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Effect of nutritional factors on fatty acid composition of lamb
fat deposits
*
P. Bas , P. Morand-Fehr
Laboratoire de Nutrition et Alimentation, (INRA-INAPG), 16, Rue Claude Bernard, 75231 Paris Cedex, France
Abstract
A literature review was made of the influence of nutritional factors on the fatty acid composition of fat deposits and muscles of lambs. A bibliographic data base, containing 979 observations from 108 papers, was made up in such way that one observation corresponded to one group of lambs in one experiment. Each nutritional factor was studied only when number of observations was sufficient. Consequently, special emphasis was given to the effect of dietary ingredients supplying energy or nitrogen on the fatty acid composition of subcutaneous (SC) and perirenal adipose (PR) tissues and in muscles of lambs. Regardless of sex, breed and breeding conditions, all the tissues examined showed a similar pattern of modification in fatty acid composition with diets. In milk fed lambs the pattern of fatty acids largely reflected the dietary fat pattern. The fatty acid of stored fats before weaning had an influence on fatty acid composition of fat deposits and muscle, several months after weaning. When the diets were rich in beet pulp or in fish meal the proportion of C18:1was significantly increased and the proportions of C18:0, C18:2 and C18:3decreased in fat tissues. Inclusion of maize in diet resulted in an increase in linoleic acid content in fat deposits. Inclusion of cotton meal increased linoleic and stearic acids in fat deposits. Grass-based diets increased C18:0and C18:3in lamb tissues. Melting point of both SC and PR were strongly associated with differences in C18:0 percentages. This approach underlined difficulties in understanding the diet effects on fatty acid composition of fat deposits and muscles without taking feeding and other management aspects into account. This study supported the extent to the modification of the fatty acid composition of lamb carcasses by choice of dietary ingredients despite the ruminal hydrogenation of dietary fat.
´ ´
Resume
ˆ ´
Effets des facteurs nutritionnels sur la composition en acides gras des depots adipeux d’agneaux. L’influence des facteurs
´ ´ ´ ´ ` ´ ´ ´
nutritionnels sur la composition en acides gras a ete etudiee a partir des donnees de la litterature. Une base de donnees
´ ´ ´ ´ `
bibliographiques de 979 observations provenant de 108 articles a ete elaboree de telle sorte qu’une observation corresponde a
´ ´ ´
un groupe d’animaux dans une experience. Chaque facteur nutritionnel etudie devait disposer d’un nombre suffisant
´ ´ ` ` ´
d’observations. De ce fait, un accent particulier a ete mis sur les effets des matieres premieres, incorporees comme source ˆ
´ ´ ´ ´ ´
d’energie ou d’azote, sur la composition en acides gras des depots adipeux sous-cutanes (SC), perirenaux (PR), et des
´ ´ ´ ´ ´ ´
muscles des agneaux. Independamment du sexe, de la race et des conditions d’elevage, les tissus etudies ont presente, selon
´ ´ ´
le regime alimentaire, des modifications similaires de profils de composition en acides gras. Les acides gras deposes avant le ˆ
´ `
sevrage influencent la composition des acides gras des depots adipeux et des muscles plusieurs mois apres le sevrage.
*Corresponding author. Tel.:133-1-4408-1767; fax: 33-1-4408-1853.
E-mail address: [email protected] (P. Bas)
Lorsque les rations sont riches en pulpes de betteraves ou en farine de poisson, la proportion de C18:1est significativement
´ ´ ¨
augmentee dans les tissus alors que les proportions de C18:0, C18:2et C18:3sont abaissees. L’incorporation de maıs dans la ˆ
´ ´
ration induit une augmentation de la teneur en acide linoleique dans les depots adipeux. L’incorporation de tourteau de coton
´ ´ `
induit un accroissement des teneurs en acides stearique et linoleique dans les tissus des agneaux. Les rations a base d’herbe ˆ
` ´ ´ ´
conduisent a des teneurs elevees en C18:0et en C18:3dans les depots adipeux des agneaux. Les variations du point de fusion
´ ´ ´ ´ `
des tissus sous-cutane et perirenal sont fortement associees a celles des teneurs en C18:0. Cette approche souligne les ˆ
´ ´ ´
difficultes pour comprendre les effets d’un regime alimentaire sur la composition des depots gras et des muscles lorsque l’on ´
ne prend pas en compte tous les aspects de l’alimentation et de l’elevage. Bien que les lipides alimentaires soient fortement
´ ´ ´ ´
hydrogenes dans le rumen, cette etude montre l’amplitude des modifications des proportions des differents acides gras dans ´
les carcasses d’agneaux qui depend du choix des aliments de la ration. 2000 Elsevier Science B.V. All rights reserved.
Keywords: Lamb; Fat deposits; Muscles; Fatty acid composition; Diet; Energy sources; Nitrogen sources
1. Introduction
feeding programs on the fatty acid composition of
lamb adipose tissues and muscles from available
Adipose deposits strongly influence the quality of
bibliographic data.
ruminant carcasses through their amount and
com-position. The effects of nutritional factors on the
proportion and body distribution of adipose deposits
2. Material and methods
´
have been reviewed by Vezinhet and Prud’hon
(1975), Kempster (1981), Robelin (1986) and Bas
Our work first consisted in selecting articles
(1993).
dealing with fatty acid composition of lamb adipose
Nutritional factors have a lesser influence on the
deposits and muscles, with a stress on those
analys-fatty acid composition of adipose tissues and muscles
ing the effects of dietary factors or having other
in ruminants than in monogastrics because of the low
objectives but supplying enough information on diets
lipid content of their diet and of the hydrogenation of
and feeding management. All the papers were taken
dietary lipids in the rumen (Wood and Enser, 1997;
except when they did not provide sufficient
infor-¨
Table 1
Main animal characteristics of the data base
a b
Sex n ADG Weaning age (WA) Slaughter LW (SLW) Slaughter age (SA)
(g / day) n (weeks) n (kg) n (weeks) n
M 425 0,ADG,100 20 0,WA,5 190 SLW,10 26 0,SA,5 38
F 136 10,ADG,150 69 5,WA,10 340 10,SLW,20 35 5,SA,10 73
C 174 150,ADG,200 73 10,WA,15 35 20,SLW,30 69 10,SA,15 141
Mixed sexes 152 200,ADG,250 122 15,WA,20 25 30,SLW,40 192 15,SA,20 188 250,ADG,300 157 20,WA,30 10 40,SLW,50 256 20,SA,50 161
ADG.300 74 SLW.50 81 SA.50 106
a
Sex: M, entire males; F, females; C, castrated males; mixed sexes, entire males1castrated males or entire males1females or castrated males1females
b
ADG, average daily gain; LW, live weight; n, number of observations.
dock (tail basis), costal, leg, inguinal and sternal
bibliographic data base thus formed contained 979
regions, the Semimembranosus and Triceps Brachii,
observations from 108 papers.
and around the fat tail. The information on the fatty
Data were analysed by the Factor, GLM, REG and
acid composition of MU came most frequently from
Nlin procedures of SAS (SAS, 1987). The GLM
Longissimus dorsi, then from Semimembranosus
procedure was used with the factors most frequently
(Table 3) and lastly from Triceps Brachii.
represented.
The variables concerning diet components were
the type of forage and the main energy, protein, and
fat sources in concentrates or in complete diets.
3. Results
Other variables, when not provided in the articles,
were calculated from available information such as
3.1. General characteristics of adipose tissues and
the proportion of roughage and concentrate in diets,
muscles
metabolisable energy, total protein, crude fibre, and
fat contents of the complete diets and concentrates,
The fatty acid composition of three adipose tissues
while missing nutritive values were assessed using
frequently mentioned in the data base, namely
sub-the INRA tables of feedstuff values (Andrieu et al.,
cutaneous, omental, perirenal, and that of one
intra-1989).
muscular tissue are reported in Table 2. In these
During milk- and post-weaning periods, ten types
tissues three main fatty acids (C
16:0, C
18:0, and C
18:1)
of diets were distinguished: ewes’ milk, milk re-
represented the major part of total FA (from 78% in
placer, ewes’ milk and concentrate, ewes’ milk and
MU to 87% in OM). The coefficients of variation of
pasture, roughage alone, pasture alone, roughage and
C
16:0and C
18:1proportions by weight were lower
concentrate, pasture and concentrate, concentrate
(CV
5
18.5 and 14.2%, respectively) than that of
alone, complete diet. The two latter diets were
C
18:0(CV
5
40%). The intra-tissue variability for
defined by using thresholds for the proportions of
C
16:0and C
18:1was approximately 10%, but was far
main feedstuffs. For example, diets were classified in
greater for C
18:0(reaching 20% in PR and 37% in
complete diets when the mixed diets contained more
SC).
