1) Sixteen pigs were randomly assigned to diets containing either 5% biohydrogenation product (BHP)-enriched beef fat or control beef fat for 7 weeks. 2) Feeding the enriched fat led to deposition of various BHPs, including trans fatty acids and conjugated linoleic acids, in the pigs' subcutaneous fat tissues. 3) While growth performance and body composition were unaffected, total and HDL cholesterol decreased in pigs fed the enriched fat diet, though LDL cholesterol and triglycerides did not change.

1. SHORT COMMUNICATION
Effects of feeding beef fat enriched with polyunsaturated
fatty acid biohydrogenation products to pigs
Payam Vahmani, W. Jon Meadus, Bethany Uttaro, Óscar L´opez-Campos, Cletos Mapiye,
David C. Rolland, William R. Caine, Jennifer L. Aalhus, and Michael E.R. Dugan
Abstract: A total of sixteen barrows were randomly assigned to diets containing 5% biohydrogenation product
(BHP)-enriched or control beef fat for 7 weeks. On completion of 7 weeks, we found that feeding enriched fat led
to deposition of BHP and isomer-specific metabolism of trans-18:1 in adipose tissue. It was also noticed that total
and HDL-cholesterol were decreased; however, LDL-cholesterol and triglycerides were not affected.
Key words: beef, kidney fat, trans 18:1 isomers, vaccenic acid, conjugated linoleic acid.
Résumé : Seize castrats ont été assignés de façon aléatoire à des diètes enrichies de 5% de produit de
biohydrogénation (BHP) ou témoin à base de gras de bœuf pendant 7 semaines. Alimenter les animaux en gras
enrichi se soldait par la déposition de BHP et le métabolisme spécifique à l’isomère du gras trans-18:1 dans les tissus
adipeux. Le cholestérol total et HDL-cholestérol ont diminué lorsque les animaux ont reçu le gras enrichi. Par
contre, il n’y a pas eu d’effet sur le LDL-cholestérol et les triglycérides. [Traduit par la Rédaction]
Mots-clés : bœuf, gras rénal, isomères trans 18:1, acide vaccénique, acide linoléique conjugué.
There is an increasing interest in developing beef and
dairy products enriched with polyunsaturated fatty acid
(PUFA) biohydrogenation products (BHPs), including
rumenic acid (RA, cis9,trans11-18:2), the most abundant
natural conjugated linoleic acid (CLA) isomer, and its
precursor vaccenic acid (VA, trans11-18:1). This new inter-
est is due to the discovery that these fatty acids have anti-
carcinogenic properties and can improve blood lipid
profiles in animal models (De La Torre et al. 2006; Dilzer
and Park 2012; Field et al. 2009; Wang et al. 2009).
Trans10,cis12-18:2, the CLA isomer most often found in
synthetic CLA preparations, has been found to reduce
body fat and increase muscle mass (Dugan et al. 1997;
Kennedy et al. 2010). However, there are many other
BHP found in beef and dairy products whose physiologi-
cal effects have not been investigated (Dugan et al. 2011;
Shingfield et al. 2013). The objective of the present study
was to examine the effects of feeding pigs two sources of
beef kidney fat with vastly different BHP profiles.
Control kidney fat was collected from steers fed a barley
grain-based diet, and kidney fat enriched with BHP from
α-linolenic acid (ALA, 18:3n-3) was harvested from steers
fed flaxseed in a grass hay-based diet (Mapiye et al.
2013). The predominant BHP in control beef fat was
trans10-18:1, which is known to negatively impact blood
lipid profiles (Bauchart et al. 2007). The most abundant
BHP in enriched kidney fat was VA followed by several
BHP specific to ALA (Table 1).
Eight Large White × Duroc barrows were individually
fed each diet. Animals were cared for according to
Canadian Council on Animal Care guidelines (CCAC
2009). From 60 to 110 kg body weight, pigs were fed a
standard 16% CP finishing diet composed of 41.7% wheat,
35.8% barely, 14.3% canola meal, and 0.95% soybean meal
with the addition of 1.14% calcium carbonate, 0.31% dical-
cium phosphate, 0.41% salt, 0.13% lysine, 0.22% vitamin/
mineral premix, and 5% (w/w) BHP-enriched or control
beef kidney fat. Feed was weighed daily into individual
Received 23 April 2015. Accepted 2 October 2015.
