ol 0.01." O.U .." 0.2~ ..• 0.0' G.n 0,1' 0.01 0,"0... 0.16 ..~ o.n 0.1' 0.1I 0.05 0.31 0.1' 0.10 0••0 o.un .., n.1l 5.51 J.16 ,.w ..~ '.31 '.61 0.51 L~0.11 2.12 ..~ 0.21 O.OJ"60 .5.12 10.52 20.'5 ".111 n,60
r.~lo VI·A. _ry" dolol. I...."" ..' (oUr ...1.. , ... lo.oM ,..- 1:. nr,rJ•.u••oul lipid ..a ..._~ ..U'I..... eo.t....l _ Upcr'-a'o' ,. -.LIol~
COn .....1 Exporl_nUI ;::~:::••:o,..",_=...:::.::;u, u-.-,.-,-..-.,-,.-..-., hI'. I £>:1'.1
Table VI-B. Unusual non-llethyll!ne-interrupted fatty acids isolated frOli C. vlrglnlca
total lipid and various lipid fractions with and without feeding on
D. tertJolecta and after six .onths stonae.
Cra51f~tr.. vlrglnlca, Control and Experi_ental Fed Dunal1ella
Exp. t
Initial
Exp. II Exp. I
6th .onth
Exp. It
Control Experi.ental Control Experi_ental Control Experl-ental Control Experi_ental
Total Lipid
(20:2s) 1.00 0.82 1.01 0.98 0.54 1.18 2.09 1.49
(22:2s) 4.57 2.11 0.87 2.55 0.73 0.58 1.82 1.55
Polar Lipid
(20:2s) 0.86 0.92 0.15 0.63 0.45 2.19 0.86
(22:2s) 6.32 2.90 6.70 5.49 1.45 1.13 2.88 1.49
Trialycerlde
(20:2s) 0.61 2.12 1. 79 1.59 2.16 2.11 2.40 2.43
(22:2s) 0.28 1.71 1.11 0.78 0.61 0.89 0.90
Steryl Esters
(20:25) 0.71 1.02
(22:25)
blllrVll. "-or.11 lin I... 1 f"lly"c:ld, I ~n'''trd rroo D. ftd"l1_ tOUI HI'
llltnM ,'(/"/f,,, C"lItr,,1 "nd "_I'rd"'1l1,,1 1.,,1 '."Chl"t/.lllnutl llIc .... t,.rllI
TOI:tl 1,ll,ltt l'e,I" .. 1.I['id
bl'f'ri ..nlltl "al><' r ~..It t n I hrcrl ..nt,,1 1_I>rr.-rnt:tl
Control r", lllochr'ldll rrd Dlcrd.rJ. rt'd Uochr\lsb fC'd Dlcuted.
I~ ,0 1.67 15.24 10.90 S.1l 1.02 7.01 I.OS4.11.12·nrrn 0.11 1..10 0.97 0.9S 0.70 0.47 0.69
". 1"'0 0.11 I. ... 1.47 0.114 0.92 0." 0.17"nld~" -15:0 O.M 0.82 0.52 1.44 0.71 0." 0.1315,/1 5.59 fl.S6 (;.71 \.\7 '.06 3.'] 2.06I-·nll,-O o. 0.57 n.65 1./.7 1.58 I.,. ....J.I',III.I .. m'nIf,,O 'I .... \9.31 31.75 ](1.01 25.12 27.19 01:.221...'-17,0 2." 1.69 l.OS 1.42 1.16 1.56 I.fl9Anl,·j ~.. 17:0 0.64 1.9] 0.71 1.111 4.61 0.71 0.6511:0 4.61 \.62 5.17 2." 2.19 2.03 \.Ott\.7,ll.15-TNllt' ..."IIt:O 1.10 0.10 2." 0,47 0.40 0.46 0.29111:/1 19."1 10.67 20.17 II.lIf, 12.24 12.'5 7.4(1I~"l 0.011 0.10 n.1I 0.01 n.1210'0 0.49 0.16 0.21 0.60 0.11IJ,II 0.0\
".'Inr:tlrd
1.. ,11.. ,1..9 L5 0." 1.92 1.42 4.39 1.70110'1 ..7 :.\." 4.02
l3.:\7 :2.\6 :2.31 : •. 695.95
Ih:I,,5 0 .... 0.1711:1 •.117111;1 .." ""11) 1.11 1.111 1.4~ 2.49 ..... 5.61 5.411,tI,1 ..1 I." 1.14 1.10 1.21 0.79 1.23 5.91111:1",5 Ira!;: 0.06 0.04 0.20 0.15 0.291!1:19 0.10 0.4\ trllce n.'" 0.15~O; 1.. 11 0.47 0.]5 0.27 0.'" 1.22 0.83 0.51~n: 1..9 0.17 0.-4\ 0.\1 0.72 0 .... 0.76 O.H'n.I ..7 1.73 0.18 n.1I> 1.(17 I.foil 2.01 2.511211:1 ...... 0.16 0.10 O.()(, 0.\0 0.1. 0.23 0.4012,1 .. 11 r·..I\l O.l' O.M 0.02n;I.~) 0.19 II. nil O.U 0.11 0.1]lZ:I",1 0 .• ' I. Ito l.2S 1.95 0.62
