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1 Meat Science 78 (2008) 343–358
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Review

Fat deposition, fatty acid composition and meat quality: A review

J.D. Wood *, M. Enser, A.V. Fisher, G.R. Nute, P.R. Sheard, R.I. Richardson,
S.I. Hughes, F.M. Whittington
Division of Farm Animal Science, Department of Clinical Veterinary Science, University of Bristol, Langford, Bristol BS40 5DU, UK

Received 25 May 2007; received in revised form 13 July 2007; accepted 14 July 2007

Abstract

This paper reviews the factors affecting the fatty acid composition of adipose tissue and muscle in pigs, sheep and cattle and shows
that a major factor is the total amount of fat. The effects of fatty acid composition on meat quality are also reviewed. Pigs have high
levels of polyunsaturated fatty acids (PUFA), including the long chain (C20-22) PUFA in adipose tissue and muscle. The full range
of PUFA are also found in sheep adipose tissue and muscle whereas cattle ‘conserve’ long chain PUFA in muscle phospholipid. Linoleic
acid (18:2n 6) is a major ingredient of feeds for all species. Its incorporation into adipose tissue and muscle in relation to the amount in
the diet is greater than for other fatty acids. It is deposited in muscle phospholipid at a high level where it and its long chain products eg
aracidonic acid (20:4n 6) compete well for insertion into phospholipid molecules. Its proportion in pig adipose tissue declines as fat
deposition proceeds and is an index of fatness. The same inverse relationships are not seen in ruminant adipose tissue but in all species
the proportion of 18:2n 6 declines in muscle as fat deposition increases. The main reason is that phospholipid, where 18:2n 6 is
located, declines as a proportion of muscle lipid and the proportion of neutral lipid, with its higher content of saturated and monoun-
saturated fatty acids, increases. Oleic acid (18:1cis 9), formed from stearic acid (18:0) by the enzyme stearoyl Co-A desaturase, is a
major component of neutral lipid and in ruminants the same enzyme forms conjugated linoleic acid (CLA), an important nutrient in
human nutrition. Like 18:2n 6, a-linolenic acid (18:3n 3) is an essential fatty acid and is important to ruminants since it is the major
fatty acid in grass. However it does not compete well for insertion into phospholipid compared with 18:2n 6 and its incorporation into
adipose tissue and muscle is less efficient. Greater biohydrogenation of 18:3n 3 and a long rumen transit time for forage diets also limits
the amount available for tissue uptake compared with 18:2n 6 from concentrate diets. A positive feature of grass feeding is that levels
of the nutritionally important long chain n 3 PUFA are increased ie EPA (20:5n 3) and DHA (22:6n 3). Future research should
focus on increasing n 3 PUFA proportions in lean carcasses and the use of biodiverse pastures and conservation processes which retain
the benefits of fresh leafy grass offer opportunities to achieve this. The varying fatty acid compositions of adipose tissue and muscle have
profound effects on meat quality. Fatty acid composition determines the firmness/oiliness of adipose tissue and the oxidative stability of
muscle, which in turn affects flavour and muscle colour. Vitamin E is an essential nutrient, which stabilises PUFA and has a central role
in meat quality, particularly in ruminants.
 2007 Elsevier Ltd. All rights reserved.

Keywords: Fatty acids; Meat quality; Pigs; Sheep; Cattle; Diets; Genetics; Lipid oxidation; Flavour

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344
2. Fatty acid composition of adipose tissue and muscle in meat animals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344
3. Fatty acid composition of triacylglycerol (neutral lipid) and phospholipid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345

*
Corresponding author. Tel.: +44 117 928 9293; fax: +44 117 928 9582.
E-mail address: jeff[email protected] (J.D. Wood).

0309-1740/$ - see front matter  2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.meatsci.2007.07.019
344 J.D. Wood et al. / Meat Science 78 (2008) 343–358

4. Effects of fat content on fatty acid composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345

4.1. Adipose tissue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
4.2. Muscle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
5. Genetic effects on fatty acid composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 348
6. Diet effects on fatty acid composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
6.1. Pigs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
6.2. Cattle and sheep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350
7. Effects of fat and fatty acids on meat quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353
7.1. Adipose tissue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353
7.2. Muscle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 353
8. Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 356
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357

1. Introduction Enser et al. (1996) was to examine muscle and fat tissues
as normally consumed, so only rough dissection was per-
In many countries, fat is an unpopular constituent of formed, as for someone separating muscle from fat on
meat for consumers, being considered unhealthy. Yet fat the dinner plate. Cores from the centre of longissimus typ-
and fatty acids, whether in adipose tissue or muscle, con- ically contain 1% total lipid in pigs. The data show that
tribute importantly to various aspects of meat quality adipose tissue has a much higher fatty acid content than
and are central to the nutritional value of meat. This review muscle but the fatty acid composition of the two tissues
considers the factors controlling fat deposition and fatty is broadly similar. However, there are important species
acid composition in adipose tissue and muscle of pigs, differences. Pigs have much higher proportions of the
sheep and cattle and the roles of fat in meat quality in these major polyunsaturated fatty acid (PUFA) linoleic acid
different species. (18:2n 6) in both tissues than cattle and sheep. In this
study, the proportions were similar in the two tissues but
2. Fatty acid composition of adipose tissue and muscle in most reports show higher proportions in pig adipose tissue
meat animals than muscle (Teye et al., 2006a; Teye, Wood, Whittington,
Stewart, & Sheard, 2006b). Linoleic acid is derived entirely
The fatty acid composition and total fatty acid content from the diet. It passes through the pig’s stomach
of subcutaneous adipose tissue and longissimus muscle unchanged and is then absorbed into the blood stream in
from loin chops or steaks of pigs, sheep and cattle pur- the small intestine and incorporated from there into tissues.
chased at retail are shown in Table 1 (Enser, Hallett, In ruminants, the fatty acid, which is at high levels in con-
Hewitt, Fursey, & Wood, 1996). The concentrations of centrate feedstuffs (grains and oilseeds), is degraded into
total fatty acids in longissimus are higher than in other monounsaturated and saturated fatty acids in the rumen
studies in which cores from the central part of the muscle, by microbial biohydrogenation and only a small propor-
with no adhering subcutaneous or intermuscular adipose tion, around 10% of dietary 18:2n 6, is available for
tissue, have been examined. The intention of the study of incorporation into tissue lipids. In both sheep and cattle,
the fatty acid is at higher levels in muscle than adipose tis-
Table 1 sue. The second most important PUFA is a-linolenic acid
Fatty acid composition (g/100 g fatty acids) and content (g/100 g total
(18:3n 3), which is present in many concentrate feed
fatty acids in subcutaneous adipose tissue and muscle) of loin steaks/chops
in pigs, sheep and cattle (Enser et al., 1996) ingredients but at lower levels than 18:2n 6. In pigs, the
proportion is higher in adipose tissue than muscle. This is
Adipose tissue Muscle
a major dietary fatty acid for ruminants since it constitutes
Pigs Sheep Cattle Pigs Sheep Cattle
over 50% of total fatty acids in grass and grass products.
14:0 1.6a 4.1b 3.7b 1.3a 3.3c 2.7b Again, a high proportion is biohydrogenated to saturated
16:0 23.9b 21.9a 26.1c 23.2b 22.2a 25.0c
fatty acids in the rumen. In a review, Doreau and Ferlay
16:1cis 2.4a 2.4a 6.2b 2.7b 2.2a 4.5c
18:0 12.8a 22.6b 12.2a 12.2a 18.1c 13.4b (1994) found that a variable proportion of dietary
18:1cis 9 35.8b 28.7a 35.3b 32.8a 32.5a 36.1b 18:3n 3 is biohydrogenated (85–100%) but this is more
18:2n 6 14.3b 1.3a 1.1a 14.2b 2.7a 2.4a than for 18:2n 6 (70–95%), so less is available for incor-
18:3n 3 1.4c 1.0b 0.5a 0.95b 1.37c 0.70a poration into tissues. As with 18:2n 6, proportions in
20:4n 6 0.2 ND ND 2.21b 0.64a 0.63a
ruminants are higher in muscle than adipose tissue.
20:5n 3 ND ND ND 0.31b 0.45c 0.28a
n 6:n 3 7.6 1.4 2.3 7.2 1.3 2.1 Muscle contains significant proportions of long chain
P:S 0.61 0.09 0.05 0.58 0.15 0.11 (C20-22) PUFAs which are formed from 18:2n 6 and
Total 65.3 70.6 70.0 2.2 4.9 3.8 18:3n 3 by the action of D5 and D6 desaturase and elong-
a,b,c
Means with different superscripts are significantly different (P < 0.05). ase enzymes. Important products are arachidonic acid
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 345

