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Departamento de Producción Animal and Departamento de Bioquímica y Biología Molecular, Facultad de Veterinaria, Universidad Complutense, 28040 Madrid, Spain
1To whom correspondence should be addressed.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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0.03) in chickens fed the sunflower
oil–enriched diets, indicating a greater rate of ß-oxidation. Liver
fatty acid synthetase activity was lower (P = 0.01)
in chickens fed the sunflower oil–enriched diet, suggesting reduced
hepatic lipogenesis in this group. Postprandial plasma triglyceride
levels were significantly lower (P < 0.05) in
birds fed the sunflower oil–enriched diet, indicating a higher rate of
dietary lipid clearance from the bloodstream to tissues. In conclusion,
the lower fat deposition observed in broilers fed sunflower
oil–enriched diets appears to be the net result of an increased rate
of lipid catabolism and lower rate of fatty acid synthesis despite
higher dietary fat absorption.
KEY WORDS: • dietary fat type • lipogenesis • lipolysis • fat deposition • broiler chickens
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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) or total body fat (Sanz et al. 2000
) deposition than do broiler chickens fed diets containing
saturated fatty acids. This is of particular interest in the rearing of
female broilers, because the amount of abdominal fat at slaughter age
(49 d) may represent >3% of the live weight (Sanz et al. 1999
). In general, body fat accumulation may be considered the net result of the balance among dietary absorbed fat, endogenous fat synthesis (lipogenesis) and fat catabolism via ß-oxidation (lipolysis). Thus, if the amount of absorbed fat is the same, lower body fat deposition may be attributed to increased fat catabolism or diminished endogenous fatty acid synthesis or to both processes.
Enhanced fat catabolism and reduced fatty acid synthesis were reported
to occur in rats fed unsaturated fatty acid–enriched diets compared
with rats fed diets enriched with saturated fatty acids.
Shimomura et al. (1990)
reported lower fat accumulation
in rats fed a diet high in sunflower oil
(SUN)2than in rats fed a beef tallow (TAL)-enriched diet. Their findings
indicated that the higher degree of fat ß-oxidation in rats fed the
SUN diet compared with those fed the TAL diet was in part responsible
for the different pattern of fat deposition. Also, fatty acid synthesis
may be reduced in rats fed unsaturated fatty acids. Clarke et al. (1977)
reported that 3% methyl linoleate in the diet
decreased fatty acid synthesis in the liver of rats by 50%. However,
8% methyl stearate, which was absorbed to the same extent as the
linoleate, had no effect on fatty acid synthesis.
As far as we are aware, the effects of dietary fat type on fat metabolism and its consequences on fat deposition in broiler chickens remain unclear. Thus, the aim of this experiment was to test the effect of two sources of dietary fat, SUN (as a source of polyunsaturated fat) and TAL (as a source of saturated fat), on lipogenesis, lipolysis and fat deposition in broiler chickens.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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Twenty-four 21-d-old Hybro-G female broiler chickens (Nutreco PRC,
Toledo, Spain) were placed in individual metabolic cages and, based on
live weight, assigned to one of two experimental treatment groups. The
experimental design included two
diets3
that differed only in the source of additional fat (added at 8 g/100
g): TAL or SUN. Diet metabolizable energy was measured by total excreta
collection according the European reference method (Bourdillon et al. 1990
) between d 35 and 40 of life. Correction for
zero-nitrogen retention was made on the basis of the difference
between nitrogen intake and nitrogen excretion, assuming that the
energy equivalent was 34.36 kJ/gN. Feed and water were consumed
ad libitum. A light/dark cycle of 23:1 h and a room temperature of
22–26°C were maintained throughout the 32-d experimental period.
Chickens were handled according to the principles for the care of
animals in experimentation (National Research Council 1985
).
To obtain preprandial and postprandial blood samples at 53 d of age, an additional group of 21-d-old-chicks (n = 32) randomly assigned to each experimental diet were established and reared as described previously.
Controls and sampling.
The final weight, feed intake and weight gain of the broiler chickens
were recorded at the end of the experimental period. The 53-d-old
broiler chickens were stunned, slaughtered, bled and plucked in a local
slaughterhouse. The heart and liver were immediately removed from hot
carcasses, packed in plastic bags and stored in liquid nitrogen until
the time of analysis. After removal of the heart and liver, carcasses
were chilled to 4°C, and the abdominal fat pad (from the
proventriculus surrounding the gizzard down to the cloaca) was removed
and weighed (Cahaner et al. 1986
).
To obtain preprandial and postprandial blood samples, an additional group of 32 chickens were deprived of food for 6 h. After food deprivation, blood samples (preprandial) were taken from eight randomly selected birds per experimental treatment. The remainder of the chickens (eight per treatment) were then refed, and blood samples were taken from these chickens 2.5 h later (postprandial). Blood was extracted via wing puncture into heparinized tubes and immediately centrifuged. Plasma samples were stored at -20°C until required.
Analysis of diets.
Samples of each diet were analyzed for nitrogen content (Kjedahl
method; Association of Official Analytical Chemists, 1990
), crude protein (N x 6.25), dry matter (dried in an
oven at 103°C for 4 h) and lipid content (6-h Soxhlet extract).
