Characteristics of the fatty acid composition and biochemistry of some fresh-water fish oils and lipids in comparison with marine oils and lipids

Comp. Biochem. Physiol., 1967, Vol. 22, pp. 907 to 922. PergamonPress Ltd. Printed in Great Britain

CHARACTERISTICS OF THE FATTY ACID COMPOSITION AND BIOCHEMISTRY OF SOME FRESH-WATER FISH OILS AND LIPIDS IN COMPARISON WITH MARINE OILS AND LIPIDS R. G. ACKMAN Fisheries Research Board of Canada, Halifax Laboratory, Halifax, Nova Scotia, Canada (Received 20 March 1967) A b s t r a c t - - 1 . T h e fatty acid composition of oils from four North American fresh-water fish (sheepshead, Aplodinotus grunniens; tullibee, Coregonus artedii; maria, Lota lota; alewife, .4losa pseudoharengus) were compared with recent data for oils from two marine species (Atlantic herring, Clupea harengus; cod, Gadus morhua). 2. In the oils from fresh-water species the total Cte fatty acids were higher than in the marine species. T h e total Cxs fatty acids were also higher but possibly less definitive as a means of distinguishing fresh-water triglyceride oils from those of marine origin. 3. Ratios among particular fatty acids and among various types of fatty acids were compared for biochemical significance. Palmitic acid was about 60 per cent of total saturates in both fresh-water and marine oils. Total di- and tetraenoic acids were twice as high in the fresh-water oils as in the marine oils; total trienoic acids were three to four times as high. It is suggested that extension of these to the marine-type fatty acids 20:5 oJ3, 22:6 oJ3, etc. is not normally obligatory in fresh-water fish. 4. The ratio of total linolenic to total linoleic types of acids was lower in the fresh-water oils, suggesting a basic difference in dietary availability of these two acids.

INTRODUCTION

THE DISTINCTIOrqbetween fresh-water and marine triglyceride fats has been defined primarily in terms of fatty acid chain length. The habitat was not ignored but the broad differences in the fat types were "that in the fats of fresh-water animals C16 and (especially) Cls acids are present in greater proportions, whilst C,o and (espeoially) C~, acids are present in smaller proportions, than in the fats of similar mar!ine animals" (Lovern, 1942; el. Lovern, 1937; Hilditch & Williams, 1964). A further "boundary" class offish fats had intermediate properties. Basically the differences in fish fats were ascribed to dietary fat intake rather than the habitat per se, but the amplification of the above trends in tropical and subtropical species was noted. A gas-liquid chromatographic examination of the fatty acids of the "oils" from four North American fresh-water fish has recently been carried out in connexion 907

908

R.G. ACKMAN

with commercial meal and oil production (Ackman et al., 1967a). This and other detailed data on fresh-water oils and lipids are compared with recent data on two marine "oils" to ascertain basic differences, particularly in depot fats. EXPERIMENTAL Sheepshead (Aplodinotus grunniens) were taken in October 1965 at the western end of Lake Erie. Tullibee (Coregonus artediz) and maria (Lota Iota) were taken in October 1965 at the south-east end of Lake Winnipeg. Alewife (Alosa pseudoharengus) were taken in September 1965 in Lake Michigan near Milwaukee. The "fat" contents of the fish as determined by a standard method (A.O.A.C., 1960) and the "oil" yields from normal commercial reduction (Ackman et al., 1967a), are given in Table 1. TABLE 1--FAT

Fat (%) Commercial oil yield (%)

CONTENT AND O I L RECOVERY ON W H O L E FISH BASIS

Sheepshead

Tullibee

Maria

Alewife

11"9 3"7

8"0 3-9

3"7 0-16

9"6 5"3

Gas-liquid chromatography of the methyl esters of fatty acids recovered after removal of non-saponifiable materials (Official Method of the American Oil Chemists' Society, 1965) was carried out as described elsewhere (Ackman & Eaton, 1966) with tentative identifications of several very minor components. Large components (Tables 2 and 3) are estimated to be accurate to + 5 per cent, moderatesize components to + 10 per cent, but very small components may be as inaccurate as + 50 per cent. Presentation of weight per cent composition data to two places of decimals is solely to show the relative levels of small components. NSA (no significant amount) indicates that the component was not detected, T R A (trace) indicates that the amount was less than 0.01 per cent. Totals of certain unidentified CIs fatty acids were respectively: sheepshead, 0.32 per cent; tullibee, 0.75 per cent; maria, 0.58 per cent; alewife, 0.54 per cent. DISCUSSION