P. Vahmani, W.J. Meadus, B. Uttaro, Ó. L ´opez-Campos, D.C. Rolland, J.L. Aalhus, M.E.R. Dugan. Agriculture and Agri-Food Canada,
Lacombe Research and Development Centre, Lacombe, AB T4L 1W1, Canada.
C. Mapiye, Department of Animal Sciences, Stellenbosch University, Stellenbosch, Western Cape, South Africa.
W.R. Caine. Caine Research Consulting, P.O. Box 1124, Nisku, AB T9E 8A8, Canada.
Corresponding author: Michael E.R. Dugan (email: mike.dugan@agr.gc.ca).
Abbreviations: ALA, α-linolenic acid; BHP, biohydrogenation products; CLA, conjugated linoleic acid; DEXA, dual x-ray absorbance;
RA, rumenic acid; TAG, triglyceride.
Copyright remains with the author(s) or their institution(s). Permission for reuse (free in most cases) can be obtained from RightsLink.
95
Can. J. Anim. Sci. 96: 95–99 (2016) dx.doi.org/10.1139/cjas-2015-0080 Published at www.nrcresearchpress.com/cjas on 29 April 2016.
Can.J.Anim.Sci.Downloadedfromwww.nrcresearchpress.combyAgricultureandAgri-foodCanadaon05/02/16
Forpersonaluseonly.

2. feeders, and feed weigh backs and animal weights were
collected weekly. Two days prior to slaughter, blood sam-
ples were collected via the jugular vein after an 8- to 10-h
overnight fast. Blood was allowed to clot (30 min at room
temperature), and serum was obtained by centrifugation
at 4 °C at 1500g for 15 min. Serum triglycerides (TAGs)
were determined using a triglyceride quantification kit
(Biovision, Mountain View, CA). Serum total cholesterol,
HDL-cholesterol, and LDL-cholesterol were measured
using an HDL and LDL/VLDL cholesterol quantification
kit (Biovision, Mountain View, CA). All pigs were killed
at the Lacombe Research Centre Abattoir. Subcutaneous
fat was collected at the 11th rib above the loin, and fatty
acids for both feed and subcutaneous fat were analyzed
as described by Dugan et al. (2007). At 45-min post-mor-
tem, the thickness of subcutaneous fat and loin muscle
were determined on the left side of the carcass between
the 3rd and 4th last ribs approximately 7 cm from the
Table 1. Fatty acid composition of experimental diets and subcutaneous fat of barrows fed these diets for 7 weeks.
Fatty acid (g 100 g−1
)b
Dieta
Subcutaneous fat
CNT (n = 3) BHP (n = 3) SEMc
P CNT (n = 8) BHP (n = 8) SEMc
P
14:0 1.88 1.84 0.01 1.43 1.53 0.04
16:0 22.17 20.2 0.05 ** 24.31 25.03 0.33
18:0 15.36 19.65 0.15 ** 13.59 14.28 3.68 **
ΣSFA 44.89 42.48 0.18 * 40.15 41.66 0.62
c9-16:1 1.42 0.91 0.02 ** 2.12 2.09 0.09
c9-18:1 28.21 21.33 0.02 ** 39.55 36.90 0.37 **
c11-18:1 1.59 1.14 0.06 * 2.83 2.59 0.07 *
Σc-MUFA 33.01 25.56 0.03 ** 46.51 43.53 0.45 **
t6-t8-18:1 0.20 0.37 0.01 ** 0.04 0.07 <0.01 **
t9-18:1 0.18 0.33 <0.01 ** 0.13 0.17 <0.01 **
t10-18:1 1.94 0.35 0.02 ** 0.51 0.12 0.02 **
t11-18:1 0.96 3.42 0.03 ** 0.14 0.54 0.02 **
t12-18:1 0.14 0.66 <0.01 ** 0.03 0.16 0.01 **
t13 + t14-18:1 0.27 1.61 <0.01 ** 0.08 0.19 0.01 **
t15-18:1 0.24 0.88 0.01 ** 0.09 0.16 0.01 **
t16-18:1 0.12 0.62 0.01 ** 0.02 0.09 <0.01 **
Σt-18:1 4.11 8.39 0.02 ** 1.04 1.50 0.06 **
t11,t15-18:2 0.02 0.18 <0.01 ** 0.01 0.08 <0.01 **
