A comparison of supplemental calcium soap of palm fatty acids versus tallow in a corn-based finishing diet for feedlot steers
© Warner et al. 2015
Received: 17 February 2015
Accepted: 13 May 2015
Published: 15 June 2015
Rumen bypass fat is commonly added to increase energy intake in dairy cattle. The objective of this study is to examine the addition of rumen bypass fat during finishing period on performance and carcass characteristics in grain fed steers. This study was conducted as a completely randomized block design with 126 cross-bred steer calves (initial BW 471.5 ± 7.5 kg) randomly assigned to pens with 9 steers/pen (n = 7 pens/treatment). Each pen was randomly assigned to one of two treatment groups; rumen bypass fat treatment (CCS, calcium soap of palm fatty acids) and control diet (CT, tallow). The diets were formulated to be isonitrogenous and isocaloric. Animals were fed twice daily at 110 % of the previous daily ad libitum intake. Blood from each sample was taken from the jugular vein. Muscle and adipose samples were collected from the longissimus dorsi regions. Feedlot performance and carcass characteristics were assessed. To examine adipogenic gene expression, quantitative real-time PCR was completed. Steers fed the CT had a greater level of performance for most of the parameters measured. The CT group had greater DMI (P < 0.05) and tended to have greater ADG (P < 0.10). Marbling score (P < 0.05) and quality grade (P < 0.05) were greater for steers fed the CT diet than those fed CCS. The longissimus muscle area tended to be greater (P < 0.10) in steers fed CT (87.60 cm2) than those fed CCS (84.88 cm2). The leptin mRNA expression was down-regulated (P < 0.05) in adipose tissue of steers fed a CCS when compared to those fed CT. These data suggest that calcium soap of palm fatty acids can be added to finishing diets without significant reduction in final body weight, although there may be modest reductions in marbling and quality scores.
KeywordsBeef Fatty acids Rumen bypass fat
Currently in the beef industry, the majority are fed high concentrate diets, which is ideal for rapid intramuscular adipose deposition, a characteristic desired for better meat quality . Ruminant fat is an important energy source consisting of predominantly saturated fatty acids (SFA), palmitic acid (C16:0) and stearic acid (C18:0), and polyunsaturated fatty acid (PUFA) . Naturally occurring trans fatty acids found in animal products are conjugated linoleic acids (CLA) and vaccenic acid (18:1, trans-11) . They are produced from PUFA via ruminal biohydrogenation and are known to be beneficial to human health . Ruminant fat has a lower PUFA: SFA ratio than non-ruminant fat . Saturated fatty acids may have negative health implications, such as increased serum low-density-lipoprotein (LDL) cholesterol concentration and increase risk of coronary heart disease in human . Thus, nutritional strategies that increase of PUFA levels in beef cattle are required to meet the demands of beef consumer. There has been interest over the past years to increase unsaturated fatty acids (UFA) levels in beef. With aims to decrease the SFA concentration of beef intramuscular adipose tissues, research is being conducted in order to minimize the SFA concentration and increase the UFA concentration in ruminant products while maintaining typical, cost efficient, grain-based feedlot finishing diets. One experimental method includes the supplementation of rumen-protected fats. They are insoluble lipids that bypass the rumen and are utilized as an energy source upon absorption in the small intestine. Fat supplementation was originally used to meet the high energy requirements of high producing dairy cattle . Unsaturated fatty acids are also more readily digestible than SFA; therefore increased UFA absorption may improve production efficiency. An increase in milk yield has also been reported by previous study in cows supplemented with 4 % bypass fat . The increased milk yield with supplemental bypass fat may be ascribed to an enhancement in energy balance. Corn based diet with protected fat also showed higher digestibility of ether extracts than corn based diet in beef cattle . Moreover, long-chain omega-3 fatty acids have been correlated to improved reproductive efficiency and carcass characteristics of ruminant animals [10, 11]. According to previous studies, UFA has been found in higher concentration in intramuscular fat of grass-fed beef cattle than in grain-fed [5, 12]. Alternatively, research indicates that grain-fed beef has a greater SFA concentration  and a lesser PUFA: SFA ratio than grass-fed beef . Therefore, calcium soap of palm fatty acids supplementation may be an alternative feeding strategy when fresh pasture is not available.