Table 2 the trends within each species are typical. In all three spe-
Fatty acid composition (g/100 g fatty acids) of subcutaneous adipose cies, oleic acid (18:1cis 9), the major fatty acid in meat,
tissue and longissimus muscle in pigs fed a control diet (Kouba et al., 2003
60 day feeding period) and Soay sheep fed a control diet (Wachira et al.,
was much more predominant in neutral lipid. This fatty
2002) acid is formed from stearic acid (18:0) by the enzyme stea-
Adipose tissue Muscle
royl Co-A desaturase, a major lipogenic enzyme. On the
other hand, 18:2n 6 was at much higher proportions in
Pigs Sheep Pigs Sheep
phospholipid than neutral lipid. The proportion of
18:2n 6 13.20 2.03 9.69 5.60 18:3n 3 was slightly higher in neutral lipid than phospho-
18:3n 3 1.47 1.09 0.65 1.73
lipid in pigs but in sheep and cattle the proportions were
higher in phospholipid. The differences between sheep
(20:4n 6) and eicosapentaenoic acid (EPA, 20:5n 3) and cattle for 18:2n 6, 18:3n 3 and the long chain
which have various metabolic roles including eicosanoid n 6 and n 3 PUFA in Table 3 are partly due to the dif-
production. Greater incorporation of 18:2n 6 into pig ferent concentrate diets fed. In the work with sheep, dried
muscle fatty acids compared with ruminants produces grass (high in 18:3n 3) formed 75% of the concentrate
higher levels of 20:4n 6 by synthesis and the net result whereas in the cattle study the concentrate contained a high
is a higher ratio of n 6:n 3 PUFA compared with the proportion of full fat soyabean meal, high in 18:2n 6.
ruminants (Table 1). Nutritional advice is for ratios <4.0 Nevertheless, we have often seen higher values for individ-
(Scollan et al., 2006a) so pig muscle is unbalanced relative ual phospholipid PUFAs in sheep compared with cattle.
to that of the ruminants. On the other hand, the ratio of all Long chain n 3 and n 6 PUFA are mainly found in
PUFA to saturated fatty acids (P:S), the target for which is phospholipid but are detected in pig and sheep muscle neu-
0.4 or above, is much higher, beneficially so, in pigs and tral lipid and adipose tissue (Enser, Richardson, Wood,
other monogastrics compared with the ruminants. Gill, & Sheard, 2000; Cooper et al., 2004). We have never
Results in Table 2 (Kouba, Enser, Whittington, Nute, & seen these fatty acids in beef muscle neutral lipid or adipose
Wood, 2003; Wachira et al., 2002) confirm those in other tissue (Scollan et al., 2001; Warren et al., in press-a), con-
studies showing that ruminants have higher proportions firming other studies showing ‘conservation’ of essential
of the two main PUFAs in muscle than adipose tissue fatty acids in cattle muscle where they are less likely to
whereas the opposite is true for pigs. be used for energy production (Crawford, Hare, & White-
house, 1984).
3. Fatty acid composition of triacylglycerol (neutral lipid) The double bonds in unsaturated fatty acids are usually
and phospholipid of the cis type, i.e. the hydrogen atoms attached to the car-
bon atoms in the fatty acid chain point in the same direc-
The major lipid class in adipose tissue (>90%) is triacyl- tion. In ruminants, as a result of biohydrogenation in the
glycerol or neutral lipid. In muscle, a significant proportion rumen, a significant proportion of double bonds are of
is phospholipid, which has a much higher PUFA content in the trans type, i.e. the hydrogen atoms point in different
order to perform its function as a constituent of cellular directions. These fatty acids have particularly low melting
membranes. Values for the fatty acid composition of lon- points as a result of this structure. A major trans fatty acid
gissimus muscle neutral lipid and phospholipid from stud- is 18:1 trans vaccenic which is a biohydrogenation product
ies on pigs, sheep and cattle conducted with collaborators of 18:2n 6. This fatty acid is converted to conjugated lin-
at Bristol are shown in Table 3 (Wood et al., 2004; Demirel oleic acid (CLA, 18:2cis 9, trans 11) in adipose tissue
et al., 2004; Warren et al., in press-a). The three studies are by the action of stearoyl Co-A desaturase, the same enzyme
not directly comparable because different diets were fed but responsible for the production of 18:1cis 9 from 18:0.
Like 18:1cis 9, both 18:1 trans vaccenic and CLA are
Table 3 at higher proportions in neutral lipid than phospholipid
Fatty acid composition (%) of longissimus muscle triacylglycerol (neutral and higher in adipose tissue than muscle. CLA is also pro-
lipid) and phospholipid in pigs (Wood et al., 2004, Durocs), sheep duced in the rumen but synthesis from 18:1 trans vaccenic
(Demirel et al., 2004, megalac diet) and cattle (Warren et al., in press-a,
in tissues is quantitatively the most important contributor
Aberdeen Angus 14 months) fed concentrate-type diets
to tissue levels (Scollan et al., 2006a). CLA has health ben-
Neutral lipid Phospholipid
efits in the human diet although meat from ruminants
Pigs Sheep Cattle Pigs Sheep Cattle makes only a small contribution towards nutritionally sig-
14:0 1.6 3.0 2.7 0.3 0.4 0.2 nificant levels.
16:0 23.8 25.6 27.4 16.6 15.0 14.6
16:1cis 2.6 2.2 3.5 0.8 1.5 0.8
4. Effects of fat content on fatty acid composition
18:0 15.6 13.6 15.5 12.1 10.4 11.0
18:1cis 9 36.2 43.8 35.2 9.4 22.1 15.8
18:2n 6 12.0 1.5 2.3 31.4 12.4 22.0 4.1. Adipose tissue
18:3n 3 1.0 1.2 0.3 0.6 4.6 0.7
20:4n 6 0.2 ND ND 10.5 5.9 10.0 As the fat content of the animal and meat increases
20:5n 3 ND ND ND 1.0 4.1 0.8
between early life and the time of slaughter, the propor-
346 J.D. Wood et al. / Meat Science 78 (2008) 343–358

tions of fatty acids change. In pig subcutaneous adipose tis-

sue, Wood (1984) showed that the C18 fatty acids 18:0 and 45

18:1cis 9 increased in proportion and 18:2n 6 declined 44

during this period. This was ascribed to an increasing role 43

for de novo tissue synthesis of saturated and monounsatu- 42

rated fatty acids and a relatively declining role for the 41

direct incorporation of 18:2n 6 from the diet. A similar 40

result was found by Kouba et al. (2003). Pigs were fed a SR
39
control diet from 40 kg live weight for 20, 60 or 100 days. 38
During this time, the proportion of 18:0 increased from 37
10% to 13%, 18:1cis 9 increased from 38% to 42% and
36
the proportion of 18:2n 6 fell from 19% to 11% of total
35
fatty acids.
34
The inverse relationship between the proportion of
33 X
18:2n 6 in subcutaneous adipose tissue and the amount 15 20
0 5 10
of fat or an index of it such as backfat thickness has been P2 fat thickness ( mm )
observed in several studies in pigs. Wood et al. (1978)
observed correlations of about 0.3 between the proportion x C 1 8-2
27
of 18:2n 6 in the inner layer of subcutaneous adipose tis- 26
25
sue and loin fat thickness in Large White pigs from a line 24
selected for fast growth and low fat thickness and a control 23
22
line. The values for 18:2n 6 were 9.3% in the control line 21
and 10.7% in the selection line. Similarly, in 300 pigs with 20
9
8 mm, 12 mm and 16 mm P2 backfat thickness, average 8
values for 18:2n 6 in subcutaneous adipose tissue fell 7
6
from 14.9% to 12.4% to 10.6% (Wood, Enser, Whittington, 5
Moncrieff, & Kempster, 1989) (Table 4). This study also 14

13
compared entire male and female pigs. Proportions of 12
11
PUFA tend to be high in subcutaneous adipose tissue from 10
entire males and this study showed this was mainly due to 9
8
their thinner backfat. However, even at the same backfat 7
thickness, there was a higher proportion of 18:2n 6 and 0 5 10 15 20

a lower proportion of 18:1cis 9 in subcutaneous adipose P2 fat thickness ( mm )

tissue from entires as the results in Fig. 1 show. At the same Fig. 1. Proportions (g/100 g fatty acids) of 18:1cis 9 and 18:2n 6 in
fat thickness as females, subcutaneous adipose tissue from subcutaneous adipose tissue lipid of pigs plotted against P2 fat thickness
entires contained a higher proportion of water and a lower (x–x entire males; o–o females) (Wood et al., 1989).
proportion of lipid, signifying a less mature tissue. These
results help explain why fat quality tends to be lower in
entire male pigs than castrates and females. ous fat of Jersey cattle of different sexes using biopsies at
The changes in adipose tissue fatty acid composition different ages. Both 16:0 and 18:0 fell in proportion as
with age and fatness are different between pigs and cattle. age increased from 3 to 30 months, whereas 18:1cis 9
Leat (1975) examined fatty acid composition in subcutane- increased, similar to the observation in pigs. In a compar-
ison of extremes, Wood (1984) found proportions of 14.7%
and 2.7% for 18:0 and 41.5% and 56.4% for 18:1cis 9 in a
Table 4 young heifer and an old fat steer respectively. We have
Water and lipid content and fatty acid composition of subcutaneous
recently observed an increase in the proportion of
adipose tissue from the loin (both layers combined) in 300 entire male and
female pigs with different backfat thickness (Wood et al., 1989) 18:1cis 9 in subcutaneous adipose tissue of Aberdeen
Angus crossbred steers fed a concentrate diet between 14
P2 fat thickness (mm)
and 24 months of age (Table 5). Carcass fat greatly
8 12 16
increased during the period as shown by the carcass fat
a
Water 22.4 17.1 14.1 *** score (values are approximately the percentage of subcuta-
Lipida 69.2 77.0 81.6 *** neous fat in the carcass ·10). The proportion of 18:0 fell
18:0b 13.1 13.8 13.9 ***
18:1cis 9b 40.3 41.8 43.1 during the same period (as in the study of Leat) and this
***
18:2n 6b 14.9 12.4 10.6 *** allowed the proportion of 18:2n 6 to remain constant
18:3n 3b 1.1 0.9 0.8 *** (Table 5). This study also showed that the proportion of
a
g/100 g fresh adipose tissue. CLA increased with fatness, as did that of 18:1 trans vac-
b
g/100 g total fatty acids. cenic acid.
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 347

