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J. Anim. Sci. 2003. 81:3174-3181
© 2003 American Society of Animal Science

Effects of forage and sunflower oil levels on ruminal biohydrogenation of fatty acids and conjugated linoleic acid formation in beef steers fed finishing diets1

J. R. Sackmann*, S. K. Duckett*,2, M. H. Gillis*, C. E. Realini*, A. H. Parks{dagger} and R. B. Eggelston{dagger}

* Department of Animal and Dairy Science, College of Agricultural and Environmental Sciences and and {dagger} College of Veterinary Medicine, University of Georgia, Athens 30602


   Abstract
Top
 Abstract
Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Six Hereford steers (295 kg) cannulated in the proximal duodenum were used to evaluate the effects of forage and sunflower oil level on ruminal biohydrogenation (BH) and conjugated linoleic acid (CLA) outflow. Steers were fed one of six treatment diets in a 3 x 2 factorial arrangement of treatments (grass hay level: 12, 24, or 36% of DM; and sunflower oil level: 2 or 4% of DM) in a 6 x 6 Latin square design. The remainder of the diet was made up of steam rolled corn and protein/mineral supplement. Duodenal samples were collected for 4 d following 10-d diet adaptation periods. Data were analyzed with animal, period, forage level, sunflower oil level, and two-way interaction between forage and sunflower oil level in the model. Dry matter intake showed a quadratic response (P < 0.04), with an increase in DMI as forage level increased from 12 to 24% followed by a decrease in DMI when 36% forage was fed. Flow of fatty acids at the duodenum was higher (P < 0.03) for 4 vs. 2% sunflower oil diets, and similar among forage levels. Apparent ruminal digestibility of NDF increased in a linear manner (P < 0.04) as dietary forage level increased. Ruminal BH of dietary unsaturated 18-C fatty acids, oleic acid, and linoleic acid increased linearly (P < 0.05) as dietary forage level increased. Linoleic acid BH tended (P < 0.07) to be greater for 4 than 2% sunflower oil level. Duodenal flow of pentadecyclic, stearic, linolenic, and arachidic acids increased linearly (P < 0.05) as dietary forage level increased from 12 to 36%. Duodenal flow of linoleic acid decreased in a linear manner (P < 0.03) with increasing dietary forage level. Flow of trans-10 octadecenoate decreased linearly (P < 0.03) as dietary forage level increased, whereas trans-11 vaccenic acid flow to the duodenum increased (P < 0.01) linearly with increased dietary forage. Dietary forage or sunflower oil levels did not alter the outflow of cis-9, trans-11 CLA. Flows of cis-11, trans-13, and cis-9, cis-11 CLA increased linearly (P < 0.05) with increased dietary forage. Flows of cis-11, cis-13, and trans-11, trans-13 CLA decreased linearly (P < 0.05) with increased dietary forage. Increasing dietary forage levels from 12 to 36% in beef cattle finishing diets increased BH of unsaturated 18-C fatty acid and outflow of trans-11 vaccenic acid to duodenum without altering cis-9, trans-11 CLA outflow.

Key Words: Beef • Biohydrogenation • Conjugated Linoleic Acid


   Introduction
Top
 Abstract
 Introduction
Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Conjugated linoleic acid (CLA) is a collective term used to describe one or more positional and geometric isomers of linoleic acid (cis-9, cis-12-octadecadienoic acid). The cis-9, trans-11 isomer of CLA is an anticarcinogen, which reduced tumor proliferation when topically applied to mice with experimentally induced epidermal carcinogenesis (Ha et al., 1987Go). Chin et al. (1992)Go reported that CLA concentrations in ground beef are 3.8 to 4.3 mg/g of lipid, with about 84% being cis-9, trans-11 CLA isomer. Ritzenthaler et al. (2001)Go reported that beef and dairy products are the predominate sources of cis-9, trans-11 CLA isomer in the human diet, and beef contributes over 25% of the total dietary intake of cis-9, trans-11 CLA. Enhancing CLA levels in beef would be advantageous to human health.