Table 2
Mean fatty acid composition of the main fat deposits in lambs
b
Fat Fatty acids
a
deposits C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C17:0
a a a a a a a a
SC 4.8 23.4 15.4 3.5 40.8 4.0 1.4 2.4
b b b b b b a b
PR 4.0 20.6 26.1 2.7 36.7 4.9 1.5 1.9
a,b a,c c a,b,c c a,b b a,b
OM 4.6 23.7 29.9 2.9 33.6 4.3 0.8 2.1
b c a c a c b c
MU 3.8 22.5 15.6 3.2 40.4 6.3 1.1 1.4
a
SC, subcutaneous adipose tissue (250,x,411); PR, perirenal adipose tissue (111,x,219); OM, omental adipose tissue (7,x,17); MU, intramuscular fat (86,x,143); x, number of observations for each fatty acid.
b
Superscripts a, b and c, means in a column with different superscripts were significantly different (P,0.05).
antagonism between the percentages of C
14:0and
were negatively correlated (r
5 2
0.33, P
,
0.05)
C
16:0and that of C
18:2and, on the second factor, an
and in SC, MP was significantly in opposition to
antagonism between the percentages of C
14:0, C
18:0odd-numbered and branched-chain fatty acid
per-and C
18:3and those of C
17:0and C
18:1. But these
centages. For these two tissues, the MP was best
relationships showed noticeable differences between
explained from the C
18:0percentage, with a
coeffi-the tissues. In PR (Fig. 1C), coeffi-the percentages of
cient of determination over 75% (Fig. 2). The two
saturated acids (C
16:0, C
17:0, and C
18:0) were op-
prediction equations became not significantly
differ-posed to those of polyunsaturated fatty acids (C
18:2ent when the C
18:0and the C
18:2percentages were
and C
18:3) on the first factor axis. In SC (Fig. 1B),
used simultaneously as explicit variables.
there was an opposition between the percentages of
MP (
8
C)
5
0.51
?
C
18:02
0.28
?
C
18:21
29.0;
C
17:0and C
18:1and those of C
14:0C
16:0, C
18:0and
2
C
18:3on the first factor, and in MU (Fig. 1D), C
18:0,
(R
5
0.86, RSD
5
1.7, n
5
101)
C
18:1, and C
18:3percentages were opposed to those
For one group of animals, the mean difference in MP
of C
17:0and C
18:2on the first factor axis also. The
between the PR and SC from caudal region, which
second factor axis revealed an opposition between
approximatively represented 75% of the SC samples,
C
18:1and C
14:0in PR (Fig. 1C), and between C
18:2analysed for MP, was about 6.4
8
C.
and C
18:1in SC (Fig. 1B).
The SI was less closely linked to fatty acid
The relationship between the melting point (MP)
proportion than was the MP probably because of its
and the percentage of the main fatty acids were
more subjective estimation and the important
vari-studied in SC and PR. The softness index (SI) was
ability of SC composition according to sampling
studied in SC only, and most particularly in samples
sites. In SC, the C
percentage was the variable
from the dorsal region. The SI values were stan-
16:0which predicted SI best. Taking into account a
dardised on a scale varying from 1 (very firm) to 5
second fatty acid percentage in the equation
im-(very soft and oily). On the whole, the MP was
proved slightly but not significantly the prediction of
linked to the same acids in PR and in SC. The
the SI.
coefficients of correlation between the MP and the
percentages of C
16:0and C
18:0were significant and
SI
5 2
0.22
?
C
1
7.3;
16:0
positive (r
5
0.32, P
,
0.05 and 0.87, P
,
0.001, in
2
(R
5
0.40, RSD
5
0.75, n
5
67)
PR, 40
,
n
,
49, and r
5
0.38, P
,
0.01, r
5
0.88,
P
,
0.001, in SC, 49
,
n
,
58, respectively). They
were significant and negative between the MP and
3.2. Variation of fatty acid composition due to
the percentages of C
16:1, C
18:1, and C
18:2(
2
0.60,
sampling site
P
,
0.001;
2
0.54, P
,
0.001;
2
0.62, P
,
0.001, in
even-numbered saturated fatty acids. But the C
18:1percentage was clearly higher in leg than in back
samples.
In spite of the low representivity of the
Semimem-branosus, one may say it was characterised by higher
percentages of unsaturated fatty acids (C
16:1, C
18:1,
C
18:2, and C
18:3) and by a lower percentage of C
18:0than Longissimus dorsi.
As variability in compositions was related to
sampling sites, and as the number of samples
avail-able varied from one site to another, because of the
higher number of samples indicated in Table 3, the
back site of SC and the Longissimus dorsi site of
MU were favourably considered to study the effects
of diets on the fatty acid composition of tissues.
3.3. Effect of milk feeding on the adipose tissue
composition
[image:6.612.58.247.72.335.2]As shown in Table 4 (ewes’ milk fed lambs) and
in comparison with Tables 5 and 6 (weaned lambs),
the fatty acid composition of tissues from lambs fed
Fig. 2. Relation between the melting point and the C18:0content
ewe milk was characterised by lower percentages of
of adipose tissues.C
18:0, C
18:2, and C
18:3and higher percentages of
C
14:0, C
16:0, and C
18:1than that of tissues from
anatomic site of sampling of OM was not precisely
lambs slaughtered after weaning. In the most cases,
known as for PR. Whereas SC and MU were
the composition of adipose tissues reflected the fatty
analysed in accurately defined locations in about
acid composition of ewe milk which is rich in C
14:090% of the cases.
and C
16:0in comparison to post-weaning diets. The
In Table 3, SC samples from the inguinal region
percentages of total saturated fatty acids did not
were characterised by the highest contents of satu-
differ between SC and PR, but SC had a lower C
18:0rated fatty acids C
16:0and C
18:0. On the opposite,
percentage and higher C
14:0and C
16:0percentages
back and leg samples were the poorest in total
than PR. During the milk feeding period, the C
18:1Table 3
1
Changes in fatty acid composition of fat deposits according to sample location
AD Sample location Fatty acids
C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C17:0
a a a a a a a a
SC Back 4.2 22.2 16.7 3.6 39.5 4.5 1.5 2.4
a b a a b a a a
Dock 4.1 24.4 16.1 3.3 41.2 3.9 1.6 2.5
a b a a b a a
Costal 4.0 24.4 15.6 3.2 42.5 3.4 2 2.7
a a,b a a b a a a
Leg 5.0 23.9 14.4 3.0 43.8 3.6 1.6 1.6
a b b a b a
Inguinal 4.0 25.2 20.2 4.5 42.9 3.5 2 2
a a a a a a a a
MU Longissimus dorsi 3.0 22.7 15.7 3.8 41.4 7.3 1.0 1.5
a a b a a b b a
Semimembranous 3.2 22.3 13.7 4.1 43.8 10.3 1.9 1.4
1
AD, adipose tissues; SC, subcutaneous adipose tissue; MU, intramuscular fat. Sample location, back (131,n,181); dock (42,n,
[image:6.612.47.499.514.624.2]Table 4
1
Fatty acid composition from three fat deposits in lambs fed ewe milk
Fat Fatty acids
deposits C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C17:0
a a a a a a a a
SC 9.4 24.8 11.7 4.0 40.0 3.1 0.8 1.5
b b b a a a b a
PR 6.8 22.2 17.1 2.9 40.2 3.8 1.2 1.5
b b a b a b b b
MU 6.6 21.7 13.0 2.3 40.4 5.9 1.4 1.0
1
[image:7.612.45.500.256.341.2]SC, subcutaneous adipose tissue (12,n,18); PR, perirenal adipose tissue (4,n,8); MU, intramuscular fat (n56); n, number of observations for each fatty acid. Superscripts a and b, means in a column with different superscripts were significantly different (P,0.05).
Table 5
1
Effects of types of diets on the mean fatty acid composition of three fat deposits in lambs
Diets Fatty acids
C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C17:0
a a a a,c a a a a
Pasture 4.2 21.5 23.6 2.8 36.9 4.2 2.4 1.2
a,b,c a,b a a,b a,b a,b,c b
Past1conc. 3.4 21.5 22.6 3.7 37.4 5.2 1.5 2
b,c a b a,c b b c b
Rough1conc. 3.6 21.9 17.6 3.0 39.5 5.1 0.5 2.6
c b b b c c c c
Concentrate 2.9 20.4 17.7 3.7 41.5 6.1 0.7 2.3
b c c c a c b c
Complete diet 3.6 23.4 19.9 2.7 38.2 6.4 2.0 2.1
1
SC, subcutaneous adipose tissue from the mid back area; PR, perirenal adipose tissue; MU, muscle (Longissimus dorsi). Pasture, pasture alone (21,n,45); Past1conc., pasture and concentrate (8,n,12); Rough1conc., roughage and concentrate (57,n,78); Concentrate, concentrate alone (161,n,236); complete diet, mixed compound diet with roughage and concentrate (64,n,120); n, number of observations for each fatty acid. Superscripts a–d, LS means in a column with different superscripts were significantly different (P,0.05); the LS means of the five diets was calculated for three adipose tissues.