c9,t13-/t8,c12-18:2 0.07 0.20 <0.01 ** 0.03 0.11 <0.01 **
t8,c13-18:2 0.02 0.09 <0.01 ** 0.01 0.06 <0.01 **
c9,t12-18:2-18:1 0.06 0.13 <0.01 ** 0.03 0.05 <0.01 **
t11,c15-18:2 0.15 0.93 0.02 ** 0.04 0.23 0.01 **
c9,c15-18:2 0.16 0.15 0.01 0.13 0.13 0.01
c12,c15-18:2 ND 0.19 <0.01 ** ND 0.07 <0.01 **
ΣAD 0.51 1.95 0.03 ** 0.27 0.71 0.02 **
c9,t11-18:2 0.16 0.40 0.02 ** 0.23 0.56 0.02 **
t11,c13-18:2 ND 0.13 <0.01 ** ND 0.04 <0.01 **
t7,c9-18:2 0.03 0.02 <0.01 * 0.02 0.03 <0.01 *
ΣCLA 0.22 0.77 0.01 ** 0.30 0.71 0.02 **
18:2n-6 17.35 15.82 0.26 9.37 9.31 0.29
20:4n-6 0.02 0.01 <0.01 0.15 0.14 0.01
Σn-6 PUFA 17.49 15.95 0.28 10.04 9.93 0.30
18:3n-3 2.12 2.23 0.02 0.99 1.08 0.03
20:5n-3 ND ND 0.01 0.02 <0.01 *
22:5n-3 0.04 0.06 0.01 * 0.07 0.08 <0.01 *
Σn-3 PUFA 2.16 2.29 0.03 1.26 1.37 0.04 *
Note: ND is not detected; *, significant at P < 0.05; **, significant at P < 0.01.
a
CNT = control diet containing 5% (w/w) kidney fat from cattle fed control barley based diet; BHP = biohydrogenation
products-enriched diet containing 5% (w/w) kidney fat from cattle fed a hay-based diet containing flaxseed.
b
c = cis; t = trans; ΣSFA = sum of saturated fatty acids; ΣMUFA = sum of monounsaturated fatty acids; ΣAD = sum of
atypical dienes; ΣPUFA = sum of polyunsaturated fatty acids; ΣCLA = sum of conjugated linoleic acids.
c
Standard error of the mean.
96 Can. J. Anim. Sci. Vol. 96, 2016
Published by NRC Research Press
Can.J.Anim.Sci.Downloadedfromwww.nrcresearchpress.combyAgricultureandAgri-foodCanadaon05/02/16
Forpersonaluseonly.

3. midline using an Anitech PG100 Grading Probe (Anitech
Information Systems Inc., ON, Canada), which was used
to estimate lean yield. Following a 24-h chill at 2 °C, the
left side of the carcass was subjected to dual X-ray
absorbance (DEXA) measurements to estimate carcass
fat and lean (L ´opez-Campos et al. 2015). Data were ana-
lyzed using the MIXED procedure of SAS (SAS 2003).
Individual pig was the experimental unit. The model
included the fixed effect of diet and the random effect
of pig. Initial weight was included in the model as a cova-
riate for the analysis of growth performance and body
composition data. The significance level was set at
α = 0.05 and treatment means were generated using the
LSMEANS option of SAS.
Fat was added at 5% of the diet in the present study as
higher levels can lead to blocked feeders. However,
feeding BHP-enriched kidney fat had no effects on pig
body composition or growth performance, including
final body weight (111.1 ± 9.8 kg), average daily gain
(1.0 ± 0.11 kg), and feed efficiency (0.34 ± 0.04). This is in
contrast to studies which demonstrate that feeding
CLA-rich oils to pigs improved feed efficiency and body
composition (Dugan et al. 2004). Moreover, the present
study indicates that feeding BHP-enriched kidney fat to
pigs may not have any economic benefit in terms of ani-
mal production. Future studies in this area will, there-
fore, likely need to focus on effects of higher dietary fat
contents or fats with greater concentrations of BHP.