By the inclusion of calcium soap of palm oil fatty acids, UFA are protected from the harsh environment of the rumen allowing them to escape microbial fermentation and biohydrogenation, therefore entering the small intestine as unsaturated and being more efficiently utilized by the ruminant . Through extrapolation of bypass fats efficiency in dairy cattle, it is hypothesized that calcium soap of palm fatty acids supplementation high in UFA during finishing period will increase the circulating UFA in blood and consequently deposition into intramuscular meat in steers fed high corn-based finishing rations. The objective of this study was to examine the influence of feeding calcium soap of palm fatty acids on meat fatty acid composition in grain fed beef steers during finishing period.
Animal care and procedure
Experimental diet on DM basis
Ingredient Basis: % of DM
Corn Silage 50 % Gr
Corn Grain Flaked
Distillers Gr. + Soluble
Dry matter, % of as-fed
Neutral detergent fiber
NEm3, Mcal/cwt DM
NEg3, Mcal/cwt DM
Vitamin A3, IU/cwt DM
Vitamin E3, IU/cwt DM
Monensin3, g/ton DM
Tylosin3, g/ton DM
Blood samples were collected on d 0, 30 and 60. Approximately 12 mL of blood was taken from the jugular vein using 20 gauge, 1½ inch vacutainer needles at each collection for fatty acid analysis. Serum from each sample was collected and stored in −80 °C for fat determination. Steers were transported and harvested at commercial slaughterhouse (JBS Swift Slaughterhouse, Greeley, CO, USA) on d 62 and were slaughtered according to industry-accepted procedures.
Animal harvest and carcass evaluation
Upon slaughter, carcasses were weighed to obtain hot carcass weight (HCW). Muscle and adipose tissue samples were dissected from the longissimus dorsi region, flash frozen and stored at −80 °C for subsequent pulverization and fatty acid determination. Following the carcass-chilling period, carcass quality characteristics were assessed according to USDA standard techniques by trained Colorado State University personnel; marbling, quality grade, fat thickness, longissimus muscle (LM) area, KPH (kidney, pelvic, and heart fat), preliminary yield grade, final yield grade, and dressing percent.
Fatty acid composition analysis
Three steers were randomly selected from each experimental unit (pen). The longissimus dorsi region tissue were pulverized using a Waring blender. Blood serum samples were pooled for each experimental unit. Total lipid from blood serum and intramuscular adipose was extracted using the procedure outlined by Folch et al.  followed by methylation as described by Morrison et al. . Fatty acid methyl esters (FAME) were determined by gas chromatography (Hewlett-Packard, 6890 series II, CA, USA) equipped with a SP-2560 fused silica capillary column (100 m × 0.25 mm i.d.; Supelco Inc., PA, USA), with a series 7683 injector and flame ionization detector. Helium was used as the carrier gas with a flow rate of 2.1 mL/min. Fatty acids were identified by comparing the retention times displayed on the chromatographs to that of reference standards.
RNA isolation and quantitative real-time PCR (qRT-PCR)
Real-time PCR primer sequences
Fatty acid synthase
This study was conducted as a completely randomized block design with repeated measures, consisting of two treatment groups with 63 animals per treatment, divided into 7 pens. In order to minimize the effect of initial body weight, steers were assigned into 1–7 weight blocks based off of body weight at d 0. Rumen bypass fat treatment was randomly assigned into 7 pens, while the other 7 were fed the CT diet. Analysis of Variance (ANOVA) was performed on feedlot performance, fatty acid composition and carcass data were analyzed on a pen mean basis using PROC GLM of Statistical Analysis Software (SAS Institute, Inc., NC, USA). The model includes treatment as fixed effect and pen as random effect for USDA quality and yield grades, incidence of morbidity and mortality and fatty acid profile analysis. The treatment difference was detected using a modified students test. Responses are reported using LS means ± standard error. Significance was determined at P < 0.05 and tendency at P < 0.10.
