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,3
*
Agriculture Victoria, Victorian Institute of Animal Science, Werribee, VIC 3030, Australia;
Swinburne University of Technology, Hawthorn, VIC 3122, Australia;
**
Charles Sturt University, Wagga Wagga, NSW 2650, Australia; and
Cornell University, Ithaca, NY 14853
4To whom correspondence should be addressed.
 |
ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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KEY WORDS: • pigs • conjugated linoleic acid • body composition • lipid deposition • growth
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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); since that time, the anticarcinogenic
activity of CLA has been demonstrated in a wide range of animal models
(Banni and Martin 1998
, Belury 1995
). The
interest in CLA has grown because it possesses several additional
biological properties that relate to health. Dietary CLA
supplementation reduced the catabolic effects of immune stimulation in
mice, rats and chickens without adversely affecting immune function
(Miller et al. 1994
). Furthermore, CLA is
antiatherogenic in hamsters (Nicolosi et al. 1997
) and
antidiabetic in the Zucker diabetic fatty acid
fa/fa rat (Houseknecht et al. 1998
).
One of the biological effects of CLA relates to fat accretion and
nutrient partitioning. CLA has been shown to increase live weight gain,
to improve feed efficiency in rats, mice and chickens (Chin et al. 1994
, Park et al. 1997
) and to decrease
carcass fat content in mice (West et al. 1998
). Recent
results suggest that similar effects occur in pigs because dietary CLA
supplementation reduced back fat thickness (Thiel et al. 1998
) and the fat content of commercial meat cuts (Dugan et al. 1997
). However, effects of CLA on carcass composition
and tissue deposition rates in pigs have not been reported. Therefore,
our objective was to examine the effect of dietary supplementation of
CLA on carcass composition and rates of accretion of fat, protein,
water and ash. We used a commercial source of CLA and supplemented the
diet of growing pigs with a range of doses up to 5.5 mg CLA isomers/kg
of diet.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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All procedures involving animals were approved by the Victorian
Institute of Animal Science Animal Ethics Committee. Female
cross-bred (Large White x Landrace) pigs (n
= 66; initial weight, 56.6 kg) were used. To obtain initial body
composition, six pigs were slaughtered at 57 kg live weight. The
remaining 60 pigs were randomly allocated to one of six dietary
treatments [0, 1.25, 2.5, 5.0, 7.5 and 10.0 mg of CLA-55/kg diet
(Natural Lipids, Hovdebygda, Norway)]. This CLA, containing 55% CLA
isomers, was prepared from sunflower oil. Thus, the CLA concentrations
for the six diets were 0, 0.7, 1.4, 2.75, 4.1 and 5.5 g/kg diet,
respectively. To ensure a homogenous mixture, CLA was mixed into
soybean oil before being added to the basal diet. Diets were mixed
every 2 wk, and the remaining CLA was stored at 4°C to minimize
oxidation. The diets (Table 1
) were formulated to be in excess of protein and lysine requirements for
this class of pigs (Dunshea et al. 1993a
). In addition,
the experimental diets were formulated to contain amino acids relative
to lysine in excess of the amino acid balance proposed as ideal by the
Standing Committee on Agriculture (1987)
. Fresh diet was
provided daily to each pig and the amount was recorded.
|
Pigs were fed their diets for 8 wk before being transported 0.5 km to a
pilot abattoir. After CO2 stunning, pigs were
exsanguinated, hair removed and bodies eviscerated. Internal organs
were weighed and the gastrointestinal tract emptied and reweighed.
Internal organs and empty gastrointestinal tract were then frozen at
-20°C. The eviscerated carcasses were hung overnight at 4°C before
being split in half. The right side of each carcass and the empty
viscera were prepared for separate proximate analyses (Campbell et al. 1985
). Dry matter was determined in triplicate by drying
samples to constant weight in a force-draft oven at 105°C. Ash
was determined by burning the oven-dried samples in a muffle
furnace at 600°C. Protein was determined by Kjeldahl analysis
(AOAC 1984
), and fat was determined after
chloroform:methanol extraction (Folch et al. 1957
). The
left side of the carcass was used for determination of meat quality and
carcass measurements (to be presented elsewhere).