However, the fatty acid composition of the diets dif-
fered significantly, and the total content of BHP in control
and enriched beef fat were 4.6% and 10.4% (% of total fatty
acids), respectively. As a result, diets affected (P < 0.05)
percentages of many fatty acids in subcutaneous adipose
tissue (Table 1). This may be the first time many BHP
(trans18:1, atypical dienes, and CLA) isomers from beef
fat have been shown to be deposited during an animal
feeding trial. This included several BHP specific to ALA
(e.g., trans11,cis15-18:2), which could have potential physio-
logical/health effects similar to CLA (Churruca et al. 2009).
To gain insight into the metabolism of BHP, we exam-
ined the distribution of BHP in the diet relative to subcu-
taneous fat (Fig. 1). Interestingly, some trans18:1 isomers
including trans11-18:1 and trans13-18:1 appeared to undergo
delta-9 desaturation in pig adipose tissue. This resulted in
decreased proportions of trans11-18:1 and trans13-18:1 and
increased proportions of their delta-9 desaturation prod-
ucts (i.e., cis9,trans11-18:2 and cis9,trans13-18:2) in adipose
tissues compared to the diets. In contrast, the proportion
of trans9-18:1 increased in adipose tissue compared to the
diets, which is likely due to the fact that trans9-18:1 does
not undergo delta-9 desaturation (Holman and Mahfouz
1981). Consistent with these findings, trans9-18:1 was
found to accumulate to a greater extent than trans11-18:1
or trans13-18:1 in mouse adipocytes cultured with individ-
ual trans-18:1 isomers (Vahmani et al. 2014). Trans10-18:1
also does not undergo delta 9-desaturation (Holman and
Mahfouz 1981). However, in contrast to trans9-18:1, the
percentage of trans10-18:1 was lower in adipose tissues
than in diets, which could be due to preferential metabo-
lism via β-oxidation or carbon chain elongation (Emken
1984). Trans9-18:1 and trans10-18:1 are the predominant
trans-18:1 isomers found in partially hydrogenated vegeta-
ble oils (Stender et al. 2008), and trans10-18:1 is also the
main BHP in beef fat from cattle fed high-grain diets
(Mapiye et al. 2012). Present results suggest that trans-18:1
isomers are metabolized differently and there is a need
to examine physiological effects of BHP on an individual
basis (Mapiye et al. 2012).
Serum concentrations of total cholesterol and
HDL-cholesterol were lower (P < 0.05) in pigs fed
BHP-enriched fat compared to those fed control beef fat
Fig. 1. Percent distribution of biohydrogenation products
(BHPs) in diets containing 5% control (CNT) or BHP-enriched
beef, and in subcutaneous fat (SQ) of barrows fed these diets
for 7 weeks. Panels (a) and (b) show distribution of major and
minor BHP, respectively. Data are expressed as mean ± SEM
(n = 3 for diet; n = 8 for SQ). t = trans; c = cis; * = not detected.
Vahmani et al. 97
Published by NRC Research Press
Can.J.Anim.Sci.Downloadedfromwww.nrcresearchpress.combyAgricultureandAgri-foodCanadaon05/02/16
Forpersonaluseonly.

4. (58.7 ± 2.0 vs. 67.5 ± 2.6 and 25.3 ± 1.0 vs. 30.2 ± 1.3 mg dL−1
,
respectively; mean ± SEM). However, LDL-cholesterol,
TAG, and atherogenic indices (LDL/HDL and total choles-
terol/HDL ratios) were not affected (data not presented).
Reductions in HDL-cholesterol have also been observed
in studies in which BHP-enriched butters were fed to
hypercholesterolemic rabbits (Bauchart et al. 2007) or
healthy human subjects (Lacroix et al. 2012; Tholstrup
et al. 2006). In contrast, Lock et al. (2005) observed that
hamsters fed a BHP-enriched butter had lower plasma
total and LDL-cholesterol concentrations, and no change
in HDL cholesterol compared with those fed standard
butter. Conversely, feeding BHP-enriched butter to grow-
ing pigs (Haug et al. 2008) or to middle-aged men (Tricon
et al. 2006) had no effect on blood lipoprotein profiles.