Conjugated linoleic acid (CLA) and trans-octadecenoic acids are produced in the rumen as intermediates in the biohydrogenation of dietary linoleic acid to stearic acid (Bauman et al., 1999Go). Concentrations of CLA in milk fat increase with dietary supplementation of unsaturated vegetable oils (McGuire et al., 1996Go; Kelly et al., 1998Go) or alterations in dietary forage source (grass vs. alfalfa hay; Dhiman et al., 1999Go) and amount (50:50 vs. 35:65 forage to concentrate; Jiang et al., 1996Go; 1/3, 2/3, or all from pasture; 50:50, 67:33, 98:2 forage to concentrate, Dhiman et al., 1999Go). Previous research (Duckett et al., 2002Go) in cattle fed high-concentrate diets has shown that oil source, high oil corn vs. corn oil, can alter ruminal biohydrogenation and outflow of specific biohydrogenation intermediates. The hypothesis of this research was to determine whether dietary forage and/or sunflower oil levels increased the flow of CLA and trans-11 vaccenic acid to the small intestine. The objective of this research project was to determine the effect of dietary forage level and/or sunflower oil level on ruminal biohydrogenation and outflow of unsaturated fatty acids in beef steers fed finishing diets.


   Materials and Methods
Top
 Abstract
 Introduction
 Materials and Methods
Results and Discussion
 Implications
 Literature Cited
 
Animals.
Six Hereford steers (295 ± 23 kg) cannulated in the proximal duodenum were fed one of six diets in 3 x 2 factorial arrangement of treatments (grass hay level: 12, 24, or 36% of DM; and sunflower oil level: 2 or 4% of DM) in a 6 x 6 Latin square design. A T-cannula (University of Wyoming, Laramie) was inserted by a clinical veterinarian into the proximal duodenum of each steer while under local anesthesia. The Animal Care and Use Committee approved the surgical procedures and use of animals in this study. Following a 4-wk recovery period, the experiment began. Steers were housed inside the Large Animal Research Unit in individual pens with concrete slatted floors. Rubber mats were provided in each pen for animal comfort. Steers had ad libitum access to plain salt blocks throughout the duration of the study. One steer was removed from the study after completing the third period due to complications with cannula placement, unrelated to dietary treatments. Data collected from the first three periods for this steer were used in the statistical analyses, and data from the other three periods were treated as missing observations in the statistical analyses.

Diets.
In addition to grass hay and sunflower oil, diets included a soybean meal-based protein supplement and steam rolled corn. Ingredient and chemical composition of the diets are presented in Table 1Go. All diets were formulated to be isonitrogenous and included Rumensin fed at a level of 250 mg steer-1•d-1 monensin activity. Bermudagrass hay was coarsely chopped to approximately 5 cm in length. Animals were individually fed daily at 0800, with total daily intake being 95% of intake determined during a preliminary period in which feed supply was unlimited to minimize refusals during the sampling period. Chromic oxide (10 g steer-1•d-1) was added to the diets as an external marker for calculating duodenal flow.


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  		Table 1. Ingredient and chemical composition of diets fed to steers on a dry matter basisa
 
Sample Collection and Analysis.
Experimental periods lasted 14 d, with 10 d for diet adaptation and 4 d for sample collection. Following the 10-d adaptation period, duodenal samples (200 mL) were taken 12 times over 4 d, with sampling at 8-h intervals and sampling times shifted ahead by 2 h each day such that samples represented duodenal contents from 12 equally spaced intervals over a 24-h day. Feed samples were collected daily during each sampling period. Feed samples were immediately frozen after collection. Duodenal samples were composited on an equal liquid-volume basis for each steer within each period and frozen for later processing. Subsequently, duodenal digesta and feed samples were thawed, lyophilized, ground through a Wiley mill (1-mm screen) and frozen at -20°C for subsequent analyses of DM, protein, and ash content (AOAC, 1990Go), and fiber analysis according to Goering and Van Soest (1970)Go using an Ankom 220 (Ankom Technology, Fairport, NY).