Table 5 proportion of 18:1cis 9 and a decrease in the proportion

Changes in proportions of some adipose tissue fatty acids between 14 and of 18:2n 6.
24 months of age in Aberdeen Angus cross steers fed a concentrate diet
(Whittington, unpublished)
Warren et al. (in press-a) examined the fatty acid con-
tent and composition of neutral lipid and phospholipid in
Age (months)
cattle of three ages, 14, 19 and 24 months. There were
14 24 two breeds, Aberdeen Angus cross and Holstein–Friesian,
Carcass fat score 55 86 and two diets, concentrate and grass silage, fed from 6
18:0 17.7 10.9 months of age. A plot of the concentrations of total neutral
18:1cis 9 28.6 35.2
18:2n 6 1.8 1.9
lipid and phospholipid fatty acids in muscle in relation to
CLA 0.71 1.17 total lipid fatty acids for all 96 steers in the trial is in
Fig. 2. This illustrates the increasing importance of neutral
lipid in total lipid as fattening proceeds and the fairly con-
4.2. Muscle stant level of phospholipid. Results for Aberdeen Angus
steers fed the concentrate diet are in Table 7. The propor-
Early work on meat fatty acid composition concentrated tion of phospholipid in total lipid fell from 30% at 14
on adipose tissue, since that is where the bulk of the body’s months to 12% at 24 months and this was accompanied
fatty acids are located. Recently, there has been more by an increase in the proportion of 18:1cis 9 and a
emphasis on muscle because of its greater significance as decrease in the proportion of 18:2n 6 in total lipid. Data
food and an increasing aversion to visible fat at retail. were statistically analysed within age group in this trial so
Muscle also contains higher concentrations of the long age groups themselves were not directly compared.
chain n 6 and n 3 fatty acids, the importance of which
in human nutrition has been recognised relatively recently.
Separation and identification procedures for low levels of 18000
Diet
unsaturated fatty acids in muscle have also greatly 16000 Concentrate o
improved in recent years. Grass silage
14000
The overall fat content of the animal and muscle have an
Lipid fraction (mg/100g)

important impact on proportionate fatty acid composition 12000

because of the different fatty acid compositions of neutral 10000 Neutral lipid
lipid and phospholipid (Table 3). Phospholipid is an essen-
8000
tial component of cell membranes and its amount remains
fairly constant, or increases little, as the animal increases in 6000
fatness. In young lean animals, genetically lean animals or 4000
animals fed a low energy diet, the lower 18:1cis 9 and
2000 Phospholipid
higher 18:2n 6 content of phospholipid has a major influ-
o
ence on total muscle fatty acid composition. But as body 0
T T T T T T
fat increases, neutral lipid predominates in overall fatty 0 2000 4000 6000 8000 10000 12000 14000 16000 18000
acid composition. Results from the study of Kouba et al. Total lipid (mg/100g)
(2003) of pigs fed a control diet from 40 kg live weight
Fig. 2. Concentrations of neutral lipid and phospholipid (mg/100 g
for 20, 60 or 100 days are shown in Table 6. Phospholipid
muscle) plotted against total lipid (mg/100 g muscle) in longissimus muscle
declined from 46% of total lipid at 20 days to 28% at 100 of steers given a concentrate or grass silage diet and slaughtered at 14, 19
days. This was associated with an increase in the or 24 months of age (Warren et al., in press-a).

Table 6 Table 7
Composition of carcass, longissimus muscle and muscle total fatty acids in Composition of carcass, longissimus muscle and muscle total fatty acids
pigs fed a control diet from 40 kg live weight for 20, 60 or 100 days in Aberdeen Angus cross steers fed a concentrate diet (Warren et al., in
(Kouba et al., 2003) press-a)
Days of feeding Months of age
20 60 100 14 19 24
Subcutaneous fata 11 19 25 *** Carcass fat scorea 55 70 86
Phospholipidb 0.45 0.45 0.36 * Phospholipidb 0.55 0.68 0.67
Neutral lipidb 0.53 0.80 0.92 *** Neutral lipidb 1.28 2.45 4.80
Total lipidb 0.98 1.25 1.28 *** Total lipidb 1.82 3.13 5.48
18:1cis 9c 37.5 42.7 43.6 *** 18:1cis 9c 29.6 31.5 35.9
18:2n 6c 14.7 9.7 8.0 *** 18:2n 6c 8.1 6.7 3.8
a a
g/100 g carcass. Visual score, range 20–145.
b b
g/100 g muscle. g/100 g muscle.
c c
g/100 g total fatty acids. g/100 g total fatty acids.
348 J.D. Wood et al. / Meat Science 78 (2008) 343–358

However, comparison with other results in the trial sug- Table 9

gests that the age effects on neutral lipid, total lipid and Composition of carcass, longissimus muscle and muscle neutral lipid and
phospholipid in Aberdeen Angus cross (AA) and Holstein–Friesian (HF)
fatty acid proportions were statistically significant. The steers fed a concentrate diet and slaughtered at 14 or 24 months of age
trends in Table 7 are similar to those in the pig study in (Warren et al., in press-a)
Table 6. In both studies, there was an increase in the pro- AA HF
portion of 18:1cis 9 and a decrease in the proportion of
14 24 14 24
18:2n 6 in neutral lipid during the periods under investi-
a
gation, evidence of the increasingly important role of stea- Carcass fat score 55 86 24 42
Phospholipidb 0.55 0.67 0.49 0.70
royl Co-A desaturase and the declining importance of 18:1cis 9c 15.8 19.4 12.9 15.8
dietary fat as a source of muscle fatty acids as fat deposi- 18:2n 6c 22.0 23.8 18.6 21.7
tion accelerates, in muscle triacylglycerol as in adipose Neutral lipidb 1.28 4.80 1.10 3.35
tissue. 18:1cis 9c 35.2 38.2 35.3 37.9
18:2n 6c 2.30 1.71 2.85 2.38
a
5. Genetic effects on fatty acid composition Visual score, range 20–145.
b
g/100 g muscle.
c
g/100 g neutral lipid or phospholipid fatty acids.
Breeds or genetic types with a low concentration of total
lipid in muscle, in which phospholipid is a high proportion
of the total, will have higher proportions of PUFA in total
lipid, for the reasons given in Section 4. This was illustrated In the study by Warren et al. (in press-a) of beef fatty
in sheep by Fisher et al. (2000) (Table 8). Welsh Mountain acid composition involving two diets and slaughter at three
and Soay sheep were reared on grass diets and slaughtered ages (Table 7), Aberdeen Angus cross and Holstein–Frie-
at the same body weight. Soays had much leaner carcasses sian breeds were compared. Results for proportions of
and less lipid in muscle. They had lower proportions 18:1cis 9 and 18:2n 6 in phospholipid and neutral lipid
of 18:1cis 9 and higher proportions of all PUFA in in the 14 and 24 month groups fed concentrate are in Table
semimembranosus muscle. 9. Aberdeen Angus had much fatter carcasses than Hol-
Raes, De Smet, and Demeyer (2001) have shown that stein–Friesian, but amounts of neutral lipid and phospho-
the double muscling genotype (mh/mh) within the Belgian lipid in longissimus muscle were not very different and
Blue Breed has low proportions of 18:1 cis 9 and high consequently proportions of 18:1cis 9 and 18:2n 6 in
proportions of 18:2n 6 in muscle lipid compared with both lipid fractions were also quite similar. Bigger differ-
the normal genotype (+/+). This is due to a low concentra- ences would have been expected if muscle lipid concentra-
tion of total lipid in muscle and a higher ratio of phospho- tion had mirrored subcutaneous fat. These results show
lipid to total lipid. Average values for the total lipid that the dairy breed Holstein–Friesian had a higher ratio
content of five muscles in young bulls were 0.9 g/100 g of muscle lipid to carcass fat than the beef breed Aberdeen
and 2.6 g/100 g in mh/mh and +/+ respectively. The pro- Angus. This is consistent with other work showing differ-
portions of 18:1cis 9 were 23.1 and 37.8 and the propor- ences in the partitioning of body fat between dairy and beef
tions of 18:2n 6 were 16.3 and 6.5 in mh/mh and +/+ breeds, with dairy breeds having more ‘‘internal’’ and less
respectively. The mh/mh animals had a P:S ratio of 0.55, ‘‘external’’ (subcutaneous) fat (Truscott, Wood, & MacFie,
above the minimum recommended for the diet as a whole 1983).
and much higher than reported elsewhere for beef (Table 1). The Duroc pig breed is notable in having a high muscle
This study, in common with others, showed only small dif- lipid (marbling fat) content relative to subcutaneous fat
ferences between muscles in fatty acid composition. compared with other breeds. Wood et al. (2004) examined
purebred Berkshire, Duroc, Large White and Tamworth
breeds fed for 12 weeks on a standard concentrate diet.
Table 8 The two traditional breeds (Berkshire and Tamworth) grew
Fat content and fatty acid composition of total lipid in semimembranosus slowly and were lighter and fatter than the two modern
muscle of Welsh Mountain and Soay sheep fed grass diets (Fisher et al., breeds at slaughter (Table 11). The amount of phospho-
2000) lipid in longissimus was similar between the breeds but
Welsh Mountain Soay the amounts of neutral lipid and total lipid were higher
Carcass fat a
12.8 6.8 *** in Berkshire and Duroc than in Large White and Tam-
Total lipidb 2.51 1.67 *** worth. Durocs had the highest ratio of muscle lipid to sub-
18:1cis 9c 33.8 28.0 *** cutaneous fat thickness. The proportion of phospholipid in
18:2n 6c 4.4 12.9 *** total lipid was 18.8, 23.8, 38.9 and 31.7 in Berkshire,
20:4n 6c 1.9 4.0 ***
18:3n 3c 1.6 3.3 Duroc, Large White and Tamworth, respectively. Values
***
20:5n 3c 1.0 1.8 *** for the proportions of 18:1cis 9 and 18:2n 6 in total
a
g/100 g carcass. lipid were as expected based on these figures except for
b
g/100 g muscle. Duroc, the proportion of 18:1cis 9 being lower and the
c
g/100 g total fatty acids. proportion of 18:2n 6 being higher than expected. A
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 349