Table 6
1
Effects of three types of diets on fatty acid composition of three fat deposits in lambs
AD Diets Fatty acids
C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C15:0 C17:0 C17:1
a a a a,b a,b a a a a a
SC Rough1conc. 3.2 21.6 11.7 3.7 40.8 3.7 0.4 1.3 3.4 3.1
a a b a a b a b b b
Concentrate 3.4 20.9 15.8 4.1 42.5 5.6 0.8 0.8 2.7 1.6
b b b b b b b a b b
Complete diet 4.7 24.9 16.2 3.2 39.0 6.0 2.2 1.1 2.6 1.5
a a a a a a a a a a
PR Rough1conc. 3.6 20.6 26.2 2.5 36.2 4.7 0.4 0.5 2.4 0.9
b b b b b a a a a a
Concentrate 2.7 19.2 24.1 3.1 40.0 5.4 0.7 0.6 2.2 1.0
ab a a a,b a b b a b
Complete diet 3.1 21.3 27.5 2.7 35.6 6.6 2.6 2 2.4 0.6
a a a a a a ab a a a
MU Rough1conc. 4.0 23.6 14.7 2.7 41.9 7.0 0.5 0.6 2.0 1.4
b b b b a a a a a a
Concentrate 2.9 20.8 13.3 4.2 41.0 7.2 0.5 0.6 1.9 1.5
c a c c a b b b
Complete diet 2.0 23.1 17.0 2.0 40.5 5.5 0.8 2 1.1 2
1
[image:7.612.45.507.470.605.2]Fig. 4. Relation between the FA content of all fat deposits in Fig. Fig. 3. Effect on age of the C18:1content of fat deposits in milk
3 and the corresponding FA content in the liquid milk fed lambs. fed lambs.
C12:0 AD50.65?exp(0.077?C12:0 D), RSD51.5; C14:0 AD50.24?
exp(0.30?C14:0 D), RSD52.1; C16:0 AD536.6235.1?exp(0.033?
C16:0 D), RSD53.5; C18:0 AD515.3214.3?exp(0.20?C18:0 D),
content in SC, PR, and MU (Fig. 3) regularly
RSD53.0; C 5105.4286.7?exp(0.012?C ), RSD55.7;18:1 AD 18:1 D
decreased with the age of lambs by about 1% per
C18:2 AD5138.22136.9?exp(0.008?C18:2 D), RSD52.2; Cn:0ADand Cn:0 5FA content in tissues and in milk replacer,
respective-week, whereas C
16:1and C
18:2contents increased.
Dly.
As with ewe milk, the fatty acid percentages of
adipose tissues from milk-replacer fed lambs varied
with the lipid composition of milk replacers (Fig. 4).
Increasing the C
12:0supply in milk increased the
high percentage of total C
18acids in post-weaning
tissular content in C
12:0but also in C
14:0and C
16:0.
diets and the hydrogenation potential of the
unsatu-Likewise, an increase in the C
16:0content in milk
rated C
18acids in the rumen. The fatty acid profile of
replacers increased the C
16:0, C
18:0and C
18:1per-
tissues, which reflects the milk feeding period,
centages in tissues. Moreover, an increase in C
18:3in
gradually disappeared after weaning.
milk replacers had favourable repercussions on the
C
18:3and in C
18:2percentages in tissues. Thus,
3.4. Influence of post weaning feeding
knowledge of fatty acid percentages in milk replacers
made predictions of the fatty acid composition in
3.4.1. Effects of the type of diets
[image:8.612.290.494.81.329.2]anatomical location of the fat deposit. For example
SC, PR and MU from lambs receiving the complete
diet were all characterised by a lower C
18:1per-centage and a higher perper-centage of C
18:3and total
saturated fatty acids than with other diets. In SC, the
variability of the C
18:0percentage reached nearly
twice that of other tissues (CV
5
33, 19, and 15%,
respectively for SC, PR, and MU). In SC again, the
percentage of C
18:0was relatively low whereas those
of odd-numbered and branched-chain fatty acids
were high. The concentrate alone diet presented the
highest C
18:1percentage and lowest percentage of
total saturated fatty acids. On the whole, the pasture
alone and pasture and concentrate diets on the one
hand and concentrate alone and roughage and
con-centrate diets on the other hand resulted in very
similar compositions in each tissue.
[image:9.612.64.243.76.329.2]When crude protein (CP), crude fibre (CF) ether
extract (EE) and energy values of diets were together
taken into account, the coefficients of determination
of most fatty acid percentages, except C
14:0, were
strongly increased. With these variables and when
Fig. 5. Variation of the C14:0 and of the C18:0 content of fat
deposits with time (t) after weaning. C18:0 SC50.12t112.6;
considering the tissue and diet effects, the
coeffi-P50.06, RSD52.4; C18:0PR50.25t119.1; P,0.01; RSD52.5;
cients of determination of the percentages of the
C18:0MU512.8, SD51.3; C14:0SC58.4?exp(20.097t), RSD5main fatty acids were increased from 27, 57, and
2.8; C14:0 PR54.8?exp(20.092t), RSD51.5; C14:0 MU55.8?
17% to 55, 65, and 35%, for C
16:0, C
18:0, and C
18:1,
exp(20.090t), RSD51.8.
respectively. The richest diets in terms of
metabolis-able energy were generally associated with the
lowest C
18:0, C
18:1and C
18:2percentages and the
fatty acid composition of tissues between the types
highest C
16:0, C
16:1, C
15:0and C
17:0percentages in
of diets was relatively limited although significant.
SC, PR, and MU. With higher crude protein content
Extreme concentrations were observed with the
in diets lamb tissues were poorer in C
16:0and in
pasture alone and concentrate alone diets.
C
18:0and richer in C
16:1, C
18:1and C
18:3. The
With the pasture alone diet, tissues were richer in
increasing crude fibre content in diets were
associ-C
14:0, C
18:0and C
18:3and poorer in C
18:1, C
18:2and
ated with increase in C
18:0percentage and with
C
17:0. In lambs reared on pasture alone, the C
18:3decrease in C
18:2percentage in tissues. The ether
percentage in all three tissues was relatively high
extract contents of diets were negatively correlated
(2.7, 2.6, and 1.7%, in SC, PR, and MU, respective-
with the percentages of C
16:0, C
18:3, C
15:0and C
17:0Table 7
1
Effects of the main sources of energy of the concentrate on the fatty acid composition of lamb fat deposits Energy source Fatty acids
C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C15:0 C17:0 C17:1
a a a a,b a a a,b a a,b
Beet pulp 3.1 21.1 15.0 2.2 41.8 3.8 0.7 0.8 1.9 2
a b a,b a a a a a a a
Barley 3.1 23.0 18.1 2.1 39.6 4.3 0.9 0.9 1.9 1.7
a a b b a b b a b a
Maize 3.1 21.4 18.9 2.7 38.8 6.5 1.2 1.1 2.4 1.3
a c c a,b b a a,b a a,b
Wheat 3.5 24.7 21.6 2.8 36.0 4.6 0.8 0.7 2.0 2
1
Beet pulp (n58); Barley (31,n,92); Maize (33,n,131); Wheat (5,n,18); n, number of observations for each fatty acid. Superscripts a,b,c, LS means in a column with different superscripts were significantly different (P,0.05); The LS means of the four sources of energy was calculated for three adipose tissues, SC: subcutaneous adipose tissue from the mid-back area, PR: perirenal adipose tissue, MU: muscle (Longissimus dorsi).
3.4.2. Energy sources
ton, 1971). The similarity between the compositions
The energy sources of the concentrates or the
of all tissues from lambs receiving semi-synthetic
complete diets significantly influenced the percent-
diets or beet pulp based diets could be the result of
ages of most fatty acids. The C
18:2and C
18:3the low fat contents of these two diets. Thus, in both
percentages in tissues were higher when maize was
cases the fatty acids in the tissues do not originate
the main energy supply (Table 7) probably because
from the diets but from a de novo synthesis. The
it has a higher fat content than the other energy
tissues of lambs fed diets rich in wheat had a low
sources and because of its high C
18:2content. As
percentage of C
18:1and high percentages of saturated
shown in Fig. 6, the C
18:2percentage in SC, PR, and
fatty acids. A rise in the level of incorporation of
MU increased as the proportion of maize increased
barley or maize in the diets led to significant
in the diet, whereas the C
16:0percentage decreased.
increases in C
17:0percentages in SC, PR, and MU,
Beet pulp decreased the percentages of saturated
and a significant decrease in the C
18:0in SC.