Analyses of CLA isomers.
Fatty acid methyl esters were prepared by adding 100
µL of CLA-55 to 1 mL of 20% methanol benzene and
reacted with trimethylsilyldiazomethane (Hashimoto et al. 1981
). Samples were then analyzed by HPLC (Sehat et al. 1998
).
Statistical analyses.
Data were analyzed by an ANOVA suitable for a dose response, with
linear and quadratic effects determined. The model included block,
replicate and CLA dose. In addition, comparisons were made between
diets containing either none or added CLA. For these analyses, the
model included block, replicate and CLA dose. All analyses were
performed using GENSTAT (Payne et al. 1993
).
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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. The major CLA isomers were cis-11 trans-13
(18%), cis-10 trans-12 (30%), cis-9
trans-11 (25%) and cis-8 trans-10
(14%). There were traces of seven other isomers.
|
). However, the gain to feed ratio was increased by dietary CLA, with
the responses being most pronounced over the first 4 wk of the
treatment. During the first 4 wk of the study, there was a linear
(0.00165 ± 0.00087 g/g for each gram per kilogram increase in CLA
concentration; P = 0.04) increase in the gain to feed
ratio with increasing dose of CLA, whereas there was no effect of CLA
over the period between 4 and 8 wk. Overall, including CLA in the diet
increased the gain to feed ratio by 6.3%. Dietary CLA had no effect on
live weight at any stage of the study. On the other hand, there was a
dose-dependent decrease in P2 back fat
thickness that became evident after 3 wk of feeding diets containing
CLA (Fig. 1
).
|
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). Although there was a significant linear increase (P = 0.02) with increasing CLA, there was no overall significant
(P = 0.14) difference in carcass protein content among
the different dietary CLA supplements. Carcass lean tissue (sum of
water and protein) increased linearly (P < 0.001) with
increasing dietary CLA supplementation, with a commensurate decrease
(P < 0.001) in carcass fat content. At the highest
level of CLA inclusion, carcass fat content was decreased by ~60 g/kg
(-20%). As a consequence, the ratio of fat to lean in the carcass was
decreased in pigs fed all but the lowest level of supplemental CLA
(Fig. 2
). The ratio of fat to protein in the carcass decreased linearly
(-0.093 ± 0.0216 g/g for each gram per kilogram increase in CLA
concentration; P < 0.001) with increasing dietary CLA.
Although there were no significant differences among the different
dietary CLA groups in the ratio of water to protein in the carcass when
data from all CLA groups were pooled, there was a significant increase
(+5%, P = 0.03) in the ratio of water to protein in
the carcass. There was no significant effect of dietary CLA
supplementation on carcass ash content.
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, P < 0.002). The carcass protein deposition response
to dietary CLA was also quadratic (P = 0.01) in nature,
with protein deposition also maximized at a dietary CLA supplementation
of 5.0 g/kg. The carcass lean tissue deposition response to dietary CLA
was also quadratic (P = 0.001) in nature with lean
tissue deposition maximized at a dietary CLA supplementation of 5.0
g/kg. Carcass fat deposition decreased linearly with increasing CLA
supplementation rates. As a consequence, the ratio of fat to lean
deposition in the carcass was decreased in pigs fed all but the lowest
level of supplemental CLA (Fig. 2)
. The ratio of fat to protein
deposition in the carcass decreased linearly (P < 0.001) with increasing dietary supplementation of CLA. Although there
were no significant differences among the different dietary CLA groups
in the ratio of water to protein deposition in the carcass when data
from all CLA groups were pooled, there was a significant increase
(+11%, P = 0.024) in the ratio of water to protein
deposited in the carcass. There was no significant effect of dietary
CLA on carcass ash deposition.
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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, Thiel et al. 1998
), although some failed to observe any improvement
(O'Quinn et al. 1998
). The earlier work demonstrated
that the gain to feed ratio was increased by 5% in rats fed CLA
supplement of up to 5 g/kg of diet (Chin et al. 1994
).