Table 10 Table 11
Proportions of n 3 PUFA (g/100 g fatty acids) in longissimus phospho- Neutral lipid, phospholipid and total lipid content of longissimus muscle
lipid of Aberdeen Angus cross (AA) and Holstein–Friesian (HF) steers fed and fatty acid composition of total lipid in four pig breeds (Wood et al.,
grass silage and slaughtered at 14 or 24 months of age (Warren et al., in 2004)
press-a)
Berkshire Duroc Large white Tamworth
AA HF b a a
P2 fat thickness (mm) 15 9 8 15b
14 24 14 24 Phospholipidd 0.39 0.42a 0.38a 0.38a
Neutral lipidd 1.67b 1.35b 0.60a 0.82a
18:3n 3 3.70 3.30 3.60 4.00
Total lipidd 2.05b 1.77b 0.97a 1.20a
20:5n 3 3.37 2.77 3.42 4.15
18:1cis 9e 33.5c 29.8b 27.9a 29.4b
22:5n 3 4.60 4.00 4.50 4.40
18:2n 6e 11.8a 16.6b 19.4c 15.9b
22:6n 3 0.85 0.47 1.00 1.02
a–c
DHA/18:3 0.23 0.14 0.28 0.25 Within a row, means with different superscripts are significantly dif-
ferent (P < 0.05).
d
g/100 g muscle.
e
g/100 g total fatty acids.
possible explanation for these results is the slightly higher
proportion of phospholipid in Duroc longissimus muscle
(Table 11) associated with their ‘redder’ muscle fibre type
profile compared with the other breeds reported in a Teye et al. (2006a, 2006b) fed concentrate diets containing
companion paper by Chang et al. (2003). Their fatty acid 2.8% added oil coming from palm kernel oil high in lauric
profile would be expected to be closer to psoas than longiss- acid (12:0), myristic acid (14:0) and 18:0; palm oil high in
imus, with higher 18:2n 6 and lower 18:1cis 9 palmitic (16:0) and palmitoleic (16:1) acids; and soyabean
proportions. oil high in 18:2n 6. The greatest dietary impact in adipose
Analysis of long chain n 3 PUFA proportions in the tissue and muscle was on proportions of 12:0, 14:0 (these
steers fed grass silage in the study of Warren et al. (in had very low proportions) and 18:2n 6, with the C16
press-a) and referred to in Tables 7 and 9, suggested that and C18 saturated and monounsaturated fatty acids hardly
Holstein–Friesians formed more docosahexaenoic acid affected by dietary concentrations. These results are
(DHA, 22:6n 3) than Aberdeen Angus from its precursor explained by the fact that 12:0 and 14:0 are mainly derived
18:3n 3 in phospholipid. Values for these phospholipid from the diet and 18:2n 6 is entirely derived from the
fatty acids and the index DHA/18:3n 3 for the 14 and diet. Conversely, the C16 and C18 saturated and monoun-
24 month silage-fed groups are in Table 10. Most fatty acid saturated fatty acids are mainly the products of synthesis in
proportions were significantly different between the breeds the animal and interconversions between them limit the
at 24 months (P < 0.05) but not at 14 months. The DHA/ impact of dietary additions. The clearest effect was that
18:3n 3 ratio was significantly different between the of soyabean oil on 18:2n 6 in adipose tissue. Proportions
breeds at both ages (P < 0.05). These results suggest that in muscle were lower than in adipose tissue and the dietary
Holstein–Friesians have a greater activity or a greater effect was smaller.
expression of D5 and D6 desaturase enzymes. Evidence that Several studies have examined the effect of 18:3n 3 in
the double muscled (mh/mh) Belgian Blue genotype con- linseed/flaxseed on its concentration in pork. The motiva-
verts a higher proportion of 18:3n 3 to 20:5n 3 and tion for this research is the high n 6:n 3 fatty acid ratio
22:5n 3 but not 22:6n 3 was presented by Raes et al. in pork and the need to reduce this for human nutritional
(2001). reasons. An example is the work of Enser et al. (2000).
Two diets were fed, differing in the ratio of 18:2n 6:
6. Diet effects on fatty acid composition 18:3n 3, to 80 entire male and female pigs between 25 kg
and 95 kg live weight,. The aim was to favour deposition
6.1. Pigs of 18:3n 3 and its long chain products in triacylglycerol
and phospholipid. The n 6 and n 3 PUFA compete
The pig, being a monogastric species, is amenable to for access to desaturase enzymes and for incorporation into
changes in the fatty acid composition of adipose tissue lipids. A control diet contained 1.5 g 18:3n 3 and 16 g
and muscle using diets containing different oils. Spectacular 18:2n 6 kg 1 and a linseed-rich diet contained 4.5 g
results can be achieved using diets with high levels of 18:3n 3 and 10 g 18:2n 6 kg 1. This gave 18:2n 6:
18:2n 6, which is a common fatty acid in grains and oil- 18:3n 3 ratios of 11.0 and 2.0 respectively. The results
seeds. In general, the proportion of this fatty acid in tissues (Table 12) show that the linseed diet increased the deposi-
increases linearly as the dietary intake increases (Wood, tion of 18:3n 3 in adipose tissue and muscle, particularly
1984). In early studies of Ellis and Isbell (1926) the propor- muscle phospholipid. Conversion of this extra 18:3n 3
tion of 18:2n 6 in subcutaneous adipose tissue increased into the C20-22n 3 PUFA 20:5n 3, docosapentaenoic
from 1.9% on a low fat diet to over 30% on diets containing (DPA, 22:5n 3) and 22:6n 3 occurred and these were
a high level of soyabeans. deposited in muscle phospholipid but not in muscle neutral
Other dietary lipid sources containing particular fatty lipid (results not shown). Only 20:5n 3 of the long chain
acids can be used to influence meat fatty acid composition. n 3 PUFA was significantly higher in muscle total lipid
350 J.D. Wood et al. / Meat Science 78 (2008) 343–358

Table 12 this diet was fed. A companion paper by Doran et al.