fatty acids (C
16:0and C
18:0) in tissues, in the same
way as with semi-synthetic diets based on glucose or
3.4.3. Protein sources
starch (Tove and Matrone, 1962; Duncan and Gar-
Ingredients included in diets to increase their
protein contents may also influence the fatty acid
composition of adipose tissues and muscles of lambs
(Table 8). Addition of cotton meal as the main
protein source brought about a significant raise in
C
18:0and C
18:2percentages in tissues, probably
because of the high content of total unsaturated C
18acids in cotton lipids. But when lambs received
complete, concentrate alone or roughage and
concen-trate diets rich in cotton meal, the tissues had low
C
18:3, C
18:1and C
16:0contents. These effects
per-sisted when crude protein, crude fibre, and ether
extract content were at even levels in the diets. When
incorporating fish meal in lamb diets, C
18:0and C
18:2percentages in the tissues were reduced, whereas
C
18:1percentage was increased. Alfalfa meal, rich in
linolenic acid, induced a higher percentage of C
18:3in tissues than the three other protein sources shown
in Table 8. When the proportion of alfalfa meal was
increased in a diet, C
16:1, C
18:1and C
18:2percentages
were generally lower in tissues. But, as shown by
Fig. 6. Relation between the C18:2content of adipose tissues and
[image:10.612.61.246.434.641.2]Table 8
1
Effects of the main sources of nitrogen of the concentrate on the fatty acid composition of lamb fat deposits Energy source Fatty acids
C14:0 C16:0 C18:0 C16:1 C18:1 C18:2 C18:3 C15:0 C17:0 C17:1
a a a a,b a a b b b a
Alfalfa meal 3.1 21.8 19.5 2.8 38.1 5.7 1.4 1.0 2.4 0.9
a a,b a a a,c a,b a a a
Soybean meal 3.2 22.5 19.3 2.3 39.3 5.1 0.8 1.3 1.7 2
a c b b b c a b c a
Cotton meal 3.3 19.3 29.2 1.4 32.0 8.3 0.4 0.6 1.1 1.4
a a,b c a,b c b a,b
Fish meal 2.9 22.4 15.8 2.7 41.2 3.9 0.6 2 2 2
1
Soybean meal (18,n,66); alfalfa meal (20,n,115); fish meal (6,n,18); cotton meal (12,n,16); n, number of observations for each fatty acid. Superscripts a, b and c, LS means in a column with different superscripts were significantly different (P,0.05). The LS means of the four sources of nitrogen was calculated for three adipose tissues, SC: subcutaneous adipose tissue from the mid back area, PR: perirenal adipose tissue, MU: muscle (Longissimus dorsi).
presented some difficulties because the number of
records differed widely between fatty acids, tissues
and diets. The number of missing data for the main
fatty acids varied from 52 for C
18:0to 338 for C
18:3and reached 423, 528, and 331 for the odd-numbered
fatty acids: C
15:0, C
17:0, and C
17:1, respectively.
The type of diet could be defined for nearly 95%
of the observations but the type of roughage and the
main sources of energy and nitrogen could not be
defined for 40–50% of the observations. The
chemi-cal compositions of the concentrate alone and
com-plete diets were determined in more than 80% of the
cases but the chemical composition of post-weaning
diets could be estimated in only 70% of the cases
because
information
about
the
unprocessed
roughage / concentrate ratio in diets was often
miss-ing.
In addition, information was often incomplete or
Fig. 7. Relation between the C18:1content of adipose tissues and
inaccurate on other aspects of the experiments,
muscles and the alfalfa content of the solid diets. C18:1 SC5
including details of the method of feeding, the
2
38.610.24x20.0035x , RSD52.4; n557; C18:0 PR531.01
number of times feed was offered each day, and of
2
0.37x20.0048x , RSD52.9; n542; C18:0 MU551.220.22x2
2
genotype, sex and stage of growth of the lambs. In
0.0013x , RSD51.74; n512.
some cases the imprecision about weight and age at
weaning or slaughter of the lambs also raised
difficulties.
to take place only when incorporation rate in the diet
was above 40–50%.
4.2. Validity of the data base contents
4. Discussion
When available or reliable information was
[image:11.612.57.250.251.460.2]and the chemical compositions of roughage and
to that of C
18:1. It is already known that unsaturated
concentrate feeds using the tables of INRA (Andrieu
fatty acids C
18:1, C
18:2, and C
18:3and odd-numbered
et al., 1989). When the diet contained no hay, the
fatty acids such as C
17:0largely contribute to lower
level of straw intake could be estimated when it was
fat melting points. When the availability of
polyun-distributed in a trough, but was not taken into
saturated fatty acids is high, there may be a
substitu-account when no description about straw distribution
tion by a decrease in C
18:1content in subcutaneous
was found. Consequently, some diets may have been
adipose tissues probably by the antagonist action on
falsely identified as roughage and concentrate or
microsomial desaturases. Moreover, an increase in
concentrate alone.
odd-numbered fatty acid percentages, often
associ-Percentages of the fatty acids studied depended on
ated with an increase in the percentages of
branched-the procedures used in branched-the extraction and esterifica-
chain fatty acids resulting from a de novo synthesis
tion of lipid samples, and identification and quantifi-
from propionate (Garton et al., 1972b), cause a
cation of the melting point of each peak separated by
decrease in the percentage of the even-numbered
gas–liquid chromatography. The values expressed in
saturated fatty acids with highest melting points.
moles were corrected to percentages by weight, but
In PR, the negative correlation between PUFA and
authors generally gave no information on the thres-
long-chain saturated fatty acid percentages indicate
hold level for quantification and the fatty acids that
an antagonist action of PUFA, when available in
were included in the total percentage.
relatively large amounts, on the storage of saturated
These methodological criteria particularly influ-
fatty acids. As the activity of desaturase may be
enced unsaturated, odd-numbered and branched-
lower in PR than in SC (Whale and Garton, 1972)
chain fatty acid percentages (Beriain et al., 2000a). A
the synthesis of C
18:1may depend on available
bias in the interpretation of the results may have
amounts of C
14:0.
occurred because of the differences in the number of
In MU lipids, which generally contain more than
observations for different predictive variables. For
50% of phospholipids, the fatty acid composition
example, with castrated males there was very limited
play an important part in the fluidity of membranes.
nutritional information and a tendency for the lambs
In MU, variations in the percentages of C
18:2and
to be slaughtered later than either entire males or
C
17:0seem to be opposed to that of C
18:0, C
18:1, and
females.
C
18:3. Moreover, the percentages of C
18:0and C
18:2seemed to be modulated by those of shorter-chain,
4.3. General characteristics of fat deposits
even-numbered saturated fatty acids: C
14:0, and
C
16:0.
The differences in compositions between tissues
Although melting point values vary widely with
and between sampling sites observed in this study,
the methodology used (Vimini et al., 1984),
varia-irrespective of physiological, dietary, climatic and
tions in Softness Indexes may be closely linked to
management conditions, reinforced well established
fatty acid composition. The causes of the lowering of
results. Generally, adipose deposits were character-
melting points are numerous. Some authors attribute
ised by a saturation index related with the location of
an essential part to increases in odd-numbered and
the tissue, and by low melting points in tissues more
branched-chain fatty acids (Garton et al., 1972a; Bas
exposed to environmental temperatures (Cramer and
et al., 1980, 1998; Miller et al., 1980) while under
Marchello, 1964; L’Estrange and Mulvihill, 1975;
other conditions monoenic (L’Estrange and
Mul-Bas et al., 1987). For example, SC showed a
vihill, 1975; Gibney and L’Estrange, 1975) and
decrease in saturation between the back which is
dienic fatty acids (Gibney and L’Estrange, 1975)
most exposed to outside temperatures, and the dock
seem to be responsible for the lowering of fat
which is in turn less saturated than the inguinal site.
melting points. The latter study indicated that
what-In SC, our results indicate that percentages of
ever experimental conditions, the lowering of fat
even-numbered saturated fatty acids were associated
melting points was best explained by a fall in C
18:0factors, like the variations of the percentages of
of the post-weaning period (Bas, 1993), the relative
position isomers and more markedly geometric iso-
prevalence of one of the two processes of lipid
mers of fatty acids, probably have a significant
storage which differ from one tissue to another,
influence on melting point, but isomers of fatty acids
namely the de novo synthesis or from circulating
have been very seldom studied in the literature
fatty acids (Payne and Masters, 1971; Ingle et al.,
analysed in this data base.
1972a,b; Haugebak et al., 1974; Vezinhet et al.,
1983). This is also affected by other factors,
includ-ing age, sex, level of fatteninclud-ing.
4.4. Influence of feeding on the fatty acid
Dietary lipid of energy and protein sources in the
composition of tissues
data base, except fish meal, contained between 75
and 90% of C
18fatty acids, mainly PUFA (dienic
In young lambs fed milk rich in lipids, the profile
fatty acids in cereals, soybean meal, cotton seeds or
of fatty acids of fat deposits and muscles largely
cotton meal and trienic fatty acids in alfalfa and
reflected the dietary fatty acid profile, but the com-
grass). These dietary lipids are hydrolysed in the
position of the various tissues revealed some differ-
rumen and the PUFA released changed into cis or
ences in relation with their functional specificity.
trans monoenic fatty acids, mostly C
18:0. The degree
Thus, this study clarified the influence of both
of hydrolysis of these PUFA varies from 70 to 100%
amount and nature of dietary fat on concomitant
when they are not protected (Doreau and Ferlay,
variations of fatty acid composition in each adipose
1994).
tissues and muscle. The influence of dietary fatty
With diets containing beet pulp or fish meal as
acids on body composition was particularly marked
main energy or nitrogen sources, lipid compositions
at this stage of growth because of the high lipid
of the different fat deposits were similar and
char-accretion almost exclusively from circulating fatty
acterised by low levels of C
18:0, C
18:2, C
18:3and total
acids. However, the results of this study suggested
saturated fatty acids, and by a high level of C
18:1.
