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OCR for page 200
JOEL BITMAN
Status Report on the
Alteration of Fatly Acid and
Stero! Composition in Lipids
in Meat, Milk' and Eggs
Demonstration of the high positive correlation between saturated fat
intake and heart disease (Figure 1) and between blood cholesterol
levels and heart disease (Figure 2) has made the American consumer
wary of the fat in meat, milk, and eggs. The percentage of fat con-
tributed by these food groups over a 20-year period is shown in Table 1
(Economic Research Service, 1965; Agricultural Statistics, 1969~. A
50% decline in butter consumption and 20% decline in egg consump-
tion were especially meaningful trends. The American consumer has
maintained a high consumption of beef, but that portion that is fat
is an unwanted obesity-inducing nutrient. Thus, in 1973 approximately
2.5 billion pounds of excess fat, valued at 1.15 billion dollars, were
trimmed from beef carcasses (Hendricks, 19741. The animal scientist,
reluctantly and belatedly, has finally recognized and accepted this
message from the marketplace.
Within the last few years, research has been directed towards altering
animal fats to make them more acceptable to the consumer, i.e., to in-
crease the polyunsaturated fats in meat and milk and to lower the
cholesterol in eggs. The task that faces the scientist who wants to change
the lipid composition of the ruminant is considerably more difficult than
it is to change the body fat composition of the nonruminant. In mono-
gastric animals, such as man, pig, and chicken, body fat can be changed
readily by changing the composition of the diet. Many experiments
200
OCR for page 201
Alteration of Fatty Acid and Sterol Composition in Lipids 201
40
20
20
10
_ CHD DEATH RATE
PER 1000
n n n n
~ CALORIES FROM SATURATED FATS
~n 11
GR JAP NETH FIN
YUGO IT US
FIGURE 1 Coronary heart disease deaths
and percentage of total calories provided by
saturated fats in the diet of men from seven
countries (Keys, 1970).
have demonstrated that if higher levels of dietary polyunsaturated fats
are fed to pigs and chickens, these polyunsaturated dietary lipids will
be absorbed and incorporated into body fat. In ruminants, however, if
increased amounts of polyunsaturated fats are fed, they are utilized
by microorganisms in the rumen or metabolized by these organisms to
form saturated and mono-unsaturated fatty acids; as a result, the meat
and milk fat do not show any increase in polyunsaturated fat. The data
in Table 2 demonstrate that although the normal plant diet of the
ruminant is primarily polyunsaturated, both meat and milk fat normally
TABLE 1 Contribution of Fat from Various Food Groups a
Percentage of Fat
Contributed by 1947-19491968 Change (% )
-
Meat, including fish 33.5%35.2% +S
Milk and dairy products,
including butter 21.6%17.1% -20
Eggs 4.3 %3.4% -20
Total 59.4%55.7% -6
a Data from ERS, 1965; and Agricultural Statistics, 1969.
OCR for page 202
202
o
o
to
Cal
LL
JOEL BITMAN
200
50
00
50
FIRST MAJOR CORONARY EVENT
_ MEN 40-59
~ :,n,n,n,J,Q,:
175 200 225 250 275 300
SERUM CHOLESTEROL mg 96
FIGURE 2 Relationship between serum cholesterol level and incidence rate of
the first major coronary event (Intersociety Commission for Heart Disease Re-
sources, 1970).
contain only 2%-4% polyunsaturated fat (Hilditch and Williams,
1964; Bitman et al., 1974b).
Recently, a process was developed that represents a breakthrough
in the attempt to increase the polyunsaturated, fatty-acid content of
TABLE 2 Fatty Acid Composition of Pasture Grass and Bovine
Milk and Meat Fat a
Weight in Lipid ( % )
Fatty Acid Grass Milk Meat
Myristic 14 01 12 3
Palmitic 16:011 31 26
Stearic 18:02 11 14
Oleic 18:15 24 47
Linoleic 18:212 3 3
Linolenic 18:362 1 1
Others 7b 18c 64
a SOURCE: grass (Hilditch and Williams, 1964); mild and meat fat (Bitmap e' al., 1974a).
b Primarily 12:0 and 16: 1.
c 4:0-12:0 comprise, 11%; 14:! and 16:1, 4%; minor acids, 3%.
Primarily 16:1.