Fatty acid composition of adipose tissue, longissimus muscle total lipid (2006) showed that low protein diets increased the expres-
and muscle phospholipid (%) in female pigs given a control or linseed-rich
diet (Enser et al., 2000)
sion of stearoyl Co-A desaturase in longissimus muscle and
there was a linear relationship between the expression of
Adipose tissue Muscle Phospholipid
stearoyl Co-A desaturase and the amount of 18:1cis 9
C L C L C L in muscle. These data also show that de novo synthesis of
18:cis 9 33.3a 34.6a 29.6a 32.2a 12.1a 13.6b fatty acids can dominate fatty acid profiles in some
18:2n 6 18.4b 13.8a 17.5b 14.1a 30.2b 27.0a circumstances.
20:4n 6 0.23b 0.16a 4.1b 3.1a 9.7b 8.1a
18:3n 3 1.74a 2.43b 0.84a 1.32b 0.9a 1.84b
20:5n 3 0.05a 0.05a 0.42a 0.73b 1.0a 2.0b 6.2. Cattle and sheep
22:5n 3 0.19a 0.24b 0.95a 1.06a 2.0a 2.5b
22:6n 3 0.08a 0.12b 0.43a 0.47a 1.0a 1.2b Several studies have shown that dietary n 6 and n 3
18:2n 6:18:3n 3 10.5b 5.7a 20.5b 10.5a 33.3b 14.8a PUFA can be incorporated into adipose tissue and muscle
Within tissue/lipid class category and within a row, means with different of ruminants despite the biohydrogenation of dietary fatty
superscripts are significantly different (P < 0.05). acids in the rumen. The study of Warren et al. (in press-a)
of steers of two breeds fed a concentrate or grass silage diet
from 6 months of age to 14, 19 and 24 months contrasts the
of pigs fed the linseed diet. However, there was evidence of incorporation of 18:2n 6 from a grain-based concentrate
extra long chain n 3 PUFA deposition (except for diet with 18:3n 3 from a grass silage diet. Results in
20:5n 3) in adipose tissue, albeit the levels of these fatty Tables 5 and 7 show that 18:2n 6 in steers fed the concen-
acids were very low. trate diet was at higher proportions in muscle than adipose
Nguyen, Nuijens, Everts, Salden, and Beynen (2003) tissue at 14 and 24 months of age. The same was true for
studied the uptake of dietary n 6 and n 3 PUFA into 18:3n 3 from the grass silage diet. For example, the pro-
pig adipose tissue and muscle in their own and in published portions of 18:3n 3 in adipose tissue lipid and total mus-
work and concluded that the efficiency of uptake, defined cle lipid at 14 months were 0.52 and 1.17 g/100 g
as the slope of the line relating tissue level to dietary intake, respectively. At 24 months, the figures were 0.45 and
was greater for 18:2n 6 than 18:3n 3 in both adipose 0.62 g/100 g respectively. These results show that rumi-
tissue and muscle. They found that in the case of nants preferentially incorporate essential fatty acids, with
18:2n 6, the slope was higher for adipose tissue than mus- their important metabolic roles, into muscle rather than
cle but for 18:3n 3, efficiency of uptake into the two tis- storing them in adipose tissue.
sues was similar. The results of Enser et al. (2000) are In the study of Warren et al. (in press-a), the proportion
consistent with these conclusions. of 18:2n 6 in total muscle lipid varied from 1% to 12%.
The study of Kouba et al. (2003) showed that incorpora- As total lipid increased, the proportion fell steeply
tion of 18:3n 3 from a 6% crushed linseed diet into mus- (Fig. 3a) before plateauing at about 6 g/100 g total lipid.
cle neutral lipid and phospholipid reached a maximum in This curvilinear pattern was explained by the high
terms of proportions after 60 days of feeding. However, proportion of 18:2n 6 in phospholipid and a declining
91% and 87% of the effect had occurred in neutral lipid
and phospholipid respectively at 20 days. For 20:5n 3
incorporation into neutral lipid and phospholipid, the 12
Diet
maximum proportions were also reached at 60 days, with Concentrate
85% and 71% of the effect having occurred at 20 days in 10
Grass silage

neutral lipid and phospholipids respectively. These results

confirm the rapid uptake of n 3 PUFA into pork found
% 18:2 in total lipid

8
by Warnants, Van Oeckel, and Boucque (1999) and show
that incorporation of chain elongation products is more 6
rapid in neutral lipid than phospholipid.
In a recent study, Teye et al. (2006a) used low protein 4
diets (18% versus 20% crude protein) with the same energy
content to increase the concentration of total lipid in lon- 2
gissimus muscle. Low protein limits muscle deposition
and the energy which would have been used for muscle syn- 0 I I I I I I I
thesis is diverted to fat synthesis. In the later stages of 0 2000 4000 6000 8000 10000 12000 14000 16000 18000
growth, intramuscular fat is particularly affected. This Total lipid (mg/100g)
strategy increased total lipid from 1.7% to 2.8% and had
Fig. 3a. Proportion of 18:2n 6 in total lipid against total lipid (mg/100 g
a marked effect on the proportion of 18:1cis 9 which muscle) in longissimus muscle of steers given a concentrate or grass silage
increased from 32.1% to 39.0% of total muscle lipid. Pro- diet and slaughtered at 14, 19 or 24 months of age (Warren et al., in
portions of all n 6 and n 3 PUFA were reduced when press-a).
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 351

T
12 5
Diet Diet
Concentrate Concentrate
Grass ilage Grass silage

T
10
4 Phospholipid (silage)

T
8
3

T
6

%18:3
2

T
4

%18:2 in total lipid

Phospholipid (conc)
1
2 Neutral lipid (silage)
o

0 Neutral lipid (conc)

0
0 10 20 30 40 50 0 2000 4000 6000 8000 10000 12000 14000 16000 18000
Phospholipid as % total lipid Total lipid (mg/100g)

Fig. 3b. Proportion of 18:2n 6 in total lipid plotted against phospho- Fig. 4b. Proportions of 18:3n 3 in neutral lipid and phospholipid in
lipid expressed as a % of total lipid in longissimus muscle of steers given a longissimus muscle of steers given a concentrate or grass silage diet and
concentrate or grass silage diet and slaughtered at 14, 19 or 24 months of slaughtered at 14, 19 or 24 months of age (Warren et al., in press-a).
age (Warren et al., in press-a).

proportion of phospholipid in total lipid as total lipid 74 g 18:2n 6 and 73.5 g 18:3n 3 from the concentrate
increased. Proportions of 18:2n 6 in muscle from steers and grass silage diets respectively.
given the two diets were closely related to the percentage Higher levels of 18:2n 6 than 18:3n 3 in tissues are
of phospholipid in total lipid (Fig. 3b). The proportions not only due to a higher affinity for incorporation into
of 18:2n 6 and 18:3n 3 in phospholipid and neutral phospholipid molecules as illustrated in Figs. 4a and 4b
lipid plotted against total muscle lipid for all steers fed but also reduced biohydrogenation in the rumen. This
the concentrate and grass silage diets are shown in Figs. occurs when the form of the diet is similar (Doreau & Fer-
4a and 4b. These graphs emphasise the much greater incor- lay, 1994) and particularly in typical 18:2n 6-rich concen-
poration of 18:2n 6 than 18:3n 3 into muscle lipids, trate diets. These have a small particle size and a shorter
especially phospholipid, and the declining proportions of rumen transit time than fibrous forage diets, limiting the
these PUFA as muscle lipid increased. The content of opportunities for microbial biohydrogenation. Our studies
phospholipid fatty acids remained fairly constant but neu- with sheep have mainly used concentrate-based diets and
tral lipid, with its high proportions of saturated and mono- this may be one reason why concentrations and propor-
unsaturated fatty acids, increased markedly as total lipid tions of PUFA in muscle are higher than in the studies
increased (Fig. 2). These differences in tissue levels of the on beef which have mainly used diets containing 60% for-
two essential fatty acids are the more surprising consider- age and 40% concentrate (Demirel et al., 2004; Scollan
ing that intakes of the fatty acids were similar. For example et al., 2001).
in the 14 month groups, the approximate daily intakes were Incorporation of 18:2n 6 from the concentrate diet
and 18:3n 3 from the grass silage diet into muscle in
T

30 the study of Warren et al. (in press-a) led to synthesis of

Diet the long chain n 6 and n 3 PUFA in phospholipid.
Concentrate
o

Grass silage
T

25 Phospholipid
Results for the 14 month Aberdeen Angus steers are
in Table 13. These data are concentrations in muscle
T

20

Table 13
T

15 Concentrations (mg/100 g muscle) of n 6 and n 3 PUFA in longiss-

%18:2

imus phospholipid of Aberdeen Angus steers fed concentrate or grass

10
silage and slaughtered at 14 months of age (Warren et al., in press-a)
Phospholipid
Concentrate Grass silage
5 18:2n 6 119.0 46.6 ***
o

20:3n 6 14.3 6.1 ***

Neutral lipid
20:4n 6 54.2 33.2
o

***
T

0
22:4n 6 6.3 2.2 ***
0 2000 4000 6000 8000 10000 12000 14000 16000 18000 18:3n 3 4.0 20.6 ***
Total lipid (mg/100g) 20:4n 3 0.8 4.4 ***
20:5n 3 4.8 18.6 ***
Fig. 4a. Proportions of 18:2n 6 in neutral lipid and phospholipid in
22:5n 3 11.2 25.7 ***
longissimus muscle of steers given a concentrate or grass silage diet and
22:6n 3 1.2 4.7 ***
slaughtered at 14, 19 or 24 months of age (Warren et al., in press-a).
352 J.D. Wood et al. / Meat Science 78 (2008) 343–358