Our study is the first to demonstrate the effect of CLA supplements on
carcass composition and tissue deposition in growing pigs. Proximate
analysis indicated that CLA reduced carcass fat in a linear manner over
the CLA dose range (Table 4)
. At the highest level of CLA
supplementation (10 g/kg diet), carcass content of fat was reduced by
~61 g/kg and carcass fat deposition was reduced by 31% (86 g/d)
(Table 5)
. Earlier studies had indicated that dietary CLA
supplementation of growing pigs resulted in less fat at slaughter as
estimated by dissection of wholesale loin cuts (Dugan et al. 1997
) or back fat thickness (Thiel et al. 1998
).
In this investigation, we also obtained serial measures of back fat
thickness by ultrasound throughout the study. Increases in back fat
occurred across the 8-wk treatment period for all groups, but the
magnitude of the increase was reduced in a linear manner with
increasing dietary level of CLA supplementation (Fig. 1)
. Although the
numerical differences progressively increased across the 8-wk treatment
period, they were not significant until wk 3 of treatment. At the
highest level of CLA supplementation, P2 back fat depth was
reduced by 25% (6 mm) at wk 8. It is interesting that despite
substantial reductions in carcass fat, the increase in protein was
relatively small (+5%). However, lean tissue (as defined as the sum of
water and protein) was increased commensurate with the reduction in
carcass fat (because ash was unchanged). Although the fat deposition
responses to CLA feeding were linear across the range of levels used,
the lean tissue response was quadratic, with the maximum rate of lean
tissue deposition occurring at a CLA level of 5.0 g/kg.
The mechanism by which CLA causes reduced body fat accretion is not
known. Effects could involve de novo lipogenesis, use of preformed
fatty acids for lipid synthesis, rates of lipolysis or some combination
of these. In this study, pigs were fed a high carbohydrate diet; thus,
de novo lipogenesis would represent the major mechanism of lipid
synthesis, and it would appear that CLA was acting to reduce these
rates. Other studies with growing animals (Chin et al. 1994
, Park et al. 1997
, 1999a
and 1999b
,
West et al. 1998
) have not had a zero time group to
allow for comparisons of rates of fat accretion over the treatment
period. However, West et al. (1998)
recently reported
that the carcass fat content at the end of the study was less in
CLA-treated mice fed either a high carbohydrate or a high fat diet.
Thus, rates of de novo synthesis and use of preformed fatty acids might
both be reduced by CLA. However, in all of the above cases, it is
possible that the effects of CLA could be on lipolysis, for which rates
of mobilization would have to be increased sufficiently to result in a
net reduction in lipid accretion. In investigations with lactating
cows, it was demonstrated that the major effect of CLA is to reduce
milk fatty acids arising from de novo synthesis (Chouinard et al. 1999
). In addition, the cis-8,
trans-12 CLA isomer decreases lipoprotein lipase
activity and triglyceride accumulation while increasing glycerol
release in cultured 3T3-L1 adipocytes. These data suggest changes in
both lipid synthesis and breakdown. Recent data from our laboratory
show that plasma triglyceride and nonesterified fatty acid
concentrations are increased during CLA feeding in pigs (E. Ostrowska,
unpublished observations), suggesting alterations in both the uptake of
preformed fatty acids and fat breakdown. Obviously, defining the
mechanism of CLA requires a more definitive understanding of the
specific dimensions of lipid metabolism that are responding to CLA
treatment.
It is obvious that dietary CLA can dramatically change the chemical
composition of the carcass. What is intriguing though is that these
favorable alterations in body composition were not associated with the
expected magnitude of improvement in the gain to feed ratio. From a
carcass gain perspective, the deposition of protein is more efficient
than fat because every gram of protein is associated with ~3 g of
water, whereas there is very little water associated with fat
deposition. Therefore, technologies or feeding strategies that have
resulted in similar alterations in the ratio of fat to lean are
generally associated with more profound improvements in the gain to
feed ratio than those observed with dietary CLA supplementation. For
example, dietary supplementation with the ß-agonist ractopamine
reduced the ratio of fat:protein by a magnitude similar to that seen
with CLA but resulted in a much greater improvements (15%) in the gain
to feed ratio (Dunshea et al. 1993b
and 1998
). A
possible explanation is that dietary CLA supplementation may result in
an increase in metabolic rate, although this was not manifest as an
increase in visceral organ size in this study (E. Ostrowska,
unpublished observations). However, studies with AKR/J mice have
indicated that CLA treatment increased metabolic rate (West et al. 1998
).