OCR for page 203
Alteration of Fatty Acid and Sterol Composition in Lipids 203
ruminant meat and milk. T. W. Scott and his colleagues in Australia
(Scott et al., 1970) coated polyunsaturated oils with a protein and then
protected these particles from microbial attack in the rumen by treat-
ment with formaldehyde. The coated oils passed through the rumen
(pH 6-7) and into the abomasum and omasum (ply 2-3), where the
more acid conditions hydrolyzed the protein-formaldehyde coat, re-
leasing the intact dietary polyunsaturated oil, which could then be ab-
sorbed and incorporated into body and milk lipid. The process had
earlier been utilized by Ferguson et al. (1967) to increase wool growth
of sheep by protecting dietary casein from microbial degradation in the
rumen. This innovative technique has been the subject of intensive
research within the last 4 years and has, for the first time, provided a
range of new polyunsaturated ruminant foods.
An attempt has been made in this review to summarize research di-
rected towards altering the fatty-acid and sterol composition of meat,
milk, and eggs. The typical fatty-acid composition of the foods of animal
origin is shown in Table 3. This is the base upon which the experimental
alterations have been made. It can be seen that ruminant fat contains
more saturated fatty acids and less polyunsaturated fatty acids than do
swine and poultry fat. Of the three major dietary components fats, pro-
teins, and carbohydrates most attention has been given to the effects of
feeding fats upon the fatty-acid and cholesterol composition of the
animal. The tables are intended to group typical experimental findings
and are not necessarily comprehensive and complete.
TABLE 3 Typical Fatty-Acid Composition (Weight Percent) of Fat
from Different Animal Sources a
Ruminant Nonruminant
Fatty Acid Milk Beef Pork Poultry Eggs
Saturated
Lower C`-C1. 11
Myristic 14:0 12 3 1 1 1
Palmitic 16:0 31 26 25 25 23
Stearic 18:0 11 14 14 4 4
Unsaturated
Palmitoleic 16:1 4 3 3 7 5
Oleic 18:1 24 47 47 43 47
Linoleic 18:2 3 3 8 18 16
Linolenic 18 :3 1 1 - 2
Others 3 3 2 2 2
a Data from Hilditch and Williams ( 1964) and Bitman et al. (1974a).
OCR for page 204
204
A SYNOPSIS OF ALTERATIONS IN THE
LIPID COMPOSITION OF SWINE
JOEL BITMAN
A summary of the changes brought about in fatty-acid composition by
experimental dietary alterations in swine is presented in Table 4. The
data clearly demonstrate the ready capacity of the pig to store fat of
the type present in its diet. A wide variety of plant and fish oils, con-
taining large quantities of polyunsaturated fatty acids, were fed to pigs;
and the polyunsaturated fats were promptly absorbed and incorporated
into their body fat. Most of these studies were conducted with growing
pigs (8-28 weeks); end the depot fat alterations, although rapid, re-
quired several months to achieve equilibrium. Experiments during the
twenties and thirties, in which polyunsaturated fats were added to the
diet, produced a soft or oily pork, characterized by a high linoleic ( 18: 2)
and a low palm~tic ( 16: 0) and low stearic ( 18: 0) content.* Consumer
acceptability was poor.
While Table 4 may appear to indicate that alterations in the fatty-acid
composition of swine have been well explored, I would suggest that this
table, based upon approximately twenty reports, instead shows that fat
~ The first number refers to the length of the carbon chain in the fatty acid; the
second, to the number of double bonds in the chain.
TABLE 4 Fatty Acid Changes in Pig Fat a
Diet 18:2 18:1 18:0 16:0 16:1 20-22 References
Plant Lipids
Corn, corn oil
+
67, 68, 69, 71, 82,
123
Soybeans,
soybean oil + - - - 16, 68, 69
Peanuts + - - - 68, 69
Cottonseed oil + - + - 70
Fish Lipids
Menhaden oil
Whale oil
Cod liver oil-lard
Animal Lipids
Tallow
Cholesterol
VFA
High carbohydrate
Vitamin D
Copper
17, 18
83
+ + 84
O O O O
+
+ +
O + - O
123
113
14
66
113
2, 30, 142, 212
a CODE: ~ = increase,-= decrease, and 0 = no change.