Biology of sheepshead, tullibee, maria and alewife Sheepshead usually weigh 0.5-1.0 kg, but occasionally much larger fish are taken in the Great Lakes. They are voracious bottom feeders, feeding mainly on snails, other molluscs and crayfish. The sheepshead is a spring-spawning fish. Tullibee are also known as fresh-water herring and resemble marine herring. Three or four species, or subspecies, are known as tullibee in the central parts of Canada. They are heavy feeders; the main food being plankton, some insects, and minnows in spring and fall. They spawn in shallow water from October to December. The average weight is usually less than 0.5 kg. The maria is a roaming, heavy-feeding fish spawning from January to March. It is classified as carnivorous, feeding mainly on small fish although aquatic insects,

F A T T Y ACID C O M P O S I T I O N O F S O M E F R E S H - W A T E R F I S H O I L S

909

crayfish and plankton are also eaten. The average weight is 4--10 kg. It has a large fatty liver like the marine cod. Although the alewife is primarily a marine fish, it is also found in fresh waters including the Great Lakes. In fresh water the average length is about 15 cm in contrast to about 25 cm in salt water. In the Great Lakes the alewife is found in open waters alad it feeds on insects and small crustaceans (plankton). Spawning may vary from late May to early August. In samples of fish examined in the course of the recent study only the maria showed signs of gonad development. The low fat content and oil yield from the particular catch of this species also set it apart somewhat from the other three species. In the tables the four fresh-water oils have been arranged in order of increasing iodine value and hence some trends in components and properties can be observed. The maria results show most of the exceptions to such trends.

Fats and fatty acid biochemistry in fish Note should be taken of the usage of the terms "oil" or "fat" to refer to an essentially triglyceride fat, with the term "lipid" including also phospholipids and implying recovery by an efficient solvent extraction process (Bligh & Dyer, 1959; Lovern, 1965). The fat content of several species of North American fresh-water fish, and the iodine value of the fat, show remarkable variations for one species from one body of water to another. Data on the four species of the present study will be found in Pugsley (1942), Schmidt (1948), Schmidt & Carter (1948), Schmidt (1949), Karriek et al. (1956), Thurston et al. (1959), Dugal (1962) and Travis (1966). Ecological considerations are therefore even more important in assessing fatty acids of fresh-water fish than of marine fish (cf. DeWitt, 1963). The samples for the present sturdy came from large and moderately warm bodies of water where an adequate and varied food supply might be assumed. The four species of fresh-water fish examined in the present study are essentially carnivorous when adult. A reasonable degree of integration of lipid originating in plants or phytoplankton through the lower life forms eaten by these fish may therefore be expected. Size can affect the total fatty acid composition of fish as shown for the mullet (Mugil cephalus) by Reiser et al. (1963). Both fat content and iodine values of fats of rainbow trout (Salmo gairdnerii irideus) of different sizes fed a standard diet showed obvious graduations (Toyomizu et al., 1963). Fatty acid variations with size may therefore not be due solely to selectivity in choice of food in the natural environment. The fish which were the source of the oils for the present study were all of commercial size and hence mature. Total component fatty acids of fish can be altered by manipulating temperature with results considered characteristic of most poikilotherms. Thus at lower temperatures there is generally an increase in the more obvious highly unsaturated fatty acids (Kayama et al., 1963a; Reiser et al., 1963; Farkas & Herodek, 1964; Johnston & Roots, 1964; Jezyk & Penienak, 1966; Knipprath & Mead, 1966a, b).