12.(,5
1/,:1,,", 0 ..\2 0.46If,:.\..1 O.IUl 1l.19110:]..,11/,:4.• 1 0 . .(7
1ft: ~..", O.H 0.011 0.26 0,82 2." 3.91 1.\1IM:J.'" 0.29 0.79 n.4~ O.(,~ 0.01 11.0\IH' "'~, 11.29 U.118 n..!~ n.os O.OK 0.""III: ,,~, l.h.l 0.81\ l.4b ~ •(of, >\.20 2.83 l.IOIH:>\.d 0.92 O. 2~1 0.\.' I.HII 1.03 0.85 0.61
2U,1.", (. _~.9 I.llt, 1.32 0.01
211: "'~, O.W 0.0\ 0.01
/!I''-' 0.411 n.19 0.:\1 0.24 0.30 0.01~II:"-"h 0.11 O.1l7 O.IIS ~.bll 1.93 1.41 0.45'Ihl,,':\ ."" 0.11 O.lIl 0.12 O.OS 0.11
':11:5...:\ "4' n.lO 10.21 6.1>2 b.45 2.16
21 ~. ! 0.02l.l',I,.(, 0.0921;5.·,(, 0.27':2: ~....1 o.n 0.l4 0.11 0.1l 0.02n,r_' O_Sf. '.2') 1.7<1 3.78 O. 9~
1\,1 ~'-lU' l2.:\2 21.21
l·ol,·"I'''-,1 l.V, n."s
:and ..... riOU5 Ilr ld rnetluM with lind without reN.nllln '" ....lb"'Mllnd .. JnorIYU.
Oller" "",,'f., co." 1 • ,tdflpt'rl.enUI Ff-d r.odlrv"' .• .ndDt"r.tllrJ.
Tr1Illyl'C'rldf- Fn'C' Flllty Acld~ Stl',-"I F..HI'r5
I'~perl-ent.1 I'xl'~rl.entlll IXI'l'rl-entnl hperi.C'nllol hllllrl_nul l:xpl'rl-ent:>1
r,d r"ochrllsJ. fed Dlcnt.rI. Control 'od t.ochr~"fll fed Dlcrater'. fed rllochrllsh fedDJcrllterlll
5.1S '.00 6.91 1.35 4.42 12.13 2.42 11.360.4' 0.12 1.56 0.08 0.65 1.\9 0.68 1.490.69 0.94 0.79 0.12 0.61 1.87 0." 1.590.13 0.11 0.25 0.05 '.30 0.72 0.21 0.571.71 1.88 12.81 0.69 2.96 5.16 1.97 2.730.42 0.67 1.12 0.48 1.14 2.97 0.85 1.35
18.01 40.49 15.12 31.04 37.12 35.86 31.69 30.941.111 1.43 1.55 1.78 1.78 1.41 2.115 1.79O.M 0.51 0.53 1.07 0.54 1.00 2.18 1.5t.2.11 2.56 4.78 '.40 11.93 1.06 l.85 0.7'
0.25 0.25 0.96 0.24 0.43 0.'1 0.50 0.331.15 1.59 1.'0 21.26 11.'1 5.73 6.13 s...0.01 0.01 0.14ll.l\ 0.19 O.B 1.79 0.99 0.68
0.491
M.1I4 15.21 68.35 15.11 10.6\ 52.95
1.11 1.84 1.16 0.63 1.01 5.11 5.03 3.806.22 6.14 3.81
:0.952.68 5.11
:7.959.38
0.50 0.61 0.19 0.11 0.47 0.45.... 4.11 5.21 4.89 3.19 6.91 9.81 6.196.42 7.37 3.69 3.26 3.68 2.19 9.05 6.350.16 0.20 0.190.23 0.040.50 0.47 0.10 1.54 I.M 0.74 0.90 \.030.26 0.40 1.05 2.61 1.04 0.63 0.90 1.14::.73 2.31 1.28 8.46 3.64 1.25 3." 0.980.29 0.25 0.23 0.21 0.07 0.49 0.21
0.11 0.380.09 0.12 0.05 0.11 0.03 2.79
0.42 0." 0.42 0.38 0.41
23.86 2<>.04 22.81 17.10 22.80
0.35 2.13 0.53 0.16 0.64 1.05 0.170.25 0.30 0.24 0.47 0.43 0.26 1.080.03
1.85 l.25 0.31 0.08 0.52 2.110.12 0.07 0.02 0.010.06 0.04 0.031.84 1.08 2.60 4.62 0.77 1.01 0.84
1.26 0.51 0.07 0.04 0.15 0.39 0.2<>
0.05 0.010.04 0.06
1.11 1.20 1.630.07 0.02 0.120.76 0.54 0.24 0.35 0.64 0.73 0.24
0.31 0.072.216.16 2.2<> 0.11 0.13
0.092." 0.18
16.08 9.04 8.82 6.58 5.62
48.29 30.06 41.90 31.81 59.10
ndvari(ll,l5 lipid fraction,wnh.nd ..ithout feflhnlonr'9'1~andZl.J""'T1IIIU.