that C
12:0, C
14:0, and C
16:0, when available at a high
This may be the result of low C
18fatty acid uptake
level, may be successively elongated to stearic acid
into tissues resulting from the low contents of these
and converted into monoenic fatty acids with 16 or
fatty acids in these two ingredients. The storage of
18 atoms of carbon by desaturation. On the other
these acids from fish meal may also be reduced by
hand, long-chain fatty acids did not appear to be
inhibition of both fatty acid uptake and rumen
converted into smaller-chain fatty acids. When the
microbial synthesis of fatty acids in the presence of
diet contained very small amounts of PUFA, the
PUFA with chains longer than 20 carbon atoms
PUFA percentage was often high in adipose tissues
(Storry et al., 1974). Long-chain PUFA from fish
and even higher in muscles, probably because of
meal seems to be hydrogenated in the rumen to a
their higher coefficients of digestibility, as compared
lesser extent than C
18fatty acids and seems to be
to other fatty acids (Walker and Stokes, 1970), and
neither reduced into shorter fatty acids nor deposited
also to their low degradation in tissues because of
in the triacylglycerols of adipose tissues and muscles
their preferential position on C2 of triacylglycerols
although they may be stored as phospholipids (Ashes
(Hawke et al., 1977).
et al., 1992).
percentages in tissues of these lambs. On the other
5. Conclusion
hand, the high levels of linoleic and linolenic acids
in tissues and unchanged percentage of stearic acid,
In spite of the difficulties met in comparing fatty
which is terminal product of PUFA hydrogenation
acid compositions obtained with different procedures
seems to indicate that PUFA from maize are less
of quantification, and a lack of information
con-easily hydrogenated than PUFA from other cereals or
cerning management conditions and diet
characteris-meals. This resistance of maize lipids to hydro-
tics, the results obtained from this review of the
genation may result from the lower ruminal de-
literature nevertheless contribute to a better
apprecia-gradability of maize protein and starch than that of
tion of the magnitude of feed effects on the fatty acid
other cereals (Nocek and Tamminga, 1991). Maize
composition of lamb tissues. All raw materials
may lead to relatively high levels of odd-numbered
contained in diets may have an influence on the fatty
fatty acids as the synthesis of fatty acids by rumen
acid composition of lamb tissues by both amount and
microorganisms is known to be at least twice as high
composition of lipids in each ingredient, in relation
with maize than with barley diets (Brumby et al.,
with nature, fibrosity and chemical composition of
1979), and as lipids synthesised by microorganisms
the total diet. This study, which consisted in making
are relatively rich in odd-numbered fatty acids
and analysing a data base, gives an objective view of
(Ifkovits and Ragheb, 1968; Viviani, 1970; Bas et al.,
the effects of dietary ingredients on the lipid
com-2000).
position of tissues and quality of lamb carcasses.
The C
18PUFA from wheat and barley appears to
Better control of the lipid composition of lamb
be more hydrolysed and hydrogenated in the rumen
carcasses would result from carefully considering
than that from maize. Wheat tended to produce a
these dietary factors when managing lambs and
higher percentage of stearic acid and a lower per-
designing their diets.
centage of oleic acid in tissues than barley as a result
of its higher levels of C
18PUFA and lower levels in
oleic acid. But, with wheat and barley, the odd-
6. For further reading
numbered and branched chain fatty acid percentages
were often increased in SC with a concomitant
AL-Shabibi and Juma, 1973; Anderson et al.,
reduction in stearic acid. These fatty acids could
1977; Aurousseau et al., 1973; Banskalieva, 1997;
result from de novo synthesis from propionate as the
Banskalieva et al., 1992; Banskalieva et al., 1988;
high content of rapidly rumen-degradable starch in
Bas et al., 1997; Bellof et al., 1995; Bellof et al.,
wheat and barley decreases ruminal pH and increases
1997; Beriain et al., 2000b; Bouillier-Oudot et al.,
production of propionate.
1992; Boylan et al., 1976; Bozzolo and
Bouillier-The particularly high percentage of linolenic acid
Oudot, 1999; Bozzolo et al., 1990; Busboom et al.,
in adipose tissues and muscles with a grass-based
1981; Caballero Garcia de Arevalo et al., 1992;
diet (2–3% fat / DM) may result from the high
Cameron et al., 1994; Casey et al., 1988; Casey and
linolenic acid content in grass (
.
50% of total fatty
Weeb, 1995; Ch’ang et al., 1980; Cirruzzi et al.,
acids), a part of which would resist to hydrogenation
1993; Ciruzzi et al., 1977; Cramer et al., 1967;
because of the high level of linoleic acid itself. There
Crouse et al., 1972a; Crouse et al., 1972b; Demise et
may also be considerable protection against hydro-
al., 1998; Devier and Pfander, 1974; Dimov and
genation of intracellular galactosylglycerols in the
Banskalieva, 1989; Duncan and Garton, 1967;
Dun-rumen when the feed was not in crushed or ground
can and Garton, 1978; Duncan et al., 1972;
El-forms (Ben Salem et al., 1993). Similarly, the
Shahat et al., 1988; Enser et al., 1996; Enser et al.,
´
castrated and non castrated hoggets fed clenbuterol. In: Proc.
et al., 1972b; Jacobs et al., 1973; Jenkins et al., 1994;
38th Internatinal Congress of Meat Science and Technology,
Johnson et al., 1988; Kemp et al., 1981; L’Estrange
Clermont-Ferrand, France, 23–23:08 / 92, Vol. II, pp. 1–4.
and Hanrahan, 1980; L’Estrange, 1977; L’Estrange,
Banskalieva, V., Shindarska, Z., Dardzhnov, T., Dimov, V., 1988.1979; Leat et al., 1997; Leth et al., 1998; Lough et
[Effect of different amounts of energy and protein in the diet on fatty acids in subcutaneous fat depots of fattening lambs].al., 1992; Manfredini and Cavani, 1979; Marinova et
Zhivotnov’dni Nauki 25, 73–78.
al., 1992; Marinucci et al., 1998; Matthes et al.,
´
Bas, P., 1993. Croissance et metabolisme lipidique du ruminant
1998; Matthes et al., 1996; Miller and Rice, 1967a;
autour du sevrage. Etude sur modele caprin. In: Thesis Dr de`´
´
Miller et al., 1967b; Molenat and Theriez, 1973;
l’Institut National Agronomique. `Bas, P., Archimede, H., Rouzeau A., Sauvant, D., 2000. Influence
Murphy and L’Estrange, 1977; Muscio et al., 1978;
of forage level and forage source on the fatty acid composition
¨
Muscio et al., 1980; Normand et al., 2000; Nurnberg
of the rumen microorganisms in the goats. FAO Meeting,
et al., 1996; Ørskov et al., 1975; Pearce et al., 1974;
Grignon, France, 03–05 / 09 / 98, Options Mediterranneenes,´ ´ ´Petit et al., 1997; Purchas et al., 1979; Quackebeke et
Serie A, Seminaires Mediterranneens, in press.´ ´ ´ ´ ´al., 1978; Ray et al., 1975; Rule et al., 1991; Safari et
Bas, P., Chilliard, Y., Morand-Fehr, P., Rouzeau, A., Mandran, N., ` 1987. Composition des principaux tissus adipeux de la chevre˜
al., 1988; Sanudo et al., 1998; Sarti et al., 1993;
Alpine en fin de lactation. Ann. Zootech. 36, 361–374.
Shindarska et al., 1992; Sinnett-Smith et al., 1989;
Bas, P., Gallouin, F., Morand-Fehr, P., 1992. Changes in lipid
Solomon et al., 1992; Solomon et al., 1990; Solomon
content, fatty acid composition and lipoprotein lipase activityet al., 1991; Spillane and L’Estrange, 1977; Stokes
in dry goat omental adipose tissue according to tissue site.Lipids 27, 470–473.
and Walker, 1970; Takahashi and Oota, 1985;
´ ´
Bas, P., Giral, B., Rouzeau, A., 1998. Defauts de qualite des gras
´
Theriez et al., 1976; Tichenor et al., 1970; Vesely,
´ ´
de couverture des agneaux de race lacaune eleves en bergerie.
1973; Vipond et al., 1995; Webb et al., 1994a; Webb
In: Flamant, J.C., Gabina, D., Espejo Diaz, M. (Eds.), Basis of˜et al., 1994b; Webb and Casey, 1995; Wu and Savell,
the Quality of Typical Mediteterranean Animal Products,1992; Ziegler et al., 1967; Zumalacarregui Rodriguez
EAAP Publication No. 90, Wageningen, pp. 451–455. Bas, P., Morand-Fehr, P., Van Quackebeke, E., Cazes, J.P., 1980.and Burgos Gonzalez, 1975; Zygoyiannis et al.,
`
Etude du caractere mou des gras de couverture de certaines
1985.
`eme´
carcasses d’agneaux.. In: 31 Reunion annuelle de la FEZ;. commission de production Ovine et Caprine. Munich. 01–04 / 09 / 1980. 10p.
Bas, P., Rouzeau, A., Poissonnet, P., Sauvant, P., 1997. Influence
References
de la densiteenergetique de la ration sur l’engraissement et la´ ´ ´´ ˆ
composition des depots adipeux chez l’agneau. In: Institut de `
AL-Shabibi, M.M.A., Juma, K.H., 1973. Fatty acid composition of l’Elevage (Eds.), Quatriemes Rencontres autour des Recher-tail, subcutaneous and kidney fats of fat-tailed Awassi sheep. J. ches sur les Ruminants, 04–05 / 12 / 97, p. 324.