Fatty acid composition of feed and duodenal samples was determined by the direct transmethylation procedure of Park and Goins (1994)Go. Methyl esters were separated by gas chromatography (Agilent 6850; Agilent Technologies, Wilmington, DE) using a 100-m Supelco SP2560 (Supelco, Bellefonte, PA) capillary column (0.25-mm i.d. and 0.20-µm film thickness) as described by Duckett et al. (2002)Go. Fatty acids were quantified by incorporating an internal standard, methyl tricosanoate (C23:0) into each sample before methylation.

Chromium concentration in digesta was measured via spectrophotometry (Fenton and Fenton, 1979Go). Concentrations of Cr and fatty acids in the feed, refusals, and digesta were used to calculate digestibility and duodenal flow of fatty acids (Schneider and Flatt, 1975Go). The percentage of biohydrogenation of total and individual unsaturated 18-carbon fatty acids was calculated according to Wu et al. (1991)Go.

Data were analyzed as a mixed linear model (SAS Inst. Inc., Cary, NC) to account for the fixed effects of forage level, oil level, and two-way interaction and the random effects of animal and period. All interactions between dietary forage and sunflower oil levels were nonsignificant (P > 0.05). Differences among dietary forage levels were separated using orthogonal contrasts to determine linear and quadratic responses. Differences between sunflower oil levels were separated using an orthogonal contrast to compare the two levels. Linear regression was used to develop equations for the change in outflow of trans-octadecenoic (trans-10 and trans-11) acids with increasing dietary forage. Significance was determined at P <= 0.05. Differences of P > 0.05 to P <= 0.10 are discussed as trends.


   Results and Discussion
Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
Implications
 Literature Cited
 
Dietary fatty acid content decreased linearly (P < 0.01) with increasing forage level, and was greater (P < 0.01) for 4% than for 2% sunflower oil addition (Table 1Go). The effects of forage and sunflower oil levels on intake, duodenal flow, and apparent ruminal digestibility are shown in Table 2Go. Dry matter intake showed a quadratic response (P < 0.04) with an increase in DMI as forage level increased from 12 to 24% followed by a decrease in DMI when 36% forage was fed. Kalscheur et al. (1997a)Go reported reductions in DM intake of dairy cows fed a low- (25%) vs. high- (60%) forage diet. Sunflower oil level did not alter (P < 0.39) DM intake. Others have reported no reduction in DM intake when supplementing corn oil (2.36%; Duckett et al., 2002Go), yellow grease (2 to 6%; Zinn et al., 2000Go), or high-oil corn (Eibs et al., 2000Go; Duckett et al., 2002Go) to finishing steer diets. Intake of long-chain fatty acids was similar when 12 or 24% forage was fed but lower when 36% forage was fed (linear, P < 0.01; quadratic, P < 0.02). Supplementing 4% sunflower oil to the diet, regardless of forage level, increased (P < 0.01) the intake of total long-chain fatty acids by 42% compared with 2% sunflower oil level.


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  		Table 2. Dry matter and long-chain fatty acid intake and ruminal digestion by steers fed different forage or sunflower oil levels
 
Dry matter flow at the duodenum showed a quadratic response (P < 0.05), with an increase as forage level increased from 12 to 24% followed by a decrease when 36% forage was fed. Duodenal DM flow did not differ between sunflower oil levels. Long-chain fatty acid (C10–20) flow at the duodenum did not differ between forage levels. Supplementing higher levels of sunflower oil to finishing cattle increased (P < 0.03) the flow of long-chain fatty acids to the duodenum by 58 g/d.