Results and discussion
Feedlot performance and carcass characteristics of steers fed CT (tallow) or CCS (calcium soap of palm fatty acids) diet
Initial BW, kg
Final BW, kg
Adjusted fat thickness, cm
LM area, cm5
Hot carcass weight was not significantly different between treatments (P < 0.19). Marbling score (P < 0.05) and quality grade (P < 0.05) were greater for steers fed the CT diet than those fed CCS. The longissimus dorsi area tended to be greater (P < 0.10) in steers fed CT (87.60 cm2) than those fed CCS (84.88 cm2). The effects of rumen bypass fat on feedlot performance and carcass characteristics were in favor of the CT diet group. Decreased feed intake and BW gain coincides with the tendency (P = 0.10) for a lesser longissimus dorsi area in the CCS group compared to CT group. Marbling score and quality grade were greater for the CT than CCS. This may be attributed to increased intramuscular fatty acid deposition in steers fed the CT diet as a result of greater feed intake and subsequently greater weight gain. Carcass characteristics were statistically analyzed as categorical data assuming binomial distribution; quality grade for all steer were either select or choice and either had slight or small marbling scores. However, no treatment effects were detected for carcass evaluation.
Effects of CT (tallow) or CCS (calcium soap of palm fatty acids) diet on fatty acid composition of beef longissimus dorsi muscle (% of total fatty acids reported)
C18:1 t (total)
Effects of CT (tallow) or CCS (calcium soap of palm fatty acids) diet on fatty acid composition of serum (% of total fatty acids reported)
Trt x Time
C18:1 t (total)3
C18:2 t (total)4
The low rumen pH caused by high concentrate diets typically fed to feedlot cattle may contribute to minimal changes of fatty acid profile. Due to the differences in feed formulation of dairy versus feedlot cattle, bypass fat function differs at the opposing rumen pH. High forage concentrations of dairy cattle yield an increased pH, while the higher concentrate diets of feedlot cattle decreases rumen pH. Rumen bypass fats are more stable and work optimally at a normal rumen pH of 5.5–6.5 . It has been reported that ruminal fluid pH was 5.8 in cattle fed corn based diet with protected fat for 41 days . As pH decreases in the rumen, calcium soaps more readily dissociate leaving the free fatty acids susceptible to biohydrogenation into their saturated form for subsequent absorption. Therefore by increasing rumen pH more fatty acids may stay intact with the calcium ion, escaping biohydrogenation and dissociate in the abomasum and duodenum for subsequent digestion. Future studies under optimized ruminal pH conditions are needed to better understand the rumen bypass fat and UFA production mechanism in grain finishing beef cattle.