OCR for page 205
Alteration of Fatty Acid and Sterol Composition in Lipids 205
modification research in pigs is relatively limited when compared to the
number of studies on the role of fat in cardiovascular disease, where
literature references run into the hundreds. The advent of modern
gas-liquid chromatography has made investigation in this area much
more practicable. Thus, the mechanism for control of fat composition in
swine would seem to be susceptible to more exact elucidation. There are
some additional, scattered literature references that indicate that several
other factors affect fatty-acid composition (sex, age, breed, starvation,
temperature), but this limited information was not included.
A SYNOPSIS OF ALTERATIONS IN THE LIPID COMPOSITION
OF POULTRY DEPOT AND EGG LIPIDS
Summaries of the changes that can be effected in fatty-acid composition
of poultry depot fat and egg lipids are presented in Tables 5 and 6.
Several major features are apparent:
1. Alterations in egg lipids are complete within 16 days, while
depot fat changes are much slower, requiring several months to reach
an equilibrium. This difference reflects ovum maturation time in the
egg-laying cycle and a relatively rapid transfer of blood lipids to a small
fat compartment (egg lipids) in contrast to a much slower balance
between blood lipids and a large lipid compartment (depot fat).
2. The chicken stores fat of the type present in its diet: dietary
saturated or unsaturated fat causes the deposition of lipids of that re-
spective type in the depot fat.
3. Dietary unsaturated fats pass readily into egg lipids, thus ingestion
of a wide variety of unsaturated plant oils results in the appearance
of the characteristic unsaturated fatty acids in the egg. Dietary saturated
fats, however, have relatively little influence upon the composition of
egg lipids.
4. An approximate inverse relationship exists between linoleic acid
and oleic acids in egg and tissue lipids in response to dietary unsaturated
fat ingestion. Particularly because of the marked resistance to change
of the saturated fatty acids in egg lipids, most compositional changes in
response to dietary fats occur in the relative proportions of 18:2 and
18:1.
To the extent that generalizations between species are valid in these
two monogastric animals, the chicken and the pig, it is apparent that
the body fat of swine and poultry will reflect either greater saturation or
unsaturation, depending upon the composition of the diet. Egg lipid
composition, however, can be altered readily only in the direction of
more unsaturated lipids.
OCR for page 206
206
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OCR for page 208
208
SYNOPSIS OF ALTERATIONS IN EGG CHOLESTEROL
CONTENT
JOEL BITMAN
Table 7 presents a summary of factors that can alter egg cholesterol
content. Such limited studies as have been carried out on age and season
as factors influencing egg cholesterol have not been included. A large
number of studies dealing with changes in serum cholesterol and liver
cholesterol in poultry, but which do not contain data on egg cholesterol,
are not discussed in this review.
Cholesterol, like the other lipids in eggs, can be altered by dietary
means. Studies with labeled acetate (Kritchevsky and Kirk, 1951,
Kritchevsky et al., 1951), cholesterol (Andrews et al., 1965, 1968;
Connor e! al., 1965), and triglycerides (Budowski et al., 1961) and the
experiments with cholesterol inhibitors in which desmosterol builds up
in the egg (Burgess et al., 1962) have demonstrated that changes in
egg cholesterol occur within hours of treatment. The time course of egg
cholesterol changes thus appears to agree well with the time course
of changes in egg fatty-acid composition (Reiser, l951b).
Data from studies with dietary lipids disclose a large number of un-
certainties. Thus, there is no agreement on whether or not corn oil,
safflower oil, linseed oil, soybean oil, or coconut oil will increase egg
cholesterol. Use of almost every oil at the same level by different
workers has yielded different results. At the present time there is no
good explanation for these serious discrepancies in research results by
reputable, competent scientists.
The inclusion of cholesterol in the diet of the hen promptly causes
increased amounts of cholesterol in the egg. Addition of fat to the diet
along with the cholesterol rather uniformly produces a doubling in egg
cholesterol, probably by increasing the absorption of dietary cholesterol
in the gut.
Agents that influence (a) the intestinal absorption or (b) the
enterohepatic circulation of cholesterol alter egg cholesterol. Thus,
surface-active agents such as Tween or lecithin improve cholesterol
adsorbability and increase egg cholesterol. Conversely, the plant sterol,
,3-sitosterol, promotes the fecal excretion of cholesterol and conse-
quently decreases egg cholesterol. Sitosterol quantitatively replaces
cholesterol in the egg.