Differences in results may relate to the basal diets and
the amount and composition of BHP, as well as species
differences.
In a recent review of 41 human clinical trials, Brouwer
et al. (2013) showed that replacement of control fat with
CLA or ruminant trans fatty acids increases plasma LDL,
and total cholesterol/HDL and LDL/HDL ratios. The
authors concluded that all trans fatty acids (i.e., partially
hydrogenated vegetable oils, ruminant trans fatty acids,
and CLA) have essentially the same effect on blood lipo-
proteins in humans. In contrast, data from epidemiologi-
cal studies have shown that ruminant trans fatty acid
(i.e., BHP) intake is not associated with an increased
cardiovascular disease risk (Ascherio et al. 1994; Jakobsen
et al. 2008; Pietinen et al. 1997). Consequently, it will be
important to determine if differences in blood lipopro-
teins when consuming ruminant trans fatty acids
actually translate into differences in the incidence of
cardiovascular disease.
Overall, feeding BHP-enriched beef kidney fat to pigs
had no effects on their growth performance or body
composition, but did lead to deposition of BHP in adi-
pose tissue. However, there also appeared to be differen-
tial metabolism of trans-18:1 isomers in pig adipose
tissue, with some undergoing delta-9 desaturation and
others being concentrated or preferentially metabolized.
Feeding BHP-enriched beef fat reduced serum total and
HDL-cholesterol concentrations in pigs, but atherogenic
indices including total cholesterol/HDL and LDL/HDL
ratios were not affected. To further elucidate the effects
of feeding BHP-enriched beef fats, these may need to be
fed at greater rates (i.e., similar to human diets, which
typically contain 30% dietary energy in the form of fat),
and employ disease models to determine if production
and consumption of these fats are of any health value.
This will then assist in development of ruminant feeding
strategies to avoid the incorporation of detrimental BHP
while promoting the uptake of beneficial BHP.
Acknowledgements
P. Vahmani acknowledges NSERC post-doctoral fund-
ing provided by the AAFC-Peer Review program. The
authors gratefully acknowledge the in-kind contribution
in animals, facilities and manpower received from AAFC
Lacombe. Ms I.L. Larsen and Mr. Adam Sebzda are
acknowledged for valuable assistance in statistical analy-
sis and ELISA assays, respectively.
References
Ascherio, A., Hennekens, C.H., Buring, J.E., Master, C.,
Stampfer, M.J., and Willett, W.C. 1994. Trans-fatty acids
intake and risk of myocardial infarction. Circulation, 89:
94–101. doi:10.1161/01.CIR.89.1.94. PMID:8281700.
Bauchart, D., Roy, A., Lorenz, S., Chardigny, J.M., Ferlay, A.,
Gruffat, D., Sebedio, J.L., Chilliard, Y., and Durand, D.
2007. Butters varying in trans 18:1 and cis-9, trans-11 conju-
gated linoleic acid modify plasma lipoproteins in the hyper-
cholesterolemic rabbit. Lipids, 42: 123–133. doi:10.1007/
s11745-006-3018-0. PMID:17393218.
Brouwer, I.A., Wanders, A.J., and Katan, M.B. 2013. Trans
fatty acids and cardiovascular health: research completed.
Eur. J. Clin. Nutr. 67: 541–547. doi:10.1038/ejcn.2013.43.
PMID:23531781.
CCAC. 2009. CCAC guidelines on: the care and use of farm
animals in research, teaching and testing. Canadian Council
on Animal Care (CCAC), Ottawa, ON.
Churruca, I., Fernández-Quintela, A., and Portillo, M.P. 2009.
Conjugated linoleic acid isomers: Differences in metabolism
and biological effects. BioFactors, 35: 105–111. doi:10.1002/
biof.v35:1.
De La Torre, A., Debiton, E., Juaneda, P., Durand, D., Chardigny,
J.M., Barthomeuf, C., Bauchart, D., and Gruffat, D. 2006. Beef
conjugated linoleic acid isomers reduce human cancer cell
growth even when associated with other beef fatty acids. Br.