Altered mRNA expression of genes involved in lipid metabolism of steers adipose tissue
1.17 ± 0.08
0.82 ± 0.13
1.75 ± 0.24
1.69 ± 0.55
2.12 ± 0.75
1.19 ± 0.49
1.17 ± 0.12
0.68 ± 0.21
Other genes related to lipogenesis, including LPL, FASN and SCD were observed. LPL hydrolyzes triglyceride-rich lipoproteins to make NEFA for uptake in peripheral tissues . FASN regulates lipid biosynthesis from NADPH and acetyl-CoA in adipose tissue . LPL and FASN are important enzymes in adipocyte lipogenesis and have been found to have greater mRNA expression in the subcutaneous fat than in the intramuscular fat tissue . This may explain the similar levels of LPL and FASN expression observed from the tissue between treatments groups. While marbling score of carcass did statistically differ between groups, similar fat thickness was observed in the present study. Stearoyl-CoA desaturase 1 is an enzyme involved in the catalyzing the desaturation of saturated fatty acids to monounsaturated fatty acids (MUFA) . It has previously been reported that increasing n-3 PUFA composition in the diet caused a significant reduction in SCD expression in subcutaneous adipose tissue of Holstein bulls . It may be explained by the correlation with reduced gene expression levels of lipogenic transcription factor (sterol regulatory element binding protein 1c, SREBP1c) in adipose tissue. These results are in agreement with the findings of the current study as CCS supplementation showed decrease in SCD mRNA expression as compared to steers fed CT diet. Moreover, alpha-linolenic acid (C18:3 n-3) inhibited SCD gene expression in the adipocytes of mice . However, MUFA composition of longissimus dorsi muscle was not affected by CCS supplementation. In some studies, there was no correlation between Δ9-desaturase index and SCD gene expression in subcutaneous fat , and the Δ9-desaturase index does not correctly reflect the enzyme activity in cattle . Similarly, Wynn et al.  observed no effect of Megalac supplementation on SCD mRNA expression level in sheep. Further study is needed to clarify whether there is difference in SCD expression depends on supplementation methods and periods.
In conclusion, these data suggest that calcium soap of palm fatty acids can be added to finishing diets without significant reduction in final body weight, although there may be modest reductions in marbling and quality scores. Few significant differences were seen in the muscle and serum fatty acid composition. Adjustment of finishing diet formulation in favor of a more basic pH may increase PUFA in intramuscular adipose, although this may result in decreased ADG and therefore a longer time requirement to reach slaughter weight. Potential future research to consider for a significantly increase UFA content include feeding rumen bypass fat for an extended time period as opposed to the 60 days fed in this trial.
This project was supported by Colorado Corn Growers Association.
- Vernon RG. Lipid metabolism in the adipose tissue of ruminant animals. Prog Lipid Res. 1980;19(1–2):23–106.PubMedView ArticleGoogle Scholar
- Cordain L, Watkins BA, Florant GL, Kelher M, Rogers L, Li Y. Fatty acid analysis of wild ruminant tissues: evolutionary implications for reducing diet-related chronic disease. Eur J Clin Nutr. 2002;56(3):181–91.PubMedView ArticleGoogle Scholar
- Hodgson JM, Wahlqvist ML, Boxall JA, Balazs ND. Platelet trans fatty acids in relation to angiographically assessed coronary artery disease. Atherosclerosis. 1996;120(1–2):147–54.PubMedView ArticleGoogle Scholar
- Kepler CR, Hirons KP, McNeill JJ, Tove SB. Intermediates and products of the biohydrogenation of linoleic acid by Butyrinvibrio fibrisolvens. J Biol Chem. 1966;241(6):1350–4.PubMedGoogle Scholar