(mg/100 g) rather than proportions in phospholipid. The 160

concentrate diet produced relatively high levels of 140
18:2n 6 and all its long chain products and the grass
120
silage diet produced high levels of 18:3n 3 and its long
chain products, including 22:6n 3. Feeding linseed in pre- 100

mg/100g
vious research had not led to synthesis of 22:6n 3 (DHA) 80
and a block on DHA synthesis or a failure to compete for
60
incorporation has been noted in other studies (Scollan
et al., 2001). It seems that grass feeding has a special ability 40

to raise DHA levels. 20

In the 19 and 24 month age groups in the study of War- 0
ren et al. (in press-a), there was evidence of extra incorpo- 14 19 24
ration of 18:2n 6 and synthesis of 20:4n 6 in months
phospholipid beyond 14 months (Fig. 5a). However, the 18:2 conc 18:3 silage

amounts of 18:3n 3 and its products remained constant, Fig. 5b. Amounts (mg/100 g muscle) of 18:2n 6 and 18:3n 3 in
despite continued consumption of the grass silage diet. longissimus neutral lipid of steers given concentrate or grass silage
These results suggest that the capacity for incorporation respectively (Warren et al., in press-a).
of PUFA into phospholipid is limited and that 18:2n 6
competes for incorporation much more effectively than
doubled the concentration of 18:2n 6 in phospholipid
18:3n 3. Evidence suggests that the n 3 PUFA are the
and substantially increased the concentration of this fatty
preferred substrates for the D5 and D6 desaturase enzymes
acid in neutral lipid compared with 18:3n 3. Because of
(Williams & Burdge, 2006) so limited access to the enzyme
the high incorporation of 18:2n 6, the P:S ratio in muscle
systems cannot explain low values for long chain n 3
was increased from 0.1 in controls to 0.4 in animals given a
PUFA.
high level of the supplement. These results again demon-
In contrast to these results for phospholipid, extra incor-
strate the higher efficiency of incorporation of 18:2n 6
poration of 18:2n 6 and 18:3n 3 into muscle triacyl-
into muscle compared with 18:3n 3. In Australian
glycerol (neutral lipid) occurred beyond 14 months of age
research, Cook, Scott, Faichney, and Davies (1972)
(Fig. 5b). The level and rate of incorporation was greater
observed that the proportion of 18:2n 6 increased to
for 18:2n 6 than for 18:3n 3.
35 g/100 g fatty acids in perirenal fat of steers given a pro-
Levels of n 3 PUFA in ruminant tissues can be
tected sunflower supplement for 8 weeks. A value of 20 g/
increased by feeding dietary lipid which is ‘protected’ from
100 g was achieved after 2 weeks.
biohydrogenation in the rumen using formaldehyde treat-
In the work of Warren et al. (in press-a), a group of
ment of linseed. In a study by Scollan, Enser, Gulati, Rich-
steers was fed fresh grazed grass rather than grass silage
ardson, and Wood (2003), in which a protected lipid
between 14 and 19 months of age. The results in subcuta-
supplement comprised of soyabean, linseed and sunflower
neous adipose tissue (Table 14) showed that the proportion
seeds was fed, the concentration of 18:3n 3 in muscle
of 18:3n 3 was slightly higher in the steers fed grazed
phospholipid increased from 12.7 to 16.0 mg/100 g, a small
grass. This group also had higher proportions of 18:1 trans
increase and no chain elongation and desaturation to long
vaccenic acid and CLA than those fed grass silage, showing
chain n 3 PUFA occurred. However, the supplement
that the process of rumen biohydrogenation is different
between fresh and conserved grass. A similar result was
found by French et al. (2000). The CLA values were similar
160 in the groups fed grazed grass and concentrate in our work
140 (Table 14).
120
100
mg/100g

80
60 Table 14
40 Fatty acid composition (g/100 g fatty acids) of subcutaneous adipose
20 tissue of steers given concentrate, grass silage or fresh grazed grass
between 14 and 19 months of age Whittington et al. (in preparation)
0
14 19 24 Concentrate Grass silage Grazed grass
months
18:0 14.00 10.09 15.20 ***
18:2 conc 20:4 conc 18:3 silage 20:5 silage 18:1cis 9 31.17 32.83 32.06 NS
18:1trans 4.07 1.48 4.32 ***
Fig. 5a. Amounts (mg/100 g muscle) of 18:2n 6 and 20:4n 6 in
CLA 0.95 0.43 0.93 ***
longissimus phospholipid of steers given concentrate and 18:3n 3 and
18:2n 6 2.39 0.90 0.97 ***
20:5n 3 in longissimus phospholipid of steers given grass silage (Warren
18:3n 3 0.22 0.59 0.68 ***
et al., in press-a).
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 353

Changes in grassland management, such as harvesting at two genetic selection lines, 18:0 and 18:2n 6 proportions
different times of the grass growing season or allowing the were correlated with the melting point of extracted lipid
grass to wilt before harvesting and conservation have an from subcutaneous fat and in this case the proportion of
effect on fatty acid proportions in grasses and also in the 18:0 provided the best prediction of melting point (Wood
meat of cattle and sheep (Wood et al., 2007). Different et al., 1978).
grasses and pasture plants also produce different concen- Changing the fatty acid composition of subcutaneous
trations of PUFA in meat due to higher levels of certain adipose tissue using different dietary oils also changes lipid
PUFA or because of differences in the way the feed is pro- melting point and fat firmness. For example, palm kernel
cessed in the rumen. Scollan et al. (2006b) showed that the oil produced firmer fat than soyabean oil in the study of
proportions of both 18:2n 6 and 18:3n 3 in muscle Teye et al. (2006b). When all the data were pooled, the pro-
were significantly increased when steers were fed silage portions of 12:0 and 14:0 (high in pigs given palm kernel
comprised of red clover rather than perennial ryegrass. oil) were strongly correlated with fat quality parameters,
Other research (Lee et al., 2004) suggests that the pattern as also were 18:0 and 18:2n 6.
of rumen fermentation and biohydrogenation for red clo- In lamb subcutaneous fat sampled throughout the year
ver is different from that of perennial ryegrass due to the in four abattoirs, Enser and Wood (1993) found that melt-
inhibition of lipolysis in clover by the plant enzyme poly- ing point varied with the time of year, being lowest in the
phenol oxidase. Spring and Summer and highest later in the year. Melting
(slip) point ranged from 30 C to 49 C and 18:0, which
7. Effects of fat and fatty acids on meat quality ranged from 7.0% to 32.9% of fatty acids (mean 18.8%)
was the fatty acid most highly correlated with melting point
7.1. Adipose tissue (r 0.89). Linoleic acid was 1.3% of fatty acids overall and its
correlation with melting point was 0.3.
Work with pigs and ruminants has shown that the fatty Lamb subcutaneous fat is unusual in having significant
acid composition of adipose tissue affects its firmness, concentrations of methyl branched fatty acids of medium
because the different fatty acids have different melting to long chain length (C8-17) with low melting points. Their
points. The composite fatty acids of meat melt between concentration reached 4% of the total in the study of Enser
about 25 C and 50 C, with saturated fatty acids melting and Wood (1993). These fatty acids are responsible for the
at higher and polyunsaturated fatty acids at lower temper- soft, oily fat found in sheep that have consumed high grain
atures e.g. 18:0 melts at 69 C and 18:2n 6 at 5 C diets which produce high levels of propionic acid in the
(Wood, 1984). rumen.
In pigs, the differences in fatty acid composition of sub-
cutaneous fat between carcasses of different P2 fat thick- 7.2. Muscle
ness have an important effect on fat quality defined in
terms of firmness and the degree of cohesiveness between The total lipid content of muscle (intramuscular fat,
lean and fat tissues (fat separation). In a study of carcasses often termed marbling fat, although this is strictly the
with 8 mm, 12 mm and 16 mm P2 fat thickness, the propor- flecks of adipose tissue composed mainly of neutral lipid)
tion of 18:0 increased and that of 18:2n 6 decreased as fat has a role in the tenderness and juiciness of cooked meat
thickness increased (Table 4). The backfat of pigs with although the strength of the correlation varies considerably
16 mm P2 was firmer and there was less separation between between studies, with some showing an important role for
fat and underlying muscle than in backfat from the 8 mm marbling fat and others showing only a weak relationship
P2 group (Wood, Jones, Francombe, & Whelehan, 1986). with sensory traits. The role of marbling fat is of particular
Firmness, measured both objectively and subjectively in interest in pigs because genetic selection for lean pigs has
the shoulder and loin regions, was correlated with fatty reduced the level of marbling fat to below 1% of muscle
acid proportions, the highest correlations being with 18:0 weight in modern pigs (e.g. Large Whites in Table 11) com-
(positive) and 18:2n 6 (negative). The proportion of pared with 2–4% in US studies in the 1960s (Wood, 1990).
18:2n 6 provided the best prediction of fat firmness In the study of four breeds of Wood et al., 2004, the highest
(Table 15). In an earlier study of Large White pigs from correlation between marbling fat concentration and sen-
sory traits across all four breeds was 0.17, for tenderness.
The correlation in Berkshires was 0.34.
The study of Wood et al. (1986) showed that total lipid
Table 15 (marbling fat) in longissimus muscle was 0.55, 0.66 and
Correlations between fatty acid proportions and firmness of subcutaneous 0.96 g/100 g in pig carcasses having 8 mm, 12 mm and
fat in the shoulder region of pig carcasses having a wide range of P2 fat
16 mm P2 fat thickness respectively. These were very light
thickness (5–20 mm) (Wood et al., 1989)
carcasses, weighing 58 kg on average. Correlations across
18:0 18:2n 6
all pigs between marbling fat and sensory traits were 0.13
Firmness objective 0.35 0.75 for tenderness and 0.31 for juiciness. Juiciness was signifi-
Firmness subjective 0.40 0.78
cantly lower in the 8 mm than the 16 mm P2 fat thickness
354 J.D. Wood et al. / Meat Science 78 (2008) 343–358