Apparent ruminal DM digestibility did not differ between forage or sunflower oil levels. However, numerical differences were observed in DM digestibility for 12 (+28%) compared with 24 and 36% forage diets. Apparent ruminal NDF digestibility increased in a linear manner (P < 0.04) with increasing forage level in the diet. Others (Kalscheur et al., 1997aGo) have shown lower apparent ruminal NDF digestibilities in dairy cows fed low vs. high levels of dietary forage. Apparent ruminal ADF and long-chain fatty acid digestibilities did not differ between forage levels. Dietary sunflower oil level did not alter apparent ruminal DM, NDF, or ADF digestibilities. Similarly, Kalscheur et al. (1997b)Go reported no changes in apparent ruminal NDF digestibilities in dairy cows supplemented with 3% sunflower oil or vegetable oil. In contrast, Zinn et al. (2000)Go reported reductions in organic matter and NDF digestibilities when supplementing high-concentrate diets (88% concentrate; wheat based) with yellow grease (2 to 6%) and/or formaldehyde-protected fat (2 to 4%). Feeding greater levels of sunflower oil increased (P < 0.02) long-chain fatty acid digestibility. These values for apparent ruminal digestibility of long-chain fatty acids are within the range of those reported by other researchers, as summarized by Jenkins (1993)Go.

Intakes (grams per day) of long-chain fatty acids are presented in Table 3Go. Intake of lauric (C12:0) and arachidic acids increased linearly (P < 0.01) as dietary forage level increased from 12 to 36%. Palmitic (C16:0) acid intake showed a quadratic response (P < 0.02), with an increase from 12 to 24% forage followed by a decrease when 36% forage was fed. Intake of palmitoleic (C16:1) and stearic (C18:0) acids was similar when 12 or 24% forage was fed, but it was lower when 36% forage was fed (quadratic, P < 0.03). Oleic (C18:1) and linoleic (C18:2) acid intakes decreased in a quadratic manner (P < 0.05) with increasing dietary forage levels. Intake of myristic acid, linolenic acid, and unidentified fatty acids increased (P < 0.05) in a quadratic manner as the level of forage increased in the diet. Supplementing higher sunflower oil levels in the finishing diet increased (P < 0.05) the intake of all individual long-chain fatty acids and unidentified fatty acids.


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  		Table 3. Long-chain fatty acid intake (g/d) by steers fed diets containing different forage and sunflower oil levels
 
Ruminal biohydrogenation of dietary unsaturated 18-carbon fatty acids as influenced by dietary forage and oil level is shown in Table 4Go. Ruminal biohydrogenation of total 18-C unsaturated fatty acids, oleic acid, and linoleic acid increased (P < 0.02) in a linear manner with increasing dietary forage levels. Biohydrogenation of dietary linolenic acid did not differ among dietary forage levels. Others (Kalscheur et al., 1997aGo; Kucuk et al., 2001Go) have shown that higher dietary forage levels increase biohydrogenation of dietary linoleic acid, linolenic acid, and overall unsaturated 18-C fatty acids. Latham et al. (1972)Go found that switching lactating dairy cows from a high- (44%) to a low- (20%) roughage diet resulted in lower levels of lipolytic activity and biohydrogenation of unsaturated fatty acids in ruminal fluid as measured by in vitro experiments. Dietary sunflower oil level did not alter ruminal biohydrogenation of total 18-C unsaturated fatty acids, oleic, or linolenic acid. Linoleic acid biohydrogenation tended (P <= 0.07) to increase with increasing dietary oil supplementation level. Wu et al. (1991)Go reported increased biohydrogenation of linoleic and linolenic acids with higher levels (3 vs. 6%) of supplemental fat (vegetable oil–animal fat blend; 35% saturated fatty acids). Duckett et al. (2002)Go reported increased biohydrogenation of all unsaturated 18-C fatty acids as dietary lipid level increased with supplementation of corn oil (2.4%) or substitution of normal corn for high-oil corn.


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  		Table 4. Effects of forage or sunflower oil level on ruminal biohydrogenation (%) of individual and overall unsaturated 18-C fatty acids
 
Flow (g/d) of long-chain fatty acids and biohydrogenation intermediates at the duodenum of steers fed diets containing different forage and sunflower oil levels is shown in Table 5Go. Flow of pentadecyclic (C15:0), stearic, linolenic, arachidic, and unidentified fatty acids to the duodenum increased (P < 0.05) in a linear manner as dietary forage level increased from 12 to 36%. Flow of myristic acid decreased linearly (P < 0.02) as dietary forage level increased. As dietary forage level increased, flow of linoleic acid decreased in a linear manner (P < 0.03). Similarly, Kalscheur et al. (1997a)Go reported increased flow of linoleic acid to the duodenum in low-fiber (25%) compared to high-fiber (60%) diets of lactating dairy cows as a result of lower biohydrogenation levels of unsaturated fatty acids. Flows of other long-chain fatty acids (capric, lauric, palmitic, palmitoleic, and oleic acids) were unaffected by dietary forage level. Supplementing higher levels of sunflower oil increased (P < 0.03) the flow of myristic and palmitic acids to the duodenum but did not alter the flow of any other long-chain fatty acid.