Agric. Sci., Camb. 80, 255–257. Bellof, G., Allmendinger, F., Freundenreich, P., 1995. Der Einsatz Anderson, B.A., Friston, G.A., Weihrerauch, J.L., 1977. Lambs von 00-rapssamen in der lammermast (The use of rapeseed in
and veal. Comprehensive evaluation of fatty acids in foods. J. lamb feeding). Wirtscaftseigene Futter 41, 327–339. Am. Diet Assoc. 70, 53–58. Bellof, G., Freundenreich, P., Mayer, P., 1997. Der Einfluss
¨ ¨
Andrieu, J., Demarquilly, C., Sauvant, D., 1989. Tables of feeds fettreicher rapsprodukte auf die fettqualitat von lammerschlak-¨
used in France. In: Jarrige, R. (Ed.), Ruminant Nutrition, orpern (The influence of fatty rapeseed products on fat quality Recommended Allowances and Feed Tables, John Libbey, in lamb carcasses). Fett / Lipid 11, S400–S404.
London, pp. 213–303. Ben Salem, H., Krzeminski, R., Ferlay, A., Doreau, M., 1993.
Ashes, J.R., Siebert, B.D., Gulati, S.K., Cuthbertson, A.Z., Scott, Effect of lipid supply in in vivo digestion in cows: Comparison T.W., 1992. Incorporation of n-3 fatty acids of fish oil into of hay and corn silages diets. Can. J. Anim. 73, 547–557. tissue and serum lipids of ruminants. Lipids 27, 629–631. Beriain, M.J., Bas, P., Purroy, A., Treacher, T., 2000a. Effect of
´
Aurousseau, B., Theriez, M., Daniel, M., 1973. Influence de la nutrition on lamb meat quality. FAO Meeting, Grignon,
` ´ ´ ´ ´ ´
nature des matieres grasses incorporees dans l’aliment d’alaite- France, 03–05 / 09 / 98, Options Mediterranneenes, Serie A,
´ ´ ´ ´ ´
ment sur le metabolisme lipidique de l’agneau de boucherie. Seminaires Mediterranneens, in press.
Ann. Biol. Anim. Bioch. Biophys. 13, 93–105. Beriain, M.J., Horcada, A., Purroy, A., Lizaso, G., Mendizabal, Banskalieva, V., 1997. Effect of age, physiological state and J.A., 2000b. Influence of carcass weight on meat quality of
nutrition on fatty acid composition in depot fat and ruminal Lacha and Rasa Aragonesa lamb. Unpublished data. volatile fatty acids in sheep. Small Ruminant Res. 24, 37–42. Bouillier-Oudot, M., Bozollo, G., Phrem, G., 1992. Optimisation Banskalieva, V., Angelos, A., Shindarska, Z., Marinova, P., 1992. du jugement de couverture des carcasses d’agneaux de
Boylan, W.J., Berger, Y.M., Allen, C.E., 1976. Fatty acid com- on fatty acid content of ovine adipose tissue. J Anim. Sci. 34, 384–387.
position of Finn sheep crossbred lamb carcasses. J Anim. Sci.
Crouse, J.D., Kemp, J.D., Fox, J.D., Ely, D.G., Moody, W.G., 42, 1421–1426.
1972b. Effects of castration on ovine neutral and phospholipid Bozzolo, G., Bouillier-Oudot, M., 1999. Composition en acides
deposition. J. Anim. Sci. 34, 388–392.
´ ´
gras, fermete et coloration du tissu sous-cutane des carcasses
¨ ¨
Demise, S., Matthes, H.D., Mohring, H., Nurnberg, K., Bittner, G., d’agneaux. Ann. Zootech. 48, 47–58.
Pilz, K., Hartung, M., Schubert, M., 1998. Untersuchungen Bozzolo, G., Bouillier-Oudot, M., de Boisseson, E., Ghassan, M.,
zum einfluss der resse, des Geschlechys und der futterung auf Grasset, D., 1990. Influence des performances zootechniques
¨ ¨
die qualitat und fettsaurenzusammensetzung des fleiches von ´
sur les caracteristiques des tissus adipeux d’agneaux de
¨
lammern (Investigations on the effects of breed, sex, and
´ ´ ´ ´ `
bergerie, sevres precocement et alimentes avec un regime a
feeding on lamb carcass composition and quality).
´ ´
forte concentration energetique. Ann. Zootech. 39, 77–94.
¨
Zuchtungskunde 70, 119–140. Brumby, P.E., Storry, J.E., Sutton, J.D., Oldham, J.D., 1979. Fatty
Devier, C.V., Pfander, W.H., 1974. Source and level of dietary fat acid digestion and utilization by dairy cows. Ann. Res. Vet. 10, on fatty acid and cholesterol in lambs. J. Anim. Sci. 38,
317–319. 669–675.
Busboom, J.R., Miller, G.J., Field, R.A., Crouse, J.D., Riley, M.L., Dimov, V., Banskalieva, V., 1989. Effect of the amount of energy Nelms, G.E., Ferrell, C.L., 1981. Characteristics of fat from and protein in diets on the fatty acid composition of lipids in heavy ram and wether lambs. J. Anim. Sci. 52, 83–92. different fat depots in fattening lambs. Zhivotnov’dni Nauki Caballero Garcia de Arevalo, R., Rioperez Garcia del Rinco, J., 26, 90–96.
Fernandez Gonzalez, E., Arauzo Sanchez, M., Hernais Algarra, Doreau, M., Ferlay, A., 1994. Digestion and utilisation of fatty P.J., 1992. Production traits, meats composition and fat quality acids by rumuinants. Anim. Feed Sci. Technol. 45, 379–396. of finishing lamb in diets containing field beans (Vicia faba). Duncan, W.R.H., Garton, G.A., 1967. The fatty acid composition Arch. Zootech. 41, 141–154. and intramolecular structure of triglycerides derived from Cameron, N.D., Bishop, S.C., Speake, B.K., Bracken, J., Noble, different sites in the body of the sheep. J. Sci. Fd Agric. 18,
R.C., 1994. Lipid composition and metabolism of subcutaneous 99–102.
fat in sheep divergently selected for carcass lean content. Duncan, W.R.H., Garton, G.A., 1971. Triglyceride fatty acids of Anim. Prod. 58, 237–242. lambs reared on a lipid-free diet. Proc. Nutr. Soc. 30, 48A– Casey, N.H., van Niekerk, W.A., Spreeth, E.B., 1988. Fatty acid 49A.
composition of subcutaneous fat of sheep grazed on eight Duncan, W.R.H., Garton, G.A., 1978. Differences in the pro-different pastures. Meat Sci. 23, 55–63. portions of branched-chain fatty acid in subcutaneous triacyl-Casey, N.H., Weeb, E.C., 1995. Influence of dietary energy levels glycerols of barley-fed ruminants. Br J. Nutr. 40, 29–33.
and form of diet on composition of fatty acids in subcutaneous Duncan, W.R.H., Ørskov, E.R., Garton, G.A., 1972. Fatty acid adipose tissue of wethers. Small Ruminant Res. 18, 125–132. composition of triglycerides of lambs fed on barley-based diets. Cazes, J.P., Vallade, C., van Quackebeke, E., 1990. Effect of Proc. Nutr. Soc. 31, 19A–20A.
concentrates on quality of subcutaneous adipose tissue of lamb Duncan, W.R.H., Ørskov, E.R., Fraser, C., Garton, G.A., 1974. carcasses. World Rev. Anim. Prod. XXV, 55–62. Effect of processing of dietary barley and of supplementary Ch’ang, T.S., Evans, R., Hood, R.L., 1980. Sire effects on fatty cobalt and cyanocobalamin on the fatty acid composition of acid composition of ovine adipose tissue. J. Anim. Sci. 51, lamb triglycerides, with special reference to branched-chain
1314–1320. components. Br. J. Nutr. 32, 71–75.
Cirruzzi, B., Marsico, G., Vonghia, G., Vicenti, A., Pinto, F., 1993. El-Shahat, A.A., Ashry, M.A., Afify, W.M., 1988. Effect of Comparison of chemical composition of meat and fatty acid slaughter weight on lamb carcass lipid. Ind. J. Anim. Sci. 58, composition of fat deposits in lambs and kids fattened with trub 848–850.
silage and slaughtered at 120 days. Agric. Medit. 123, 106– Enser, M., Hallet, K., Hewitt, B., Fursey, G.A.J., Wood, J.D.,
113. 1996. Fatty acid content and composition of English beef, lamb
Ciruzzi, B., Belliti, E., Muscio, A., Casamassima, D., 1977. and pork at retail. Meat Sci. 42, 443–456.
Variazioni delle carratterische chimiche delle carni e del grasso Enser, M., Hallett, K.G., Hewett, B., Fursez, G.A.J., Wood, J.D., di agnelli svezzati e macellati in epoche diverse. [Variations in Harrington, G., 1998. Fatty acid content and composition of the chemical characteristics of the meat and fat of lambs UK beef and lamb muscle in relation to production system and weaned and slaughtered at different ages]. Ann. Fac. Agr. Bari implications for human nutrition. Meat Sci. 49, 329–341.