- French P, Stanton C, Lawless F, O’Riordan EG, Monahan FJ, Caffrey PJ, et al. Fatty acid composition, including conjugated linoleic acid, of intramuscular fat from steers offered grazed grass, grass silage, or concentrate-based diets. J Anim Sci. 2000;78(11):2849–55.PubMedGoogle Scholar
- Siri-Tarino PW, Sun Q, Hu FB, Krauss RM. Saturated fatty acids and risk of coronary heart disease: modulation by replacement nutrients. Curr Atheroscler Rep. 2010;12(6):384–90.PubMed CentralPubMedView ArticleGoogle Scholar
- Palmquist DL. Influence of source and amount of dietary fat on digestibility in lactating cows. J Dairy Sci. 1991;74(4):1354–60.PubMedView ArticleGoogle Scholar
- Purushothaman S, Kumar A, Tiwari DP. Effect of feeding calcium salts of palm oil fatty acids on performance of lactating crossbred cows. Asian Aust J Anim Sci. 2008;21(3):376–85.View ArticleGoogle Scholar
- Hill GM, West JW. Rumen protected fat in kline barley or corn diets for beef cattle: digestibility, physiological, and feedlot responses. J Anim Sci. 1991;69(8):3376–88.PubMedGoogle Scholar
- Mattos R, Staples CR, Thatcher WW. Effects of dietary fatty acids on reproduction in ruminants. Rev Reprod. 2000;5(1):38–45.PubMedView ArticleGoogle Scholar
- Ponnampalam EN, Sinclairt AJ, Egan AR, Blakeley SJ, Leury BJ. Effect of diets containing n-3 fatty acids on muscle long-chain n-3 fatty acid content in lambs fed low- and medium-quality roughage diets. J Anim Sci. 2001;79(3):698–706.PubMedGoogle Scholar
- Dannenberger D, Nuernberg K, Nuernberg G, Scollan N, Steinhart H, Ender K. Effect of pasture vs. concentrate diet on CLA isomer distribution in different tissue lipids of beef cattle. Lipids. 2005;40(6):589–98.PubMedView ArticleGoogle Scholar
- Realini CE, Duckett SK, Hill NS, Hoveland CS, Lyon BG, Sackmann JR, et al. Effect of endophyte type on carcass traits, meat quality, and fatty acid composition of beef cattle grazing tall fescue. J Anim Sci. 2005;83(2):430–9.PubMedGoogle Scholar
- Duckett SK, Wagner DG, Yates LD, Dolezal HG, May SG. Effects of time on feed on beef nutrient composition. J Anim Sci. 1993;71(8):2079–88.PubMedGoogle Scholar
- Folch J, Lees M, Sloane Stanley GH. A simple method for the isolation and purification of total lipides from animal tissues. J Biol Chem. 1957;226(1):497–509.PubMedGoogle Scholar
- Morrison WR, Smith LM. Preparation of fatty acid methyl esters and dimethylacetals from lipids with boron fluoride–methanol. J Lipid Res. 1964;5:600–8.PubMedGoogle Scholar
- Schmittgen TD, Livak KJ. Analyzing real-time PCR data by the comparative C(T) method. Nat Protoc. 2008;3(6):1101–8.PubMedView ArticleGoogle Scholar
- Araujo DB, Cooke RF, Hansen GR, Staples CR, Arthington JD. Effects of rumen-protected polyunsaturated fatty acid supplementation on performance and physiological responses of growing cattle after transportation and feedlot entry. J Anim Sci. 2010;88(12):4120–32.PubMedView ArticleGoogle Scholar
- Spector AA. Fatty acid binding to plasma albumin. J Lipid Res. 1975;16(3):165–79.PubMedGoogle Scholar
- Ha YL, Storkson J, Pariza MW. Inhibition of benzo(a)pyrene-induced mouse forestomach neoplasia by conjugated dienoic derivatives of linoleic acid. Cancer Res. 1990;50(4):1097–101.PubMedGoogle Scholar
- Ip C, Singh M, Thompson HJ, Scimeca JA. Conjugated linoleic acid suppresses mammary carcinogenesis and proliferative activity of the mammary gland in the rat. Cancer Res. 1994;54(5):1212–5.PubMedGoogle Scholar
- Parodi PW. Conjugated linoleic acid and other anticarcinogenic agents of bovine milk fat. J Dairy Sci. 1999;82(6):1339–49.PubMedView ArticleGoogle Scholar