The fatty acid content of the feeds was determined by extracting the
fat in chloroform/methanol (Bligh and Dyer 1959
).
Previously methylated fatty acid samples were identified by gas
chromatography as described elsewhere (Lopez-Bote et al. 1997
).
Enzyme assays.
Crude heart and liver extracts were used to determine the activity of
the enzymes under study. Mitochondria were isolated from defrosted
hearts for carnitine palmitoyltransferase I (CPT I; EC 2.3.1.21) and
L-3-hydroxyacyl-CoA dehydrogenase (L3HOAD; EC 1.1.135)
activity determinations as previously described by Saggerson (1982)
. CPT I was assayed directly using whole mitochondrial
isolates. For L3HOAD determination, mitochondrial pellets were
resuspended in 0.3 mol sucrose containing 10 mmol Tris-HCl (pH 7.4
at 0°C) and 1 mmol EGTA per L and sonicated to obtain a mitochondrial
supernatant after centrifugation at 20,000 x g for
40 min at 4°C. FAS (EC 2.3.1.38) was determined in liver homogenates
prepared as previously described by Dias et al. (1998).
)
CPT I was assayed via modification of a previously described isotopic
procedure using radiolabeled carnitine and unlabeled fatty acyl-CoA
(Singh and Poulos 1995
). Briefly, mitochondrial extracts
(
5 µg of protein) were preincubated for 4 min at 30°C with 50
mmol sodium phosphate buffer, pH 7, containing 1 mmol dithiothreitol,
5 g fatty acid–free bovine serum albumin and 90 µmol
palmitoyl-CoA per L. The reaction was initiated by the addition of
250 µmol of
L-[methyl-3H]carnitine/L
(1.48 Bq/mol; Amersham, Barcelona, Spain) and terminated after 20 min
by the addition of 1:1 chloroform/methanol. The radiolabeled product of
CPT activity was then extracted and quantified. Two control assays were
conducted in parallel: one contained no mitochondrial extract and a
second one inhibited the activity of CPT I with 0.2 mmol of
malonyl-CoA/L (Derrick and Ramsay 1989
). The activity of
L3HOAD was measured according to the spectrophotometric method proposed
by Bradshaw and Noyes (1975).
FAS activity was
determined according to the isotopic method proposed by Hsu et al. (1969).
Soluble protein content was measured according to
the method of Bradford (1976)
, using bovine serum
albumin as standard (chemicals from Sigma, Alcobendas, Spain).
Plasma triglycerides.
Plasma triglycerides were enzymatically determined (Bucolo and David 1973) with a commercial kit (Sigma Diagnostics) in which triglycerides are hydrolyzed by lipase to glycerol, which is then subjected to coupled enzymatic reactions to yield a molecule with a maximum absorbance at 500 nm.
Statistical analysis.
The SAS General Linear Models (GLM) procedure was used for the
statistical analysis of data (SAS Institute, 1988
).
Plasma triglyceride data were analyzed by considering a 2 x 2
factorial design, with two levels of dietary fat (TAL and SUN) and two
times (preprandial and postprandial). Because the interaction between
the effects (fat type x time) was significant (P
= 0.01), data were reanalyzed with the Duncan test to separate
means (
= 0.05).
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RESULTS AND DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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). As expected, the metabolizable energy of the TAL-enriched diet
tended to be lower than that of the SUN-enriched diet (P
= 0.09). Because the experimental diets were identical except in
the fat source, observed differences in dietary apparent metabolizable
energy corrected for zero-nitrogen retention are attributable to
the different absorptions of the dietary fats (Wiseman et al. 1991
).
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).
|
. Similar findings
have been reported for the comparison of abdominal fat deposition in
birds fed diets containing unsaturated or saturated fatty acids
(Sanz et al. 1999
). In addition, Sanz et al. (2000)
demonstrated lower total body fat and lower total body
fat gain in chickens fed diets rich in SUN compared with those fed
TAL-enriched diets.
As mentioned, body fat deposition depends on the net balance among
absorbed fat, endogenous fat synthesis and fat catabolism. Because
abdominal fat is positively correlated with total body fat
(Becker et al. 1979
) and given that the amount of
absorbed fat was higher in birds fed the SUN-enriched diet,
observed lower body fat deposition should be accounted for by a higher
rate of fat oxidation, lower rate of FAS or both.
To explore whether different dietary fat sources may result in
different fat metabolism, leading to changes in fat deposition, the
activities of three enzymes involved in fat metabolism were determined.
Mitochondrial CPT I exchanges CoA for carnitine to facilitate the
transfer of activated fatty acids inside the mitochondria for
ß-oxidation. L3HOAD participates in the oxidation phases of fatty
acid ß-oxidation. Heart and skeletal muscle CPT I have been reported
to be identical (Weis et al. 1994
). Because
mitochondrial oxidation rate is mainly dependent on the actual capacity
of the carnitine transport system (Brouns and Vusse 1998
), the activities of both CPT I and L3HOAD in heart extract
were used as indicators of muscle fatty acid ß-oxidation. In
contrast, FAS is a multienzyme complex responsible for elongating
chains of newly synthesized fatty acids via successive acetyl-CoA
linkage. Because the liver is considered the primary site of FAS in
birds (Saadoun and Leclercq 1983
, Yeh and Leveille 1973
), the activity of this enzyme was estimated in
liver tissue as an indicator of FAS.