´
29, 153–172. Fernandez, C., Gallego, L., Lopez-Bote, C.J., 1998. Effect of
Cramer, D.A., Marchello, J.A., 1964. Seasonal and sex patterns in betaine on fat content in growing lambs. Anim. Feed Sci. fat composition of lambs. J. Anim. Sci. 23, 1002–1010. Technol. 73, 329–338.
Cramer, D.A., Barton, R.A., Shorland, F.B., Czochanska, Z., 1967. Ferruzi, G., Secchiari, P., Pistoia, A., Berni, P., Martini, A., A comparison of the effects of white clover (Trifolium repens) Trimarchi, G., 1985. La prolificita’ delle pecore in funczione and a perennial ryegrass (Lolium perenne) on fat composition della produzione di carne. Nota II: composizione acidica dei and flavour of lamb. J. Agric. Sci., Camb. 69, 367–373. grassi di deposito in agnelli nati da parto gemellare (Appen-Crouse, J.D., Kemp, J.D., Fox, J.D., Ely, D.G., Moody, W.G., ninici e metticci F ed F Romanov2 3 3Appeninina). Atti della
Field, R.A., Rule, D.C., Riley, M.L., 1992. Characteristics of fat the ruminant: In vivo site of fatty acid synthesis in sheep. J. from heavy lambs fed a 90% corn diet or alfalfa pellets. Sheep Nutr. 102, 617–624.
Res. J. 8, 77–80. Jacobs, J.A., Field, R.A., Botkin, M.P., Kaltenbach, C.C., Riley,
Gaili, E.S., Ali, A.E., 1985. Meat from Sudan desert sheep and M.L., 1972a. Effects of testosterone enanthate on lamb carcass goats: Part 2 — composition of the muscular and fatty tissues. composition and quality. J. Anim. Sci. 34, 30–36.
Meat Sci. 13, 229–236. Jacobs, J.A., Field, R.A., Botkin, M.P., Riley, M.L., Roehrkasse, Garrett, W.N., Yang, Y.T., Dunkley, W.L., Smith, L.M., 1976. G.P., 1972b. Effects of weight and castration on lamb carcass
Increasing the polyunsaturated fat content of beef and lamb. J. composition and quality. J. Anim. Sci. 35, 926–930. Anim. Sci. 42, 845–853. Jacobs, J.A., Ray, A., Field, M.P., Botkin, M.P., Riley, M.L., 1973. Garton, G.A., De Hovell, F.D., Duncan, W.R.H., 1972a. Influence Effect of dietary stress on lamb carcass composition and
of dietary volatile fatty acids on the fatty-acid composition of quality. J. Anim. Sci. 36, 507–510.
lamb triglycerides, with special reference to the effect of Jenkins, T.C., Thies, E.J., Fotouhi, N., 1994. Dietary soybean oil propionate on the presence of branched-chain components. Br. changes lipolytic rate and composition of fatty acids in plasma
Jr. Nutr. 28, 409–416. membranes of ovine adipocytes. J. Nutr. 124, 566–570.
Garton, G.A., De Hovell, F.D., Duncan, W.R.H., 1972b. Effect of Johnson, C.B., Purchas, R.W., Birch, E.J., 1988. Fatty acid dietary propionate on the fatty acid composition of lamb composition of fats of differing melting points extracted from triglycerides. Proc. Nutr. Soc. 31, 20A–21A. ram subcutaneous tissue. Lipids 23, 1049–1052.
Geissler, C., Fix, H.P., Hoffmann, M., Peschke, I., 1981. Un- Kemp, J.D., Mahyuddin, M., Ely, D.G., Fox, J.D., Moody, W.G., tersuchungen zum einfluss des alters der tiere und des 1981. Effect of feeding systems, slaughter weight and sex on
¨
futterungsregimes auf quantitative und qualitative merkmale organic properties, and fatty acid composition of lamb. J.
¨ ¨
von intensiv gemasteten lammern. 4. Anteil der einzelnen Anim. Sci. 51, 321–330.
¨ ¨
fettsauren in verschiedenen korperfettproben (Effect of age of Kempster, A.J., 1981. Fat partition and distribution in the carcass the animals and feeding on quantitative and qualitative charac- of cattle, sheep and pigs. Meat Sci. 5, 83–98.
teristics of intensively fattened lambs. 4. Proportions of in- L’Estrange, J.L., Hanrahan, J.P., 1980. Some breed effects on the dividuals fatty acids in different samples of body fat). Arch. melting point and fatty acid composition of carcass fat in
¨
Tierenahrung 31, 849–859. lambs. J. Agric. Sci., Camb. 95, 73–76.
Gibney, M.J., L’Estrange, J.L., 1975. Effects of dietary unsatu- L’Estrange, J.L., Mulvihill, T.A., 1975. A survey of fat charac-rated fat and protein source on melting point and fatty acid teristics of lamb with particular reference to the soft fat composition of lamb fat. J. Agric. Sci. 84, 291–296. condition in intensively fed lambs. J. Agic. Sci., Camb. 84, Hansen, R.P., Czochanska, Z., 1978. Fatty acid composition of the 281–290.
subcutaneous fats of lambs fed 71% barley grain, and a L’Estrange, J.L., 1977. The performance and carcass fat charac-comparison with those of lambs grazed on pasture, or fed teristics of lambs fattened indoors on concentrate diets. 3. higher levels of barley grain. N.Z. J. Sci. 21, 85–90. Effects of whole versus ground barley or roughage supple-Hansen, R.P., Czochanska, Z., 1976. Fatty acid composition of the mentation with chopped hay or straw. Ir. J. Agric. Res. 16,
subcutaneous and perinephric fats of lambs grazed on pasture 221–232.
in New Zealand. N.Z. J. Sci. 19, 413–419. L’Estrange, J.L., 1979. The performance and carcass fat charac-Haugebak, C.D., Hedrick, H.B., Asplund, J.M., 1974. Adipose teristics of lambs fattened on concentrate diets. 4. Effects of tissue accumulation and cellularity in growing and fattening barley fed whole, ground or pelleted and of a high level of zinc lambs. J. Anim. Sci. 39, 1016–1025. supplementation. Ir. J. Agric. Res. 18, 173–182.
Hawke, J.C., Morrison, I.M., Wood, P.R., 1977. Structures of Leat, W.M.F., Kemp, P., Lysons, R.J., Alexander, T.J.L., 1997. triacylglyceriols in lamb fat with elevated levels of linoleic Fatty acid composition of depots fats from gnotobiotic lambs. acid. J. Sci. Fd. Agric. 28, 293–300. J. Agric Sci. 88, 175–179.
Horcada, A., Beriain, M.J., Purroy, A., Lizaso, G., Chasco, J., Leth, T., Ovesen, L., Hansen, K., 1998. Fatty acid composition of 1998. Effect of sex on meat quality of Spanish lamb breeds meat ruminants, with special emphasis on trans fatty acids. J. (Lacha and Rasa Aragonesa). Anim. Sci. 67, 541–547. Am. Oil Chem. Soc. 75, 1001–1005.
Ifkovits, R.W., Ragheb, H.S., 1968. Cellular fatty acid composition Lough, D.S., Solomon, M.B., Rumsey, T.S., Elsasser, T.H., Slyter, and identification of rumen bacteria. Appl. Microbiol. 16, L.L., Kahl, S., Lynch, G.P., 1992. Effects of dietary canola
1406–1413. seed and soy lecithin in high-forage diets on cholesterol content
Illian, M.A., Razzaque, M.A., Al-Awadi, A., Salman, A.J., 1988. and fatty acid composition of carcass tissues of growing ram Use of fat in diets of sheep in hot environments. I. Effects on lambs. J. Anim. Sci. 70, 1153–1158.
performance, carcass characteristics and lipid composition of Manfredini, M., Cavani, C., 1979. Utilizzazione zootecnica degli plasma. Anim. Feed Sci. Technol. 19, 327–341. effuenti di distilleria: la borlanda di mellaso di bientola Ingle, D.L., Bauman, D.E., Garrigus, U.S., 1972a. Lipogenesis in concentrata (BMBC) nell’alimentazione degli agnelli. Zoot.
the ruminant: In vitro study of tissue sites,carbon source and Nutr. Anim. 5, 147–153.
reducing equivalent generation for fatty acid synthesis. J. Nutr. Marinova, P., Shindarska, Z., Banskalieva, V., 1992. Deposition
102, 609–616. and composition of muscular tissue in lambs under different
´ ´ ´ ´ ´ ´ ´ ´ Internatinal Congress of Meat Science and Technology, Cler- Mediterranneenes, Serie A, Seminaires Mediterranneens, in mont-Ferrand, France, 23–23:08 / 92, Vol. II, pp. 89–92. press.
¨
Marinucci, M.T., Ippedico, V., Martino, G., Polidori, P., Loschi, R., Nurnberg, K., Grumbach, S., Papstein, H.J., Matthes, H.D., Ender, ¨
Severini, M., 1998. Use of agro-industrial by-products for K., Nurnberg, G., 1996. Fettzusammensetzung von fattening lambs: pasta factory residues. Influence on meat Lammfleisch (Fat composition of lamb). Fett / Lipid 2, S77–80.