It is generally considered that the cis-9,
trans-11 CLA isomer is responsible for the
anticarcinogenic effects of CLA (Ip et al. 1994
). This
study, as well as all of the aforementioned studies looking at CLA
effects on growth, used a commercial source of CLA that contains a
number of isomers. Recently, Park et al. (1999b)
showed
that the cis-8, trans-12 CLA isomer of
CLA was much more potent than the cis-9,
trans-11 CLA isomer in reducing the body fat content of
mice. Similarly, work in dairy cows indicates that the CLA-induced
reduction in milk fat synthesis involves isomers with a
trans-10 double bond (Bauman et al. 1998
). Identifying the role of specific CLA isomers should aid
in understanding the mechanisms responsible for the diverse biological
functions reported for CLA.
In conclusion, dietary CLA supplements increased lean tissue deposition and decreased fat deposition in pigs. Chemical analysis of carcass composition revealed that the rate of lean tissue deposition was maximized at a CLA inclusion level of 5.0 g/kg, whereas the depression in fat deposition was linear up to at least 10 g CLA/kg.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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2 Supported by Pig Research and Development Corporation.
3 Recipient of a Swinburne University of Technology postgraduate award.
Manuscript received March 29, 1999. Initial review completed May 28, 1999. Revision accepted July 19, 1999.
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A. H. Terpstra Effect of conjugated linoleic acid on body composition and plasma lipids in humans: an overview of the literature Am. J. Clinical Nutrition, March 1, 2004; 79(3): 352 - 361. [Abstract] [Full Text] [PDF]
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G. J. Hausman and S. Poulos Recruitment and differentiation of intramuscular preadipocytes in stromal-vascular cell cultures derived from neonatal pig semitendinosus muscles J Anim Sci, February 1, 2004; 82(2): 429 - 437. [Abstract] [Full Text] [PDF]
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S. Viswanadha, J. G. Giesy, T. W. Hanson, and M. A. McGuire Dose Response of Milk Fat to Intravenous Administration of the trans-10, cis-12 Isomer of Conjugated Linoleic Acid J Dairy Sci, October 1, 2003; 86(10): 3229 - 3236. [Abstract] [Full Text] [PDF]
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A. H. M. Terpstra, M. Javadi, A. C. Beynen, S. Kocsis, A. E. Lankhorst, A. G. Lemmens, and I. C. M. Mohede Dietary Conjugated Linoleic Acids as Free Fatty Acids and Triacylglycerols Similarly Affect Body Composition and Energy Balance in Mice J. Nutr., October 1, 2003; 133(10): 3181 - 3186. [Abstract] [Full Text] [PDF]
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L. Granlund, L. K. Juvet, J. I. Pedersen, and H. I. Nebb Trans10, cis12-conjugated linoleic acid prevents triacylglycerol accumulation in adipocytes by acting as a PPAR{gamma} modulator J. Lipid Res., August 1, 2003; 44(8): 1441 - 1452. [Abstract] [Full Text] [PDF]
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J. M. Brown, M. S. Boysen, S. S. Jensen, R. F. Morrison, J. Storkson, R. Lea-Currie, M. Pariza, S. Mandrup, and M. K. McIntosh Isomer-specific regulation of metabolism and PPAR{gamma} signaling by CLA in human preadipocytes J. Lipid Res., July 1, 2003; 44(7): 1287 - 1300. [Abstract] [Full Text] [PDF]
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S. R. Demaree, C. D. Gilbert, H. J. Mersmann, and S. B. Smith Conjugated Linoleic Acid Differentially Modifies Fatty Acid Composition in Subcellular Fractions of Muscle and Adipose Tissue but Not Adiposity of Postweanling Pigs J. Nutr., November 1, 2002; 132(11): 3272 - 3279. [Abstract] [Full Text] [PDF]