Two published reports, which disagree, are inadequate to determine
whether dietary fiber reduces or increases egg cholesterol. The few
studies with vitamins do not demonstrate striking or consistent effects.
D-thyroxine was found to increase egg cholesterol, apparently by stimu
lating cholesterol turnover and excretion via the egg.
OCR for page 209
Alteration of Fatty Acid and Sterol Composition in Lipids
TABLE 7 Effect of Dietary Agents on Egg Cholesterola
209
Group Agent Effect References
Oils
Corn
Safflower
Linseed
+
o
39
32,63,80,122,143,
158, 220
7, 80, 209, 219, 220
220,222
80, 219
222
Soybean + 80, 209
0 7,51
Coconut + 7, 220
0 32
Cottonseed 0 167
Lard 0 32,63
Tallow 0 39, 51, 63, 80, 122,
158, 220
Rapeseed 0 122
Sterols Cholesterol + 32, 52, 53, 60, 64, 90,
102, 135, 182, 220,
226
0 121, 136
Sitosterol - 36
0 220
Protein 0 39, 63, 143, 158
Surface Active Tween 0 220
Lecithin 0 220
Cholestyramine 0 111, 134
Tween-cholesterol + 220
Lecithin-cholesterol + 220
Fiber Cellulose + 135
Cellulose - 216
Pectin - 216
Vitamins Niacin 0 220
Vitamin C 0 155
Vitamin A + 182
0 60, 220
Drugs Clofibrate 0 220
MER-29 - 25
Azasterols - 184
Diethyl-aminoethyl
diphenyl valerate - 146
Probucol - 146, 147
Hormones D-thyroxine + 220
a CODE: + = increase,-= decrease, and 0 = no change.
OCR for page 210
210
JOEL BITMAN
Drugs that inhibit the biosynthesis of cholesterol at the reductive
step in the pathway from desmosterol to cholesterol have been successful
in lowering egg cholesterol, but there is an accompanying quantitative
replacement of cholesterol by desmosterol. This alteration raises several
questions:
1. Are eggs containing large quantities of desmosterol satisfactory
as human foods?
2. What level of cholesterol is necessary in the developing ovum to
permit satisfactory egg production?
3. What levels of desmosterol can the chicken successfully cope with?
What are the long-range physiological consequences for the hen of in-
creased levels of desmosterol?
A number of studies have demonstrated that egg cholesterol concen-
tration varies genetically. Eggs from broiler-breeder strains contained
more cholesterol than did those from commercial layer strains (Edwards
et al., 1960; Miller and Denton, 1962; Harris and Wilcox, 1963a; Col-
lins et al., 1968; Turk and Barnett, 1971; Marks and Washburn, 1973;
Cunningham et al., 1974; Washburn and Nix, 1974~. Whether these
differences are large enough to be nutritionally and physiologically
meaningful to humans has not yet been determined.
Many of the substances that have been used in attempts to alter egg
cholesterol are agents that reduce serum cholesterol in other species.
Review of the serum cholesterol changes of the studies summarized in
Table 7 indicated that there was no simple direct relationship between
plasma and egg cholesterol concentration. Thus, both D-thyroxine and
,3-sitosterol lowered blood cholesterol; thyroxine increased egg cho-
lesterol, while sitosterol lowered it. Cholestyramine caused a very large
decrease in serum cholesterol but had no effect upon egg cholesterol.
Feeding dietary oils or oils with cholesterol raised serum cholesterol
and also raised egg cholesterol.
A diagrammatic representation of major cholesterol compartments
of the laying hen is shown in Figure 3. The lack of consistent results
and the lack of understanding of cholesterol relationships in the laying
hen suggest that the time has arrived for complete metabolic balance
studies in egg cholesterol reduction research. Although studies of this
type would be expensive, the many studies listed in Table 7 attest to
the large amount of money already expended on this problem. Measure-
ment of cholesterol levels in the diet, plasma, fat, liver, body, egg, and
excrete; the use of labeled cholesterol; and gas chromatography to
identify egg sterols could bring order to the cholesterol picture order
OCR for page 227
Alteration of Fatty Acid and Sterol Composition in Lipids 227
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OCR for page 228
228
JOEL BITMAN
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OCR for page 229
Alteration of Fatty Acid and Sterol Composition in Lipids 229
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Representative terms from entire chapter:
fatty acids