The specific activities of CPT I and L3HOAD were both significantly
higher in heart extracts of broiler chickens fed SUN diets than in
those fed high TAL diets, suggesting a higher rate of ß-oxidation in
the former (Table 3
). The enhanced CPT I and L3HOAD activity observed in chickens fed the
SUN-enriched diet is in agreement with the findings of
Leyton et al. (1987)
, who reported a greater rate of
ß-oxidation of unsaturated compared with saturated fatty acids.
Similarly, Takeuchi et al. (1996)
reported significantly
decreased CPT activity in the brown adipose tissue of rats fed
lard-enriched diets versus rats fed high oleic SUN-, SUN- or
linseed oil–enriched diets. In addition, Powers and Newsholme (1997)
found higher CPT I activity in heart mitochondrial
preparations of rats fed diets enriched with SUN, evening primrose oil
(both are rich in linoleic acid) and menhaden oil [rich in long chain
(n-3) fatty acids] compared with that of rats fed low fat diets or
diets enriched with olive oil or hydrogenated coconut oil. These
authors suggest that high dietary linoleic acid concentration (an
essential fatty acid) may induce an increase in the rate of
ß-oxidation (and thus higher CPT I activity) due to the consequent
reduced risk of loss of linoleic acid through catabolism, leading to
essential fatty acid deficiency.
|
. Tanaka et al. (1983)
showed that increasing dietary energy through the
addition of soybean oil led to a higher concentration of nonesterified
fatty acids in plasma and reduced citrate lyase activity (cytosol
enzyme involved in the generation of acetyl-CoA from citrate) with
a concomitant decrease in FAS.
Plasma triglyceride concentrations were significantly greater after the
ingestion of a meal (Fig. 1
), and postprandial values were significantly lower in broilers fed the
SUN diet (Fig. 1)
. Given the higher fat absorption in chickens fed the
SUN diet, due to the greater digestibility of SUN, this finding may
suggest a relatively higher postprandial rate of fat uptake from the
bloodstream to the tissues in chickens fed the SUN diet. Possibly
consistent with the present findings, Lai and Ney (1995)
observed that rats fed diets high in saturated fatty acids (palm oil
and butterfat) compared with those fed diets high in corn oil showed
significantly higher postprandial plasma triacylglycerol
concentrations. In their experiment, postheparin plasma lipoprotein
lipase activity was greater after the intake of corn oil than after the
intake of saturated fatty acids. These authors suggested that the
greater postprandial lipemia observed in saturated fatty acid–fed rats
was probably in part due to slower plasma triacylglycerol clearance.
Shimomura et al. (1990)
also reported higher plasma
triacylglycerol concentrations in rats fed a TAL-enriched diet
compared with rats fed an SUN-enriched diet. In this experiment,
the heart and soleus muscle of rats fed the SUN-enriched diet
showed higher LPL activity after feeding, suggesting that blood
triacylglycerol muscle uptake occurred at a greater rate in the
SUN-fed rats than those fed TAL. In contrast, these authors found
no difference in adipose tissue lipoprotein lipase activity.
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In conclusion, the lower abdominal fat deposition occurring in broiler chickens fed a SUN-enriched diet compared with those fed a TAL-enriched diet may be explained by an increased rate of ß-oxidation of unsaturated dietary fats and reduced endogenous fatty acid synthesis despite higher dietary fat absorption.
The lower plasma triglyceride concentration after meal ingestion
recorded in chicks fed the SUN diet may be explained by a higher rate
of uptake of dietary polyunsaturated lipid (Lai and Ney 1995
, Shimomura et al. 1990
), leading to greater
substrate availability and thus faster ß-oxidation of polyunsaturated
fatty acids (Leyton et al. 1987
).
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FOOTNOTES |
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3 Experimental diets contained (per kg of diet):
106 g wheat, 448 g corn, 298 g soybean meal, 30 g
full fat soybean, 80 g added fat (tallow or sunflower oil, 3 g sodium chloride, 6.2 g calcium carbonate, 22 g bicalcium
phosphate, 22 g dl-methionine and 5 g vitamin
and mineral premix. The vitamin and mineral premix provided (per kg of
diet) 2.58 mg retinal, 37.5 mg cholecalciferol, 7.52 mg
dl--tocopheryl acetate, 5.28 mg riboflavin, 8 mg
pantothenic acid, 1.84 mg pyridoxine (PN), 0.5 mg folic acid, 12.5 mg
cobalamin, 350 mg choline, 0.15 mg Se, 1.9 mg I, 0.2 mg Co, 6 mg Cu,
30.8 mg Fe, 50 mg Zn, 80 mg Mn and 232 mg S. Analyzed average nutrient
composition was (per kg of diet) 193 g crude protein and 104 g crude fat.
Manuscript received May 15, 2000. Revision accepted August 22, 2000.
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