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J. W. Perfield II, G. Bernal-Santos, T. R. Overton, and D. E. Bauman Effects of Dietary Supplementation of Rumen-Protected Conjugated Linoleic Acid in Dairy Cows during Established Lactation J Dairy Sci, October 1, 2002; 85(10): 2609 - 2617. [Abstract] [Full Text] [PDF]
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L. A. Gatlin, M. T. See, D. K. Larick, X. Lin, and J. Odle Conjugated Linoleic Acid in Combination with Supplemental Dietary Fat Alters Pork Fat Quality J. Nutr., October 1, 2002; 132(10): 3105 - 3112. [Abstract] [Full Text] [PDF]
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J.-C. Bouthegourd, P. C. Even, D. Gripois, B. Tiffon, M.-F. Blouquit, S. Roseau, C. Lutton, D. Tome, and J.-C. Martin A CLA Mixture Prevents Body Triglyceride Accumulation without Affecting Energy Expenditure in Syrian Hamsters J. Nutr., September 1, 2002; 132(9): 2682 - 2689. [Abstract] [Full Text] [PDF]
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A. H. M. Terpstra, A. C. Beynen, H. Everts, S. Kocsis, M. B. Katan, and P. L. Zock The Decrease in Body Fat in Mice Fed Conjugated Linoleic Acid Is Due to Increases in Energy Expenditure and Energy Loss in the Excreta J. Nutr., May 1, 2002; 132(5): 940 - 945. [Abstract] [Full Text] [PDF]
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M. Evans, X. Lin, J. Odle, and M. McIntosh Trans-10, Cis-12 Conjugated Linoleic Acid Increases Fatty Acid Oxidation in 3T3-L1 Preadipocytes J. Nutr., March 1, 2002; 132(3): 450 - 455. [Abstract] [Full Text] [PDF]
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E. E. Mosley, G. L. Powell, M. B. Riley, and T. C. Jenkins Microbial biohydrogenation of oleic acid to trans isomers in vitro J. Lipid Res., February 1, 2002; 43(2): 290 - 296. [Abstract] [Full Text] [PDF]
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J. M. Brown, Y. D. Halvorsen, Y. R. Lea-Currie, C. Geigerman, and M. McIntosh Trans-10, Cis-12, But Not Cis-9, Trans-11, Conjugated Linoleic Acid Attenuates Lipogenesis in Primary Cultures of Stromal Vascular Cells from Human Adipose Tissue J. Nutr., September 1, 2001; 131(9): 2316 - 2321. [Abstract] [Full Text] [PDF]
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M. B. Sisk, D. B. Hausman, R. J. Martin, and M. J. Azain Dietary Conjugated Linoleic Acid Reduces Adiposity in Lean but Not Obese Zucker Rats J. Nutr., June 1, 2001; 131(6): 1668 - 1674. [Abstract] [Full Text]
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L. H. Baumgard, J. K. Sangster, and D. E. Bauman Milk Fat Synthesis in Dairy Cows Is Progressively Reduced by Increasing Supplemental Amounts of trans-10, cis-12 Conjugated Linoleic Acid (CLA) J. Nutr., June 1, 2001; 131(6): 1764 - 1769. [Abstract] [Full Text]
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K. L. Ritzenthaler, M. K. McGuire, R. Falen, T. D. Shultz, N. Dasgupta, and M. A. McGuire Estimation of Conjugated Linoleic Acid Intake by Written Dietary Assessment Methodologies Underestimates Actual Intake Evaluated by Food Duplicate Methodology J. Nutr., May 1, 2001; 131(5): 1548 - 1554. [Abstract] [Full Text]
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G. Bee Dietary Conjugated Linoleic Acid Consumption during Pregnancy and Lactation Influences Growth and Tissue Composition in Weaned Pigs J. Nutr., December 1, 2000; 130(12): 2981 - 2989. [Abstract] [Full Text] [PDF]
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G. Bee Dietary Conjugated Linoleic Acids Alter Adipose Tissue and Milk Lipids of Pregnant and Lactating Sows J. Nutr., September 1, 2000; 130(9): 2292 - 2298. [Abstract] [Full Text]
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