o.t ... , ""..Jh. e.:.ntrol and hperl.-nul Fed rloehryah and DJcTlt.r/,
Free fatty Acid5 Steryl Euen
5.11 1.00
1.71 I.U
S1.SJ
23.7. 22.93
1.61
6.97 1.3S 4.U 12.83 2.42 11.:56
12.81 , 0.69 ,." 5.16 1.117 2.71
33.26 42.64 1S.1l3 44.79
4.71 0.7'
37.23 26.15 16."
6.44 7.53 6.52
0 . .(9 0.93 3.16
Table.\'IJ·A. ~ry of chain lenJths of fatty atid, holaad f". o. -.;full_ total
_1.0-.. eduJia. Control and ~ri_nUI 'od Z_odIryai_ and Dierauria
Toul Upid Polar Lipid
Exper1_nul Experillenul Uperiwnul Experillental
fed laochry.Ja fed DJeTatada Control red laoc:hryai. fed DJer.tad. COntrol
'I< 8.67 15.24 10.90 5.83 7.02 7.08 8.05
'IS S.Sg '.SO 6.71 3.37 3.0. 3.83 2.06
'" 37.64 46.28 34.27 32." 33.0!l 50.67
'" 3.62 5.77 2.36 2.8g 2.03 3.06
'" 26.80 2S.S6 22.73 26.03 27.S6 22.22
'" 0.10 0.51 0.10 0.11 0.52 0.15 0.12
'"7.67 3.gl 3.'" 17.37 13.34
'" 0.02
'"l.48 0.02 0.22 '.00 3.60 S.g8 1.71
"''1. VIII No_I ••1_' ,.", .<ld. 1001> f t'. .'r"al.......1 IIF'd. """ ••d .... lIF'd fTO"I .... vlt~ 0lI<l vl._. food'"1 .. I _ID.I_••
C• ..-u_ ....,.._. e-.....I -.I lqa,l_ul ,0<1 ,_....,. _III~ ,.. c.._u.. 0'''''0''', e-...., oaoI ~.,.'I .....l fod I~.'......111 '.
~d_••1 l.,.rl_..1 bpnl_UI 1""'MOtol --==::-:--==::;_ ..... , fod '-.,.,10 fod 01.......". _.",1 fod '_I. fod ~.......,I. _.",1
',05',U
I.U__
,d_"".,I',,'·~
..10....11,.
1.1I.IS·'"
• u,...
,.....•.U
,.'.S1
,.1.1•,.
...)
..'.tI
'.1',.'.tI'.1'
...'.1.
...)
..,..."U.tS
I.U•. U
'.ff...)
....,.•. ll
,..''."•. 01'.OJI .•'
• .•1
o.n
,~...,
_"'0' thai" I.III.h. 01 ,.ny acid. 1101..... ,.,.. C. ,bll'J~I"," ....1 lipid........1..... lIpl. , ....1""••I,h wI,,,,,,,. 'oodl"l 00 I. ",'kno , D. 1_......
e............ ''',''''",", <'- ....1 ..... txpo,l ...tal F" 1__10 """ Nera.ul. cr._or.. ""Imea COO I oM to...I...... ' , .. ro_rya.lo &IWI1tl«ratul.
to l...to' (.,.,I tal (","1'1"'''1 (>po.I 1 tqo.l.....1 tJlparl ...tol f_,I .....1 txporl 1
COol.... I f _I. f .. Dle I. Cooo....1 F'" ,_J. '... DI"' I. COol .... ' F... ,_b '''0''''''.''. C>a ....1 F... ,_Ja F....J ··,,·
'abl. VIII-I. UnU~UIlI non-_thyIVle-inuTT\JPud fatty .dd~ holaUd he.
C. virgJnJI:. total lipid~ and vanous lipid frattiOll~ .,Ith and
.,i thout feedin. on 1. ~lhanlo and D. inorlWlt..
Cr...._er.. .,Jrgillic., COIltrol and ~eriM1lUI fed l~brll.j. and Di~At.,.j.
ExperiM1lUl ExperiM1lul
Control Fed J.oebry.i. Feel DJer.tu-J.
Total Lipid
(20:2~) 2.31 I." 2.02
(22;2~) 2 .•' •• OJ 2.13
Polar Lipid
(20;2~) 1.19 1.27 1..6
(22:2~) '.09 6.39 6.63
Tri.lytulde
(20;2~) 2 .• 2 1.87
(22: 2~) 0." 0.67 0."
Suryl Ener~
(20:25) 2.53 en(22:2~)