- Nugent AP, Roche HM, Noone EJ, Long A, Kelleher DK, Gibney MJ. The effects of conjugated linoleic acid supplementation on immune function in healthy volunteers. Eur J Clin Nutr. 2005;59(6):742–50.PubMedView ArticleGoogle Scholar
- Moloney F, Yeow TP, Mullen A, Nolan JJ, Roche HM. Conjugated linoleic acid supplementation, insulin sensitivity, and lipoprotein metabolism in patients with type 2 diabetes mellitus. Am J Clin Nutr. 2004;80(4):887–95.PubMedGoogle Scholar
- Nicolosi RJ, Laitinen L. Dietary conjugated linoleic acid reduces aortic fatty streak formation greater than linoleic acid in hypercholesterolemic hamsters. Faseb J. 1996;10(3):2751–1.Google Scholar
- Sukhija PS, Palmquist DL. Dissociation of calcium soaps of long-chain fatty acids in rumen fluid. J Dairy Sci. 1990;73(7):1784–7.PubMedView ArticleGoogle Scholar
- McFadin EL, Keisler DH, Schmidt TB, Lorenzen CL, Berg EP. Correlations between serum concentrations of leptin and beef carcass composition and quality. J Muscle Foods. 2003;14(1):81–7.View ArticleGoogle Scholar
- Nkrumah JD, Li C, Yu J, Hansen C, Keisler DH, Moore SS. Polymorphisms in the bovine leptin promoter associated with serum leptin concentration, growth, feed intake, feeding behavior, and measures of carcass merit. J Anim Sci. 2005;83(1):20–8.PubMedGoogle Scholar
- Geary TW, McFadin EL, MacNeil MD, Grings EE, Short RE, Funston RN, et al. Leptin as a predictor of carcass composition in beef cattle. J Anim Sci. 2003;81(1):1–8.PubMedGoogle Scholar
- Merkel M, Eckel RH, Goldberg IJ. Lipoprotein lipase: genetics, lipid uptake, and regulation. J Lipid Res. 2002;43(12):1997–2006.PubMedView ArticleGoogle Scholar
- Ranganathan G, Unal R, Pokrovskaya I, Yao-Borengasser A, Phanavanh B, Lecka-Czernik B, et al. The lipogenic enzymes DGAT1, FAS, and LPL in adipose tissue: effects of obesity, insulin resistance, and TZD treatment. J Lipid Res. 2006;47(11):2444–50.PubMed CentralPubMedView ArticleGoogle Scholar
- Pickworth CL, Loerch SC, Velleman SG, Pate JL, Poole DH, Fluharty FL. Adipogenic differentiation state-specific gene expression as related to bovine carcass adiposity. J Anim Sci. 2011;89(2):355–66.PubMedView ArticleGoogle Scholar
- Li X, Ekerljung M, Lundstrom K, Lunden A. Association of polymorphisms at DGAT1, leptin, SCD1, CAPN1 and CAST genes with color, marbling and water holding capacity in meat from beef cattle populations in Sweden. Meat Sci. 2013;94(2):153–8.PubMedView ArticleGoogle Scholar
- Hiller B, Herdmann A, Nuernberg K. Dietary n-3 fatty acids significantly suppress lipogenesis in bovine muscle and adipose tissue: A functional genomics approach. Lipids. 2011;46:557–67.
- Sessler AM, Kaur N, Palta JP, Ntambi JM. Regulation of stearoyl-CoA desaturase 1 mRNA stability by polyunsaturated fatty acids in 3 T3-L1 adipocytes. J Biol Chem. 1996;271(47):29854–8.PubMedView ArticleGoogle Scholar
- Corazzin M, Bovolenta S, Sepulcri A, Piasentier E. Effect of whole linseed addition on meat production and quality of Italian Simmental and Holstein young bulls. Meat Sci. 2012;90(1):99–105.PubMedView ArticleGoogle Scholar
- Archibeque SL, Lunt DK, Gilbert CD, Tume RK, Smith SB. Fatty acid indices of stearoyl-CoA desaturase do not reflect actual stearoyl-CoA desaturase enzyme activities in adipose tissues of beef steers finished with corn-, flaxseed-, or sorghum-based diets. J Anim Sci. 2005;83(5):1153–66.PubMedGoogle Scholar
- Wynn RJ, Daniel ZC, Flux CL, Craigon J, Salter AM, Buttery PJ. Effect of feeding rumen-protected conjugated linoleic acid on carcass characteristics and fatty acid composition of sheep tissues. J Anim Sci. 2006;84(12):3440–50.PubMedView ArticleGoogle Scholar
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