J. Nutr. 95: 346–52. doi:10.1079/BJN20051634.
Dilzer, A., and Park, Y. 2012. Implication of conjugated
linoleic acid (CLA) in human health. Crit. Rev. Food
Sci. Nutr. 52: 488–513. doi:10.1080/10408398.2010.501409.
PMID:22452730.
Dugan, M., Aldai, N., Aalhus, J., Rolland, D., and Kramer, J.
2011. Review: Trans-forming beef to provide healthier fatty
acid profiles. Can. J. Anim. Sci. 91: 545–556. doi:10.4141/
cjas2011-044.
Dugan, M., Kramer, J., Robertson, W., Meadus, W., Aldai, N., and
Rolland, D. 2007. Comparing subcutaneous adipose tissue
in beef and muskox with emphasis on trans 18:1 and
conjugated linoleic acids. Lipids, 42: 509–518. doi:10.1007/
s11745-007-3051-7. PMID:17492324.
Dugan, M.E., Aalhus, J.L., and Kramer, J.K. 2004. Conjugated
linoleic acid pork research. Am. J. Clin. Nutr. 79: 1212s–1216s.
Dugan, M.E.R., Aalhus, J.L., Schaefer, A.L., and Kramer, J.K.G.
1997. The effect of conjugated linoleic acid on fat to lean
repartitioning and feed conversion in pigs. Can. J. Anim.
Sci. 77: 723–725. doi:10.4141/A97-084.
Emken, E.A. 1984. Nutrition and biochemistry of trans and posi-
tional fatty acid isomers in hydrogenated oils. Annu. Rev.
Nutr. 4: 339–376. doi:10.1146/annurev.nu.04.070184.002011.
PMID:6432011.
Field, C.J., Blewett, H.H., Proctor, S., and Vine, D. 2009. Human
health benefits of vaccenic acid. Appl. Physiol. Nutr. Metab.
34: 979–991. doi:10.1139/H09-079. PMID:19935865.
Haug, A., Sjogren, P., Holland, N., Muller, H., Kjos, N.P., Taugbol,
O., Fjerdingby, N., Biong, A.S., Selmer-Olsen, E., and Harstad,
O.M. 2008. Effects of butter naturally enriched with conju-
gated linoleic acid and vaccenic acid on blood lipids and
LDL particle size in growing pigs. Lipids Health Dis. 7: 31.
doi:10.1186/1476-511X-7-31. PMID:18759970.
98 Can. J. Anim. Sci. Vol. 96, 2016
Published by NRC Research Press
Can.J.Anim.Sci.Downloadedfromwww.nrcresearchpress.combyAgricultureandAgri-foodCanadaon05/02/16
Forpersonaluseonly.

5. Holman, R.T., and Mahfouz, M.M. 1981. Cis and trans-
octadecenoic acids as precursors of polyunsaturated acids.
Prog. Lipid Res. 20: 151–156. doi:10.1016/0163-7827(81)90028-X.
Jakobsen, M.U., Overvad, K., Dyerberg, J., and Heitmann, B.L.
2008. Intake of ruminant trans fatty acids and risk of
coronary heart disease. Int. J. Epidemiol. 37: 173–182. doi:10.
1093/ije/dym243. PMID:18077475.
Kennedy, A., Martinez, K., Schmidt, S., Mandrup, S., LaPoint, K.,
and McIntosh, M. 2010. Antiobesity mechanisms of action of
conjugated linoleic acid. J. Nutr. Biochem. 21: 171–179. doi:10.
1016/j.jnutbio.2009.08.003. PMID:19954947.
Lacroix, E., Charest, A., Cyr, A., Baril-Gravel, L., Lebeuf, Y.,
Paquin, P., Chouinard, P.Y., Couture, P., and Lamarche, B.
2012. Randomized controlled study of the effect of a
butter naturally enriched in trans fatty acids on blood
lipids in healthy women. Am. J. Clin. Nutr. 95: 318–325.
doi:10.3945/ajcn.111.023408. PMID:22205319.
Lock, A.L., Horne, C.A.M., Bauman, D.E., and Salter, A.M. 2005.
Butter naturally enriched in conjugated linoleic acid and
vaccenic acid alters tissue fatty acids and improves
the plasma lipoprotein profile in cholesterol-fed hamsters.