¨
production and meat quality. J. Anim. Feed Sci. 7, 283–292. Nurnberg, K., Wegner, J., Ender, K., 1998. Factors influencing fat ¨
Matthes, H.D., Hillmann, D., Demise, S., Mohring, H., 1998. composition in muscle and adipose tissue of farm animals. Fleischqualitatsmerkmale von schlaflammern unterschiedlicher Livest. Prod. Sci. 56, 145–156.
genotypen aus okologisch orientierter haltung (Meat quality Ørskov, E.R., Duncan, W.R.H., Carnie, C.A., 1975. Cereal Pro-traits of lambs of different genotypes under ecologically cessing and food utilisation by sheep. 3. The effect of replacing orientated management). Zuchtungskunde 70, 282–297. whole barley by whole oats on food utilization and firmness
¨ ¨
Matthes, H.D., Nurnberg, K., Mohring, H., Demise, S., Bittner, G., and composition of subcutaneous fat. Anim. Prod. 21, 51–58. ¨
Pilz, K., 1996. Fettsaurenzusammensetzung von Lammfleisch Ørskov, E.R., Fraser, C., Gordon, J.G., 1974. Effect of processing
¨ ¨
in abhangigkeit von der futterung (Dependence of the fatty acid of cereals on rumen fermentation, digestibility, rumination composition of lambs’ meat on feeding). Fleischwirtschaft 76, time, and firmness of subcutaneous fat in lambs. Br. J. Nutr.
907–909. 32, 59–69.
Melton, S.L., 1990. Effects of feeds on flavor of red meat: a Payne, E., Masters, C.J., 1971. The in vitro incorporation of
14
review. J. Anim. Sci. 68, 4421–4435. C-labelled fatty acids into ovine liver and omental adipose Miller, G.J., Kunsman, J.E., Field, R.A., 1980. Characteristics of tissue. Int. J. Bioch. 2, 623–643.
soft subcutaneous fat in ram lambs fed corn and corn-silage Pearce, J., Chesnutt, M.B., Todd, J.R., 1974. A comparison of the diets. J. Food Sci. 45, 279–285. fatty acid composition of adipose tissue triglyceride from Miller, G.J., Rice, R.W., 1967a. Lipid metabolism in lambs as grass-fed and intensively-reared lambs. Proc. Nutr. Soc. 33,
affected by fattening rations of roughage and concentrate. J. 99A–100A.
Anim. Sci. 26, 1153–1159. Petit, H.V., Rioux, R., D’Oliveira, P.S., do Prado, I.N., 1997. Miller, G.J., Varnell, T.R., Rice, R.W., 1967b. Fatty acid com- Performance of growing lambs fed grass silage with raw or position of certain ovine tissues as affected by maintenance extruded soybean or canola seeds. Can. J. Anim. Sci. 77, level ration of roughage and concentrate. J. Anim. Sci. 26, 455–463.
41–45. Purchas, R.W., O’Brien, L.E., Pendleton, C.M., 1979. Some
´ ´ ´
Molenat, G., Theriez, M., 1973. Influence du mode d’elevage sur effects of nutrition and castration on meat production from ´
la qualite de carcasse de l’agneau de bergerie. Etude des effets male Suffolk cross (Border Leicester–Romney cross) lambs.
´ ` ´
de l’allaitement artificiel et des regimes entierement condenses N.Z. Agric. Res. 22, 375–383.
´ ´ ˆ
sur la qualite des depots adipeux de couverture. Ann. Zootech. Quackebeke, E. van, Cazes, J.P., Vallade, C., 1978. Alimentation et
´ `
22, 279–293. qualite des gras. In: L’Alimentation de la brebis et de la chevre.
`emes
´
Murphy, J., L’Estrange, J.L., 1977. The performance and carcass In: INRA-ITOVIC (Eds.), 4 Journees de la Recherche fat characteristics of lambs fattened on concentrate diets. 1. Ovine et Caprine, pp. 295–311.
Effects of maize and barley as the cereal source and dietary Ray, E.E., Kromann, R.P., Cosma, E.J., 1975. Relationships supplementation with roughage, vitamine E, cobalt and vitamin between fatty acid composition of lamb fat and dietary B12. Ir. J. Agric. Res. 16, 187–204. ingredients. J. Anim. Sci. 41, 1767–1774.
Muscio, A., Marsico, G., Celi, R., Casamassima, D., Maiorana, Robelin, J., 1986. Composition corporelle des bovins: Evolution
´ ´
M., 1978. L’utilizazzazione dei gemelli per la produzione au cours du developpement et differences entre races. In: `
dell’agnello pesanto bianco. Qualita delle carni e del grasso di Thesis Dr es Sciences, Univ. Clermont-Ferrand II.
deposito (Use of twins for the production of heavy lambs with Rule, D.C., Koch, D.M., Jones, R.R., Kercher, C.J., 1991. Brassica white meat. Quality of the meat and fat deposits). Ann. Fac. and sugarbeet forages for lambs — growth performance of
Agr. Bari 30, 865–883. lambs and composition of forage and dock-fat fatty acids. J.
Muscio, A., Zezza, L., Marsico, G., Nicastro, F., 1980. Com- Prod. Agric. 4, 29–33.
posizione chimico-bromatologica delle carni di agnelli di 100 Safari, E., Sefidbakht, N., Farid, A., 1988. Effect of castration and giorni alimentati con crescenti di fieno (Chemical and crytorchidism on fatty acid content of ovine adipose tissue. bromatological composition of meat from lambs 100 days old Meat Sci. 23, 65–69.
˜
fed on increasing proportions of hay). Ann. Fac. Agr. Bari 31, Sanudo, C., Sierra, I., Olleta, J.L., Martin, L., Campo, M.M.,
571–590. Santolaria, P., Wood, J.D., Nute, G.R., 1998. Influence of
Nocek, J.E., Tamminga, S., 1991. Site of digestion of starch in the weaning on carcass quality, fatty acid composition and meat gastrointestinal tract of dairy cows and its effect on milk yield quality in intensive lamb production systems. Anim. Sci. 66, and composition. J. Dairy Sci. 74, 3598–3629. 175–187.
´
Normand, J., Bas, P., Aurousseau, B., Theriez, M., Sauvant, D., Sarti, D.M., Morbidini, L., Panella, F., Sarti, F.M., Binaglia, L., 2000. Effect of diet and sire origin on the fatty acid com- Iacurto, M., 1993. Carattecistiche chimiche della carne di
`
´
lamb meat at different slaughtering ages and nutritive level of Vezinhet, A., Noughes, J., Teissier, J., 1983. Influence d’un mode
´ ´
the diet). Zoot. Nutr. Anim. 19, 103–112. d’elevage sur la mise en reserve des graisses par le tissu
SAS, 1, 1987. Statistical Analysis System. SAS / STAT SAS adipeux chez les ovins. Reprod. Nutr. Develop. 23, 837–846. User’s Guide: Statistics, 6th Edition, SAS, Cary, NC. Vipond, J.E., Lewis, M., Horgan, G., Noble, R.C., 1995. Malt Shindarska, Z., Banskalieva, V., Marinova, P., Dardjonov, T., distillers grains as a component of diets for ewes and lambs 1992. Changes in both quality and quality and composition of and its effects on carcass tissue lipid composition. Anim. Feed meat in castrated and noncastrated hoggets fed clenbuterol. In: Sci. Technol. 54, 65–79.
Proc. 38th Internatinal Congress of Meat Science and Technol- Vimini, R.J., Field, R.A., Crouse, J.D., Miller, G.J., 1984. Factors ogy, Clermont-Ferrand, France, 23–23:08 / 92, Vol. II, pp. 137– affecting melting point of subcutaneous fat from heavy ram and
140. weather lambs. Int. Goat Sheep Res. 2, 105–113.
Sinnett-Smith, P.A., Woolliams, J.A., Warriss, P.D., Enser, M., Viviani, R., 1970. Metabolism of long-chain fatty acids in the 1989. Effects of recombinant DNA-derivated bovine somatot- rumen. Adv. Lipid Res. 8, 267–346.
ropin on growth, carcass characteristics and meat quality in Walker, D.M., Stokes, B.B., 1970. The nutritive value of fat in the lambs from three breeds. Anim. Prod. 49, 281–289. diet of the milk-fed lamb. 1. The apparent and corrected Solomon, M.B., Lynch, G.P., Lough, D.S., 1992. Influence of digestibilities of different dietary fats and their constituent fatty
dietary palm oil supplementation on serum lipid metabolites, acids. Br. J. Nutr. 24, 425–433.
carcass characteristics, and lipid composition of carcass tissues Whale, K.W.J., Garton, G.A., 1972. Desaturation of stearic acid by of growing ram and ewe lambs. J. Anim. Sci. 70, 2746–2751. sheep tissue microsomes. Proc. Nutr. Soc. 31, 110A–111A. Solomon, M.B., Lynch, G.P., Ono, K., Paroczay, E., 1990. Lipid Webb, E.C., Bosman, M.J.C., Casey, N.H., 1994a. Dietary
in-composition of muscle and adipose tissue from crossbreed ram, fluences on subcutaneous fatty acid profiles and sensory wether and cryptorchid lambs. J. Anim. Sci. 68, 137–142. characteristics of Dorper and SA Mutton Merino wethers. S. Solomon, M.B., Lynch, G.P., Paroczay, E., Nort