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In this type of experiment insufficient attention has perhaps been paid to the quantitative status and distribution throughout the fish of different basic lipids such as phospholipids and triglycerides. In relating these results to natural conditions for fres:h-water fish it must also be borne in mind that low water temperatures would correspond to a period of food scarcity for most species. It is also well known that total fish fatty acids can be altered within certain limits by varying the fatty acid composition of the food (Kelly et al., 1958; Mead et al., 1960; Brenner et al., 1961, 1963 ; Kayama et al., 1963a, b; Reiser et al., 1963 ; Toyomizu et al., 1963; Farkas & Herodek, 1964; Lovern, 1964; Vazza, 1964; Jezyk & Penicnak, 1966). The general conclusion to be drawn from these experiments is that fish require 18:2 co6 (or 18:3 06) and 18:3 w3 as "essential" fatty acids* (Nicolaides & Woodall, 1962; Higashi et aL, 1964, 1956), and that fish cannot synthesize these particular fatty acids de novo. When supplied with these essential fatty acids in suitable proportions fish perform chain extention by the usual metabolic pathways to give longer-chain polyunsaturated fatty acids such as 20:4 co6, 20:5 co3, 22:5 03 a n d 2 2 : 6 w3. The relative proportions of all of the fatty acids are kept in balance in normal fish on a varied diet as best suited to the species, environmental conditions, age, stage of sexual development and other factors (Loverrc, 1964). Phospholipids are the most essential lipid and should show the least variation in overall fatty acid composition in a given species (Lovem, 1964). This has been shown for marine fish with cod (Gadus morhua) from both sides of the Atlantic (Ackman & BuJ~gher, 1964b) and also in cod from a given population (Jangaard et al., 1966, 1967), although very large fish tended to be different in some details. A similar situation exists in fresh-water fish as shown by the study of wild and cultivated sweet smelt (Plecoglossum altivelis) by Shimma & Taguchi (1964b). Part of the sweet smelt analysis data is reproduced in Table 4. Minor variations in the polyunsaturated C20 and C ~ acids are reduced in significance if these acids are regarded as interchangable (Ackman & Burgher, 1964b) and totalled (Table 4). Jezyk & Penicnak (1966) have observed that the phospholipid fatty acids of brine shrimp (Artemta salina) in naupli and adult stages are more similar than are the neutral lipid fatty acids. A distinction which can affect even fatty acids of phospholipids is that of a prolonged period on a fat-free or low-fat diet (Brenner et al., 1963; Reiser et al., 1963). De novo synthesis of certain fatty acids (saturated acids and monoenoie fatty acids) is possible under these conditions, but it is not known if the proportions of these are temperature-independent as in the mosquito Aedes sollicitans (Van Handel, 1966). Johnston & Roots (1964) pointed out that in addition to the changes in fatty acid composition of goldfish (Carassius auratus) brain with temperature, the percentage of lipid in the brain dropped from 9-32 to 8.30 per cent over the temperature range * The shorthand notation of chain length--number of double bonds and number of carbon atoms from centre of ultimate double bond to, and including, the terminal methyl group--is used for particular polyunsaturated fatty acids.

914

R.G. ACKMAN

TABLE 4----PERCENTAGES OF SOME FATTY ACIDS OF PHOSPHOLIPIDS OF WILD AND CULTIVATED SWEET SMELT (SHIMMA & TAGUCHI, 1964b)