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  		Table 5. Flow (g/d) of long-chain fatty acid and biohydrogenation intermediates at the duodenum of steers fed diets containing different forage and sunflower oil levels
 
Biohydrogenation of dietary linoleic acid to stearic acid is sometimes incomplete, yielding several intermediates, including various trans- or cis-octadecenoic acids and CLA isomers (Bauman et al., 1999Go). Duodenal flow of trans-9-octadecenoic acids showed a quadratic response (P < 0.05), with an increase from 12 to 24% forage followed by no change with further increases in forage level. trans-10-Octadecenoic acid flow to the duodenum decreased linearly (P < 0.03; outflow, g/d = 54.24 - 1.06 x percentage of dietary forage; r2 = 0.21) as dietary forage level increased, whereas trans-11-vaccenic acid flow to the duodenum increased linearly (P < 0.01; outflow, g/d = 0.33 x percentage of dietary forage - 0.25; r2 = 0.47) as dietary forage level increased. trans-12-Octadecenoic acid increased (P < 0.02) in a linear manner as dietary forage level increased from 12 to 36%. cis-11-Octadecenoic acid flow increased in a quadratic manner (P < 0.03) with maximal levels for 24% forage diet. Flow of cis-12-octadecenoic acid was unchanged by dietary forage level. Sunflower oil level did not alter the flow of trans-9, -10, and -11 or cis-12-octadecenoic acids to the duodenum. Flow of trans-12-octadecenoic acid tended (P < 0.10) to be greater for 4 vs. 2% sunflower oil treatment. Flow of cis-11-octadecenoic acid increased (P < 0.05) as dietary sunflower oil level increased.

The predominant CLA isomers detected in duodenal digesta were trans-10, cis-12; cis-9, trans-11; and cis-9, cis-11 isomers, and these accounted for over 74% of total CLA present. The cis-9, trans-11; cis-8, cis-10; and trans-9, trans-11 isomers of CLA were unchanged by dietary forage or sunflower oil levels. Similarly, Beaulieu et al. (2002)Go reported no changes in cis-9, trans-11 isomer of CLA in ruminal contents or tissues of finishing cattle fed varying levels of soybean oil. Previous research in our laboratory (Duckett et al., 2002Go) also found no change in cis-9, trans-11 isomer of CLA with the addition of lipid to the diet either as corn oil or substitution of normal corn for high-oil corn. Duodenal flow of trans-10, cis-12 isomer of CLA decreased linearly (P < 0.05) as dietary forage levels increased and was greater (P < 0.04) for 4 vs. 2% sunflower oil. Beaulieu et al. (2002)Go and Duckett et al. (2002)Go both showed increased production of trans-10, cis-12 CLA isomer when supplemental oil was added to a high-concentrate diet. Flow of cis-11, trans-13 CLA increased linearly (P < 0.04) as dietary forage level increased, and tended (P < 0.10) to decrease with higher oil supplementation levels. Flow of cis-9, cis-11 CLA increased (P < 0.01) in a linear manner as dietary forage level increased. Flows of cis-11, cis-13 and trans-11, trans-13 CLA isomers decreased (P < 0.05) in a linear manner as dietary forage level increased. Flow of cis-10, cis-12 CLA isomer decreased in a linear manner (P < 0.03) with increasing dietary forage. Flow of other CLA isomers (cis-8, cis-10; cis-9, cis-11; cis-10, cis-12; cis-11, cis-13; trans-11, trans-13; trans-9, trans-11) was unchanged by dietary sunflower oil level. Regardless of dietary forage or sunflower oil level, the trans-10, cis-12 isomer of CLA was present in greater concentrations than the cis-9, trans-11 isomer of CLA in duodenal digesta. Bauman et al. (1999)Go has also shown that the trans-10, cis-12 isomer of CLA increases in concentration when lactating dairy cows are consuming a low-fiber/high-concentrate diet. The cis-9, trans-11 isomer of CLA was not influenced by dietary forage or sunflower oil level and flow of this isomer was less than 240 mg/d, a value 16 to 122 times lower than the outflow of trans-11-vaccenic acid depending on dietary forage level. Similarly, Duckett et al. (2002)Go reported low levels of cis-9, trans-11 isomer of CLA in duodenal digesta of steers fed high-concentrate diets containing high oil corn or added corn oil. Piperova et al. (2002)Go also reported low levels of cis-9, trans-11 CLA compared to trans-octadecenoic acids in duodenal digesta of lactating dairy cows fed low-fiber (25%) or high-fiber (60%) diets with or without the addition of 2% buffer. These results continue to demonstrate the importance of enhancing trans-11-vaccenic acid flow at the duodenum for increasing cis-9, trans-11 CLA isomer in beef fat.