group. In a recent comparison of lamb chops produced in with brine to a target level of 10%. The loin was immersed
organic and conventional production systems, Angold in brine for 3 days after which the bacon was sliced, blast
et al. (in press) showed that the correlations between the frozen, stored at 18 C for 8 weeks, thawed, packed in
total fatty acid content of longissimus (marbling fat) and overwrapped trays and kept under retail display conditions
eating quality scores given by the taste panel were 0.36 for up to 9 days. Injection of salt followed by freezing and
and 0.06 for juiciness and toughness respectively. It seems thawing were probably the most important contributors to
from these results that juiciness is the trait most affected by lipid oxidation. Even after this treatment, TBARS values
increasing levels of marbling fat, associated with greater were below 0.5 at 5 days of retail display, increasing to
retention of water in meat during cooking. The location about 1.5 after 9 days. The bacon with a high level of
of marbling fat in the perimysial connective tissue between 18:3n 3 (1.3%) had a similar TBARS value to the con-
muscle fibre bundles may also be important in ‘opening up’ trols (18:3n 3 0.85%). The bacon was assessed by the
the structure of muscle, allowing it to be more easily bro- trained taste panel after the freezing stage and no differ-
ken down in the mouth (Wood, 1990). There are therefore ences n flavour characteristics between feeding treatments
good reasons to expect a positive role for marbling fat in were detected.
meat quality. It is possible that in the study of Sheard et al. (2000),
The use of low protein diets to increase marbling fat in 18:3n 3 proportions in muscle of treated pigs were below
pigs has sometimes produced a higher score for tenderness those likely to produce oxidation products having adverse
and juiciness in cooked pork. In the study of Teye et al. effects on pork flavour. In the work of Kouba et al. (2003),
(2006a), total lipid was increased to 2.8% of longissimus in which the proportion of 18:3n 3 was increased to 3.0%
using an 18% protein diet compared with 1.7% in a stan- of muscle fatty acids compared with 0.65% in controls, an
dard diet containing 20% protein. The scores for tenderness increase in TBARS after 7 days of retail display was
and juiciness (1–8 range) were increased from 4.2 and 3.9 in observed, although only to 0.15 mg malondialdehyde/kg
the 20% protein diet to 4.8 and 4.4 in the 18% protein diet compared with 0.10 in controls. A slightly higher level of
(both P < 0.01). abnormal odour was detected by the taste panel in subcu-
Despite contradictory results between studies for the taneous adipose tissue compared with the controls, and the
role of marbling fat in the tenderness and juiciness of fresh ‘flavour liking’ score of longissimus muscle was significantly
pork, beef and lamb, incorporation of adipose tissue at dif- reduced. It is at levels of around 3% 18:3n 3 in muscle
ferent levels into burgers or patties has been linked posi- fatty acids that other workers have detected off flavours
tively to tenderness and juiciness in several studies (e.g. as a result of feeding linseed (Shackelford, Miller, Haydon,
Kregel, Prusa, & Hughes, 1986). In these cases, positive & Reagan, 1990).
effects on tenderness and juiciness are observed at between Feeding fish oils to pigs increases levels of long chain
10% and 20% lipid rather than at the lower levels seen for n 3 PUFA in adipose tissue and particularly in muscle
marbling fat. and fishy odours and flavours are detected when critical tis-
The fatty acid composition of muscle affects its oxidative sue levels are exceeded. In the work of Overland, Taugbol,
stability during processing and retail display, the polyun- Haug, and Sundstol (1996), feeding a 1% fish oil diet
saturated fatty acids in phospholipid being liable to oxida- between 10 and 100 kg live weight increased the propor-
tive breakdown at this stage. A standard test for lipid tions of 20:5n 3 and 22:6n 3 from 0.6 and 0.9% of mus-
oxidative stability in foods is the thiobarbituric acid react- cle fatty acids in controls to 1.5% and 1.8% respectively.
ing substances (TBARS) test of Tarladgis, Watts, Youna- This caused significantly higher ‘off odour’ and ‘off flavour’
than, and Dugan (1960) which measures the oxidation scores in cooked subcutaneous adipose tissue sampled both
product malondialdehyde. Values above about 0.5 are con- fresh and after 6 months frozen storage. A 3% fish oil diet
sidered critical since they indicate a level of lipid oxidation increased these scores even more.
products which produce a rancid odour and taste which Several studies with pigs have shown that high levels of
can be detected by consumers. vitamin E in the diet are incorporated into tissues where
Values of TBARS in our studies with pork have usually they are effective in reducing lipid oxidation in stored and
been well below 0.5, even when PUFA proportions have displayed pork (Buckley, Morrissey, & Gray, 1995). How-
been increased to high levels using soyabean oil or linseed ever, a level of 150 mg/kg diet did not prevent the deterio-
(Riley, Enser, Nute, & Wood, 2000; Kouba et al., 2003; ration in flavour when linseed feeding raised the level of
Sheard et al., 2000). In the studies of Riley et al. (2000) 18:3n 3 to 3% of muscle fatty acids in the study of Kouba
and Sheard et al. (2000), minced and comminuted products et al. (2003) and extra vitamin E did not increase storage
were produced which develop higher levels of oxidation stability in pigs given diets containing 0.5% fish oil in a
because the fatty acids are exposed to pro-oxidants such study by Hertzman, Goransson, and Ruderus (1988). In
as iron released from muscle cells. Even here, TBARS val- some studies, ‘supranutritional’ vitamin E has reduced drip
ues remained below 0.5 except in the case of bacon in the loss and improved colour stability in pork, probably by
work of Sheard et al. (2000). In this study, several factors preventing oxidation of muscle pigments by lipid oxidation
contributed to high levels of oxidation. The loin joint was products but in others, limited effects on drip loss and col-
conditioned at 1 C for 10 days, after which it was injected our stability have been seen (Jensen et al., 1997).
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 355

In ruminants, we have frequently seen higher values of ments were made at 4 and 7 days of display. The results
TBARS than in our studies on pork. In a recent study, (Table 17) show that the concentrate diet increased lipid
Nute et al. (in press) examined oxidative stability and eat- oxidation in the steaks, values increasing to over 2.0 after
ing quality in lambs which had been fed different levels of 7 days of display in both breeds. On the other hand,
n 3 PUFA from linseed oil, fish oil, a protected lipid sup- TBARS values remained at low levels in the grass silage-
plement (PLS) made from linseed, sunflower seed and fed groups, albeit these were higher than we normally see
soyabean meal, marine algae (contains long chain n 3 in pork. These TBARS values were apparently inversely
PUFA) and combinations of these different oil sources. related to the vitamin E concentration in muscle and
Leg steaks were conditioned for 6 days at 0 C then packed plasma. Steers fed grass silage had higher values for muscle
in modified atmosphere packs (O2:CO275:25) and dis- and plasma vitamin E than those fed concentrate. Muscle
played under retail conditions. Lipid oxidation was mea- values for the grass silage groups were around the 3.3–
sured on the semimembranosus muscle at 7 days of 3.5 mg/kg level found by Arnold, Scheller, Arp, Williams,
display. The lowest TBARS value was in muscles from and Schaefer (1993) to give optimum lipid stability in lon-
the group given linseed oil (0.6 mg/kg) and all other groups gissimus while values in the concentrate group were consid-
had values above 2.0 mg/kg, the highest value being 6.2 in erably lower. The high vitamin E values in the steers fed
the group given a combination of algae and fish oil. All the grass silage are partly the result of lower PUFA levels in
groups, except those given linseed, had low taste panel muscle but mainly due to greater uptake of vitamin E from
scores for lamb flavour and high scores for abnormal lamb the diet. When the data from all animals slaughtered at 14,
flavour. These scores were correlated with fatty acid pro- 19 and 24 months was combined, TBARS were higher in
portions in phospholipid, negative correlations being found the concentrate-fed animals than in those fed grass silage
between long chain n 3 PUFA and lamb flavour (Table
16). The fatty acid composition of semimembranosus
phospholipid was greatly affected by diet in this study. Table 17
For example, 18:2n 6 varied from 11.5 (fish oil) to Lipid oxidation (TBARS) and vitamin E content of longissimus muscle
and vitamin E content of plasma in Aberdeen Angus cross (AA) or
33.7% (PLS), 18:3n 3 from 1.4 (combination of fish oil
Hostein–Friesian (HF) beef steers given concentrate or grass silage diets
and marine algae) to 6.9% (linseed) and 22:6n 3 from between 6 and 14 months of age (Warren et al., in press-b)
0.6% (PLS) to 5.35% (combination of PLS and marine
AA HF Significance
algae). Significant proportions of long chain n 3 PUFA
Concentrate Grass Concentrate Grass Breed Diet
were also detected in adipose tissue. A companion paper
silage silage
by Elmore et al. (2005) showed that very high levels of lipid
TBARSa
oxidation products were produced during the cooking of
day 4 1.2 0.3 1.4 0.3 NS ***
these samples to affect the flavour scores. Muscle samples day 7 2.1 0.4 2.7 0.4 NS ***
from the fish oil/marine algae treatments had the highest
Vitamin E
lipid oxidation and the lowest concentration of vitamin muscleb 1.3 3.6 1.3 3.2 NS ***
E. Other work has shown low vitamin E levels in pig tissues plasmac 2.2 5.8 2.1 4.5 * ***
containing high PUFA proportions, suggesting utilization a
mg malondialdehyde/kg.
of the vitamin to control oxidation (Kouba et al., 2003). b
mg/kg.
In cattle, consumption of concentrate produced higher c
mg/l.
TBARS values in steaks than consumption of grass silage
in a study of Warren et al. (in press-b). This is a companion
12
study to that of Warren et al. (in press-a). Aberdeen Angus
cross and Holstein–Friesian steers were fed the diets from 6
10
to 14 months of age. After slaughter, loin joints were con- Concentrate
ditioned at 1 C for 10 days. Steaks were then placed in o Grass silage
TBARS (mg /kg meat)