Female muscle

Male muscle

Milt

F a t t y acid

Wild

Cultivated

Wild

Cultivated

Wild

Cultivated

14:0 16:0 16:1

1"6 24"2 6-3

0"8 25"6 2-6

1-4 22"8 6"6

0-8 24"3 2"7

1"7 34"0 4"5 4"3 20"3 5"6 3-2 0"6 5-2 2-4 18"0 26"2

18 : 0

8"6

9"0

9"5

18:1 18:2 18 : 3 20:4 20:5 22:5 22:6

14"6 4"0 4-8 2-7 5-7 4-4 19"0

17"8 6"3 4-9 3-0 5"4 Trace 24-2

16"6 4"6 5"2 3"6 6-5 4"7 19"0

15"9 4"7 5"0 1"0 5-2 2"5 28"4

2-8 33"3 11"5 6"6 20"7 3-8 2-4 1"8 4"5 4"9 7"7

ZC~o+Cv,

31"8

32"6

33"8

37"1

18"9

9"4

5-30°C, while conversely liver lipid increased from 1.76 to 3.69 per cent. The ratio of phospholipid to triglyceride would probably be altered accordingly and influence total fatty acid composition. The inverse relation of iodine value of fatty acids and water temperature noted by Farkas & Herodek (1964) for selected fresh-water crustacea was correlated with increased C20 and C ~ fatty acids, presumably highly unsaturated, in three species of copepods. It was also shown that melting points of fats from the copepods were directly related to water temperature. This is an instance of adaption to environment by keeping extracellular fat fluid at lower temperatures. Reiser et al. (1963) showed that approximately 1 per cent 18:3 co3 (linolenic) acid in a fish diet increased the level of 22: 6 co3 whereas 5 per cent was deposited without this conversion. These authors also stated that fish in which the longerchain, more highly unsaturated fatty acids had been replaced by high levels of 18:2eo6 and 18:3 co3 converted these Cls acids to their successor acids when placed on a depletion diet. Although triglycerides and phospholipids were examined separately in some of this work the proportions present in the fish were not given. It is likely that fresh-water fish directly deposit considerable proportions of ingested Cas polyunsaturated fatty acids when feeding actively in warm water periods, but only convert these fatty acids in their depot fats to more highly unsaturated fatty acids of longer chain length when necessary as water temperature falls or food becomes scarce. Competitive inhibition among fatty acids (Brenner & Peluffo, 1966; Castor et al., 1966; Lindstrom & Tinsley, 1966) may be the factor restricting these conversions except when overuled by circumstances such as changes in environment or availability of food. As discussed below the diet of fresh-water fish, relative to that of marine fish, appears to be enriched in co6 acids as compared to oJ3 acids.

FATTY ACID C O M P O S I T I O N OF SOME FRESH-WATER FISH OILS

915

Differentiation offish oils of fresh-water and marine origin There may be wide variations in the percentages of individual fatty acids in different lots of commercial marine oils from the same species, either of the wholebody type (Atlantic herring, Clupea harengus; Ackman & Eaton, 1966) or fatty-liver type (cod; Dc;Witt, 1963; Lambertsen & Braekkan, 1965; Jangaard et al., 1967). Such samples average the composition for thousands of individual fish and are therefore more reliable than fats and lipids from a single fish (Jangaard et al., 1966). The four fresh-water oils examined in detail in the present study are good averages for these particular lots of fish only and too much stress should not be placed on comparisons ~Lmong the four fresh-water species examined in this study. Two species (tullibee and maria) came from the same lake, the others from different although interconnecting bodies of water. Environmental influences may possibly be different in the respective water systems. The basic comparisons to be made are therefore with commercial marine oils. Clupeids (herring) are processed for whole body oils, and a recent survey of Atlantic herring oils includes iodine values overlapping with the sheepshead and tullibee, while other clupeids provide limited data for oils of higher iodine values (Ackman & Eaton, 1966). Cod liver oil (Jangaard et al., 1967) also provides data for comparison with high iodine value fresh-water oils, and particularly for the maria which has a similar fatty liver. Hilditch & Williams (1964), following views of Lovern (1942), point out that the triglyceride oils and lipids from fresh-water fish, in comparison with those of marine origin, are richer in C16 and C1s fatty acids, with correspondingly lower proportions for C20 and C22 fatty acids. In some tropical species there are particularly low levels of C2~ fatty acids. In general these views are confirmed by recent studies on various subtropical and temperate fresh-water species (Brenner et al., 1961, 1963; Pathak & Reddy, 1962, 1964; Vazza, 1964). In a study (Shimma & Taguchi, 1964a) of fat from dorsal flesh of fresh-water Japanese fish some species such as sweet smelt and loach (Misgurnus fossilis) showed low levels of C20 and Cz2 acids. On the other hand, fat from the flesh of rainbow trout and pond smelt (Hypomesus olidus), and from whole whitebait (Salangichthys microdon), had higher levels of Cz0 and C22 acids more commonly associated with marine-type fats. These results were from samples containing relatively low (10.7-5.6 per cent) fat levels. Lipids of three North American species were examined by Gruger et al. (1964). Flesh of lake herring or tullibee, rainbow trout and lake whitefish (Coregonus clupeaformis) had 2.2-2.5 per cent lipid, and fatty, acids of these lipids showed a marine-type pattern of high levels of Cz0 and Cz2 fatty acids, but the samples may have been largely phospholipids (see below). Sheepshead and tullibee oil data support the chain-length view, being relatively high in C16 and ClS fatty acids, and particularly low in C2z fatty acids (Table 3). The maria and alewife oils do not fall into quite the same pattern, but maria oil was from relatively lean fish, possibly including some free fatty acids liberated from phospholipids. The alewife was originally a marine clupeid, naturalized to a