J. Nutr. 135: 1934–1939. PMID:16046719.
L´opez-Campos, Ó., Larsen, I., Prieto, N., Juárez, M., and Aalhus,
J.L. 2015. Using dual energy X-ray absorptiometry technology
for rapid and non-invasive fat and lean predictions in
beef carcasses. Meat. Sci. 99: 144. doi:10.1016/j.meatsci.2014.
07.015.
Mapiye, C., Aalhus, J.L., Turner, T.D., Rolland, D.C., Basarab, J.A.,
Baron, V.S., McAllister, T.A., Block, H.C., Uttaro, B., Lopez-
Campos, O., et al. 2013. Effects of feeding flaxseed or
sunflower-seed in high-forage diets on beef production,
quality and fatty acid composition. Meat. Sci. 95: 98–109.
doi:10.1016/j.meatsci.2013.03.033. PMID:23669875.
Mapiye, C., Aldai, N., Turner, T.D., Aalhus, J.L., Rolland, D.C.,
Kramer, J.K.G., and Dugan, M.E.R. 2012. The labile lipid
fraction of meat: From perceived disease and waste to health
and opportunity. Meat. Sci. 92: 210–220. doi:10.1016/j.meatsci.
2012.03.016. PMID:22546816.
Pietinen, P., Ascherio, A., Korhonen, P., Hartman, A.M., Willett,
W.C., Albanes, D., and Virtamo, J. 1997. Intake of fatty
acids and risk of coronary heart disease in a cohort of
Finnish men. The alpha-tocopherol, beta-carotene cancer
prevention study. Am. J. Epidemiol. 145: 876–887. doi:10.
1093/oxfordjournals.aje.a009047.
SAS. 2003. SAS user’s guide: Statistics. SAS for Windows.
Version. 9.1. SAS Institute, Inc., Cary, NC.
Shingfield, K.J., Bonnet, M., and Scollan, N.D. 2013. Recent
developments in altering the fatty acid composition of
ruminant-derived foods. Animal, 7: 132–162. doi:10.1017/
S1751731112001681. PMID:23031638.
Stender, S., Astrup, A., and Dyerberg, J. 2008. Ruminant and
industrially produced trans fatty acids: Health aspects.
Food. Nutr. Res. 52. doi:10.3402/fnr.v52i0.1651. PMID:19109659.
Tholstrup, T., Raff, M., Basu, S., Nonboe, P., Sejrsen, K., and
Straarup, E.M. 2006. Effects of butter high in ruminant
trans and monounsaturated fatty acids on lipoproteins,
incorporation of fatty acids into lipid classes, plasma
C-reactive protein, oxidative stress, hemostatic variables,
and insulin in healthy young men. Am. J. Clin. Nutr. 83:
237–243. PMID:16469980.
Tricon, S., Burdge, G.C., Jones, E.L., Russell, J.J., El-Khazen, S.,
Moretti, E., Hall, W.L., Gerry, A.B., Leake, D.S., Grimble, R.F.,
et al. 2006. Effects of dairy products naturally enriched with
cis-9, trans-11 conjugated linoleic acid on the blood lipid
profile in healthy middle-aged men. Am. J. Clin. Nutr. 83:
744–53. PMID:16600923.
Vahmani, P., Meadus, W.J., Turner, T.D., Duff, P., Rolland,
D.C., Mapiye, C., and Dugan, M.E.R. 2014. Individual trans
18:1 isomers are metabolised differently and have distinct
effects on lipogenesis in 3T3-L1 adipocytes. Lipids1–10.
PMID:16600923.
Wang, Y., Jacome-Sosa, M.M., Ruth, M.R., Goruk, S.D., Reaney,
M.J., Glimm, D.R., Wright, D.C., Vine, D.F., Field, C.J., and
Proctor, S.D. 2009. Trans-11 vaccenic acid reduces hepatic lipo-
genesis and chylomicron secretion in JCR: LA-cp rats. J. Nutr.
139: 2049–2054. doi:10.3945/jn.109.109488. PMID:19759243.
Vahmani et al. 99
Published by NRC Research Press
Can.J.Anim.Sci.Downloadedfromwww.nrcresearchpress.combyAgricultureandAgri-foodCanadaon05/02/16
Forpersonaluseonly.