8
modified atmosphere packs (O2:CO275:25) and displayed
under retail-type conditions. Lipid oxidation measure- 6

Table 16 4
Correlations between lamb flavour scores and proportions of phospho-
lipid fatty acids in lambs given different dietary oil sources (Nute et al., in 2
press) o
Lamb flavour Abnormal flavour 0 T T T 1

100 200 300 400 500

18:2n 6 0.25 0.11
18:3n 3 0.51 0.49 Muscle total PUFA (mg/100g)
20:4n 6 0.11 0.20
Fig. 6. Relationships between lipid oxidation (TBARS) and longissimus
20:5n 3 0.13 0.24
total PUFA (mg/100 g) in steers given concentrate or grass silage diets
22:6n 3 0.28 0.32
(Warren et al., in press-b).
356 J.D. Wood et al. / Meat Science 78 (2008) 343–358

at all levels of PUFA in muscle (Fig. 6). TBARS increased Table 18

with PUFA level only in those fed concentrate. For the Correlations (Spearmans rho) between lipid oxidation (TBARS) and beef
flavour terms in samples in which oxidation was promoted (Campo et al.,
steers fed grass silage it appears that vitamin E was suffi- 2006)
ciently high to protect the PUFA from oxidation, at least
Beef flavour 0.80***
for the 7 days during which the beef was displayed. Abnormal flavour +0.82***
In the study of Warren et al. (in press-b), the concentrate Rancid +0.84***
diet contained a standard level of vitamin E (25 mg/kg). In Greasy +0.70***
work with sheep, Demirel et al. (2004) fed concentrate diets Bloody 0.60***
containing different oil sources (5%) and either 100 or Metallic 0.36***
Livery 0.60***
500 mg/kg vitamin E. Despite these ‘supranutritional’ lev-
***
els, concentrations of the vitamin in muscle were 0.27 P < 0.001, n = 216.
and 0.52 mg/kg for the 100 and 500 mg/kg diets respec-
tively. Values around 5 mg/kg would be expected. The rea-
son for these very low levels is unclear but these results, mal flavour, rancid and greasy increased (Table 18). This
together with those of Warren et al. (in press-b) point to study identified a TBARS value of 2.3 as the point where
excessive utilisation of antioxidants in concentrate-type rancid and other abnormal flavours overpower beef flavour
diets. to produce an unacceptable flavour profile in beef. Below
A change in muscle colour seen during retail display in this, rancidity was detected but beef flavour remained high.
the study of Warren et al. (in press-b) suggested a link These results suggest that the upper limit for TBARS of 0.5
between lipid oxidation, vitamin E concentration and col- suggested by Tarladgis et al. (1960) based on pork may not
our. The intensity of the red colour, termed saturation or be appropriate for beef (or lamb) where the natural level of
chroma, declined gradually as the display period pro- lipid oxidation is higher.
gressed but the decline was more rapid in the muscles from
the groups fed concentrate than the groups fed grass silage. 8. Conclusions
A value of 18 for colour saturation, which has been used as
an index of the end of shelf life, was reached 2–3 days This review has shown that the fatty acid composition of
sooner in the concentrate groups. adipose tissue and muscle in pigs, sheep and cattle depends
Elmore et al. (2004) examined the flavour volatile com- on the amount of fat in the carcass and in muscle. Effects of
pounds produced when beef samples from the study of diet and breed have to be judged against the amount of fat.
Warren et al. (in press-b) were grilled. Products of Also, there are important differences between the species
18:2n 6 oxidation such as pentanal and hexanal were which are only partly explained by differences in the diges-
detected in steaks produced using concentrate and the alco- tive process. These include: ruminants conserve PUFA in
hols 1-penten-3-ol and cis 2-penten-1-ol, products of muscle whereas in pigs, concentrations are higher in adi-
18:3n 3 oxidation, were detected in steaks from the pose tissue; long chain (C20-22) PUFA are found in adi-
grass-silage-fed group. A compound formed from chloro- pose tissue and muscle neutral lipid in pigs and sheep but
phyll, 2-phytene, was higher in samples from the grass- not in cattle; and the ratio of 18:0/18:2n 6 in adipose tis-
fed groups. Despite these differences in lipid oxidation, sue increases as fattening proceeds in pigs but declines in
the trained taste panel at Bristol could detect few clear dif- ruminants.
ferences in sensory (eating) quality between the concentrate In muscle, the high proportion of 18:2n 6 in phospho-
and grass silage groups. On balance, the panel preferred lipid compared with neutral lipid in all species means that
loin steaks from steers fed grass silage. This result is consis- muscle from lean animals has high proportions of this
tent with papers which have shown that when grain- and major PUFA. As the animal is fattened for meat, the
grass-fed cattle grow at similar rates, sensory scores are decline in PUFA proportions is more dramatic in the rumi-
similar (Bidner et al., 1986). Comparisons of well finished nants because PUFA levels in neutral lipid are so low.
grain-fed with poorly finished grass-fed steers have often Since desirable sensory characteristics tend to increase with
found results in favour of those fed grain (Medeiros, Field, fatness, there is potentially an inverse relationship between
Menkhaus, & Russell, 1987). nutritional value and eating quality in ruminants.
Campo et al. (2006) examined eating quality in 73 beef Of the 2 major PUFA, 18:2n 6 is more rapidly taken
loins produced using different feeding treatments and con- up into meat tissues than 18:3n 3 and reaches much
taining different concentrations of n 6 and n 3 PUFA. higher levels. Synthesis of long chain PUFA from these
Lipid oxidation was promoted by conditioning for 10 days, precursors occurs in phospholipid but the available sites
freezing, thawing and storing steaks in modified atmo- for incorporation are soon filled and long term feeding of
sphere packs (O2:CO2 75:25) before sensory analysis. This linseed to pigs or grass to cattle does not increase levels
series of procedures promoted high levels of lipid oxida- and so proportions decline.
tion, TBARS values up to 12.0 being recorded. As the Oxidation of fatty acids proceeds at a naturally higher
TBARS value increased, the scores for beef flavour, level in ruminants than pigs after slaughter despite the
bloody, metallic and livery declined, and scores for abnor- lower PUFA proportions. Vitamin E is a vital meat quality
 J.D. Wood et al. / Meat Science 78 (2008) 343–358 357

enhancing nutrient, particularly in ruminants where high acids, breed and dietary vitamin E on the fatty acids of lamb muscle,
concentrations resulting from grass feeding prevent fatty liver and adipose tissue. British Journal of Nutrition, 91, 551–565.
Doran, O., Moule, S. K., Teye, G. A., Whittington, F. M., Hallett, K. G.,
acid oxidation and extend colour shelf life. Low concentra- & Wood, J. D. (2006). A reduced protein diet induces stearoyl-CoA
tions, often seen after the feeding of concentrate diets, lead desaturase protein expression in pig muscle but not in subcutaneous
to a shorter colour shelf life and unfavourable beef flavour adipose tissue: relationship with intramuscular lipid formations.
notes. British Journal of Nutrition, 95, 609–617.
Doreau, M., & Ferlay, A. (1994). Digestion and utilisation of fatty acids
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We are grateful to many collaborators in the research Elmore, J. S., Cooper, S. L., Enser, M., Mottram, D. S., Sinclair, L. S.,
presented including Professor Nigel Scollan and Helen Wilkinson, R. G., et al. (2005). Dietary manipulation of fatty acid
Warren of the Institute of Grassland and Environmental composition in lamb meat and its effect on the volatile aroma
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per Adams University College and Professor Don Mottram Elmore, J. S., Warren, H. E., Mottram, D. S., Scollan, N. D., Enser, M.,
Richardson, R. I., et al. (2004). A comparison of the aroma of
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