916

R.G. ACKMAN

fresh-water habitat, and moreover the oil must have a high proportion of one or more polyunsaturated C20 and C2z fatty acids to conform to the high iodine value since the contents of 18:4 co3 and 18:3 acids, the only alternative high-iodine value fatty acids, are not sufficiently different in the fresh-water and marine oils. The C16 and Cls chain lengths respectively averaged 35-5 and 35.6 per cent for the four fresh-water oils. The Cls fatty acids are higher in the cod liver oil than in the herring oil (see DeWitt, 1963 ; Ackman & Burgher, 1964a; Lambertsen & Braekkan, 1965 ; Jangaard et al., 1966, 1967, for other cod liver oil analyses) and therefore the C xe fatty acids are perhaps the better indicator of a fresh-water oil, since they appear to be more consistent in these triglyceride marine oils. Lewis (1962, 1965) has pointed out that with increasing depth of water there is a corresponding increase in 18:1 in the fats of marine organisms. In the depot (muscle) triglyceride fat of the sablefish (Anoplopoma fimbria), a marine species in which mature fish are commonly found at 100-300 fathoms, the percentages of Ct6 fatty acids in the total composition have been found by two laboratories to be respectively 19.8 and 28.4 per cent, and the ClS fatty acids 32.1 and 45.2 per cent (Ackman et al., 1967b; Dolev & Olcott, 1965). These findings also indicate that the Cxs fatty acids are less important than the Cle fatty acids in distinguishing between fresh-water and marine oils. A common feature of all of these fresh-water fish for which detailed data is available, excepting the pond smelt, is an obvious enrichment in linoleic (18:2 oJ6) and linolenic (18:3 co3) fatty acids, with percentages running approximately twice those of most marine species. This property extends to both phospholipids (Table 4) and triglycerides in the case of sweet smelt, and is not drastically modified by artificial rearing of this species (Shimma & Taguchi, 1964b). Generally the presence of high proportions of polyunsaturated Cls fatty acids is a more typical fresh-water lipid characteristic than are low levels of C~0 and C2~ fatty acids. The detailed fatty acid composition also indicates that 20:4 co6 (arachidonic) acid is much higher in fresh-water oils than in marine oils, but this acid occurs in marine phospholipids at higher levels than in the triglycerides (Olley & Duncan, 1965; Jezyk & Penicnak, 1966; Jangaard et al., 1967) so it may not be a basic characteristic distinguishing all types of fresh-water and marine lipids. The results of the present gas-liquid chromatographic study of fatty acids of commercial oils from four fresh-water species (Tables 2 and 3) are more detailed than data hitherto available. The previous analysis of tullibee flesh (Gruger et al., 1964) differs considerably in detail and chain-length totals. In particular the proportion of 22: 6 co3 is very high, and cannot be due to inclusion of gonads which might be relatively rich in this acid. Since the flesh lipid was found to be 2-5 per cent by Gruger et al., and fat was 8-0 per cent in the present analysis of whole fish, it must be concluded that the flesh lipid sample was particularly rich in phospholipids. Comparison of published data on triglyceride and phospholipid fatty acids for marine species (Olley & Duncan, 1965) and for the sweet smelt (Table 4) supports this view in some of the other details of fatty acids from the flesh analysis such as low level of 16:1 when compared to 16:0.