Figure 1Go shows the duodenal outflow of trans-octadecenoic acids (trans-10 and trans-11) and CLA isomers (cis-9, trans-11 and trans-10, cis-12) by dietary forage level calculated as a percentage of linoleic acid intake. In typical high-concentrate diets (12% forage), trans-10-octadecenoic acid was present in the duodenal digesta at levels 12.2-fold greater than trans-11-vaccenic acid. Increasing dietary forage level in the finishing diet increased (P < 0.05) the outflow of trans-11-vaccenic acid and decreased (P < 0.05) the outflow of trans-10-octadecenoic acid as a percentage of linoleic acid intake. trans-10-Octadecenoic acid levels were 3.5-fold greater than trans-11-vaccenic acid for 24% forage diet. At the 36% forage level, trans-11 vaccenic acid flow to the duodenum as a percentage of linoleic acid intake approached the level of trans-10 octadecenoic acid. Attempts to increase CLA outflow to the duodenum by altering dietary forage or oil level were not successful because low levels (less than 0.23% of linoleic acid intake) of CLA are produced in the rumen regardless of dietary forage or oil level. However, the flow of trans-11-vaccenic acid to the duodenum seems to depend on dietary forage levels. Increasing dietary forage levels resulted in a linear increase in trans-11-vaccenic outflow. trans-11-Vaccenic acid can be desaturated to the cis-9 trans-11 isomer of CLA through the {Delta}-9 desaturase enzyme present in bovine mammary gland (Griinari et al., 2000Go). This enzyme is also present in bovine adipose tissues and is responsible for the desaturation of stearic acid to oleic acid, the predominant fatty acid in beef tissues (St. John et al., 1991Go). Research in dairy cattle has shown that 64% (Griinari et al., 2000Go) to 78% (Corl et al., 2001Go) of CLA in milk fat originated from desaturation of trans-11-vaccenic acid. Estimates from our laboratories (Gillis et al., 2003Go) suggest that over 86% of tissue CLA in beef originates from desaturation of trans-11-vaccenic acid based on ratios of trans-11-vaccenic acid to cis-9, trans-11 CLA in duodenal and adipose tissues. Because the outflow of CLA is very low compared to that of trans-11-vaccenic acid, selecting dietary parameters that increase the outflow of trans-11-vaccenic acid to the greatest degree should be most helpful in attempts to increase tissue CLA levels in beef cattle. Based on this research, feeding the high-forage diet (36%) resulted in the highest flows of trans-11 vaccenic acid to the duodenum. However, dietary DM intake and ruminal DM digestibility were lower for this high-forage diet, which may impact animal performance and warrants further research in this area.



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  		Figure 1. Duodenal output of trans-octadecenoic acids (trans-10 and -11) and conjugated linoleic acid (trans-10, cis-12 and cis-9, trans-11) isomers by dietary forage level as a percentage of linoleic (C18:2) acid intake (n = 11 per treatment).