FATTY ACID C O M P O S I T I O N OF SOME F R E S H - W A T E R F I S H OILS

917

Total saturated fatty acids (Table 5) appear to be slightly higher in fresh-water oils than in average marine oils but are comparable to marine clupeid oils of higher iodine values (Ackman & Eaton, 1966). Monoenoic fatty acids are somewhat lower than in marine oils of comparable iodine values. This is a basic difference (cf. Ackman, 1966) following from the higher proportions of dienoic and particularly trienoic fatty acids in fresh-water oils, especially 18:2 co6, 18:3 oJ6, 18:3 oJ3 and successor acids intermediate in metabolic pathways to longer-chain fatty acids such as 20: 4 co6, 2,9: 5 oJ3 and 22: 6 co3. Tetraenoic fatty acids are also higher than in marine species, but this difference is not as marked in the totals since in marine oils total tetr.-tenoic fatty acids invariably include metabolically inactive 16:4 col. If this acid is excluded the totals are more significant, and the high proportions of 20: 4 oJ6 and universal occurrence of 22: 4 oJ6, which is seldom observed in appreciable amounts in marine oils (Ackman & Burgher, 1965), are noteworthy in the fresh-water oils. Total levels of pentaenoic and hexaenoic acids are not exceptional in the fresh-water oils since these are quite variable in different samples of marine oils. Total polyunsaturated fatty acids are somewhat higher in the fresh-water oils than in marine oils of comparable iodine value for reasons noted above. The key J~ole of 16:0 (palmitic) acid in saturated fatty acid metabolism in marine depot fats has been stressed previously (Brenner et al., 1963; Ackman & Sipos, 1965 ; Ackman & Eaton, 1966). In agreement with this view the fresh-water fish show the same proportions (about 60 per cent) as do other fresh-water and marine fish. In the fresh-water fish the higher levels of 16:1 and 18:1 give slightly lower ratios of 16:0/16:1 and 16:0/(16:1 + 18:1) than are observed in marine fish; but the ratio i[6:1/18:1 is essentially the same for the whole-body oils. The chain, extension (from Cxs dietary intake) ratios of fatty acids of the oJ6 (linoleic) series fall within the same range of values as found for the marine oils, but in the co3 (linolenic) series the ratios (excepting the maria) are lower than for the marine oils for both the 20:x oJ3/18:x co3 and 22:x ~o3/18:x oJ3 values, while the 20:x co3/22:x co3 ratios are of the same order as in marine species examined. The ratio of total oJ3 to oJ6 fatty acids shows a much lower value in freshwater species than in marine oils, indicating a diet richer in 18:2 oJ6 and 18:3 co6 than in 18:3 ,~3. In river-caught coho salmon (Oncorhynchus kisutch) fry, further fed briefly on a fresh-water invertebrate, the ratio of ~03 to co6 acids was approximately 1 (Saddler et al., 1966; data revised by Ackman, 1967). As a juvenile form of an anadromous fish this is a somewhat special case (Lovern, 1964; Stansby, 1967), but otherwise conforms to the characteristics suggested herein for freshwater fats. The overall summation of related ¢o3 fatty acids differing by the introduction of a new double bond closer to the carboxyl group is lower for fresh-water species. Probably this arises from the inclusion of a much higher proportion of 18:3 oJ3, instead of co3 :mccessor acids, in the diet of fresh-water species, leading to deposition of this acid in depot fat without further chain extension. As suggested above conversion to longer-chain fatty acids such as 20:5 oJ3, 22:5 co3 and 22:6 co3 is apparently not obligatory in fresh-water fish in the natural state.

918

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FATTY ACID C O M P O S I T I O N OF SOME FRESH-WATER FISH OILS

919

T h i s reasoning probably also applies to the oJ6, series, but in marine species the amounts of t]aese acids are too low to provide a reliable comparison. T h e consiideration of biochemical interrelationships a m o n g various fatty acids and types of :fatty acids of fresh-water fish have been based, in the case of sheepshead, tuUibee and alewife, on fall-caught fish p r e s u m a b l y in p r i m e condition for winter survival and not affected b y gonad development. Generally there was a coherent pattern for fatty acids of these three species, and frequently for the maria as well, although results for the latter m a y have been modified by the low lipid level and gonad development. T h e s e observations m a y apply only to fresh-water fish f r o m m o r e northerly latitudes, and further work would be required to establish their validity for fresh-water fish f r o m tropical and subtropical areas. Acknowledgement--The author expresses his thanks for the cooperation of Dr. E. G. Bligh of Fisheries Research Board of Canada, Freshwater Research Institute, University of Manitoba, Winnipeg 19, Manitoba.

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