 

   Implications
Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
Literature Cited
 
Increasing dietary forage levels from 12 to 36% in beef cattle finishing diets increased the biohydrogenation of linoleic acid and outflow of trans-11-vaccenic acid to duodenum, without altering cis-9, trans-11 conjugated linoleic acid outflow. Feeding higher levels of forage in finishing diets seems to alter ruminal biohydrogenation of linoleic acid, resulting in greater outflow of intermediates via the trans-11 pathway.


   Footnotes
 
1 Supported in part by Cattlemen’s Beef Board and National Cattlemen‘s Beef Association. Back

2 Correspondence: Animal and Dairy Science Complex, Athens 30602-2771 (E-mail: sduckett{at}uga.edu).

Received for publication April 14, 2003. Accepted for publication August 22, 2003.


   Literature Cited
Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 


AOAC. 1990. Official Methods of Analysis. 15th ed. Assoc. Offic. Anal. Chem., Washington, DC.

Bauman, D. E., L. H. Baumgard, B. A. Corl, and J. M. Griinari. 1999. Biosynthesis of conjugated linoleic acids in ruminants. Proc. Am. Soc. Anim. Sci., available at http://www.asas.org/jas/symposia/proceedings/0937.pdf. Accessed Sept. 1, 2002.

Beaulieu, A. D., J. K. Drackley, and N. R. Merchen. 2002. Concentrations of conjugated linoleic acid (cis-9 trans-11 octadecadienoic acid) are not increased in tissue lipids of cattle fed a high concentrate diet supplemented with soybean oil. J. Anim. Sci. 80:847–861.[Abstract/Free Full Text]

Chin, S. F., W. Lui, J. M. Storkson, Y. L. Ha, and M. W. Pariza. 1992. Dietary sources of conjugated dienoic isomers of linoleic acid, a newly recognized class of anticarcinogens. J. Food Comp. Anal. 5:185–197.

Corl, B. A., L. H. Baumgard, D. A. Dwyer, J. M. Griinari, B. S. Phillips, and D. E. Bauman. 2001. The role of delta-9 desaturase in the production of cis-9, trans-11 CLA. J. Nutr. Biochem. 12:622–630.[Medline]

Dhiman, T. R., G. R. Anand, L. D. Satter, and M. W. Pariza. 1999. Conjugated linoleic acid content of milk from cows fed different diets. J. Dairy Sci. 82:2146–2156.[Abstract]

Duckett, S. K., J. G. Andrae, and F. N. Owens. 2002. Effect of high oil corn or added corn oil on ruminal biohydrogenation of fatty acids and conjugated linoleic acid formation in beef steers fed finishing diets. J. Anim. Sci. 80:3353–3360.[Abstract/Free Full Text]

Eibs, M. S., B. J. Johnson, D. M. Wulf, B. D. Rops, and F. N. Owens. 2000. Effects of high-oil corn on feedlot performance, carcass characteristics, and meat quality. Page 76 in Proc. Plains Nutr. Counc. Spring Conf. Texas A&M Research and Education Center, Amarillo.

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Griinari, J. M., B. A. Corl, S. H. Lacy, P. Y. Chouinard, K. V. V. Nurmela, and D. E. Bauman. 2000. Conjugate linoleic acid is synthesized endogenously in lactating dairy cows by {Delta}9-desaturase. J. Nutr. 130:2285–2291.[Abstract/Free Full Text]

Goering, H. K., and P. J. Van Soest. 1970. Forage fiber analyses (apparatus, reagents, procedures, and some applications). Agric. Handbook 379. ARS, USDA, Washington, DC.

Ha, Y. L., N. K. Grimm, and M. W. Pariza. 1987. Anticarcinogens from fried ground beef: heat altered derivatives of linoleic acid. Carcinogenesis 8:1881–1887.[Abstract/Free Full Text]

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Jiang, J., L. Bjoerck, R. Fonden, and M. Emanuelson. 1996. Occurrence of conjugated cis-9 trans-11 octadecadienoic acid in bovine milk: effects of feed and dietary regimen. J. Dairy Sci. 79:438–445.[Abstract]

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