J. Anim Sci.
HOME HELP FEEDBACK SUBSCRIPTIONS ARCHIVE SEARCH TABLE OF CONTENTS
 QUICK SEARCH:   [advanced]


     


This Article
Right arrow Abstract Freely available
Right arrow Full Text (PDF)
Right arrow Alert me when this article is cited
Right arrow Alert me if a correction is posted
Services
Right arrow Similar articles in this journal
Right arrow Similar articles in PubMed
Right arrow Alert me to new issues of the journal
Right arrow Download to citation manager
Right arrow reprints & permissions
Citing Articles
Right arrow Citing Articles via HighWire
Right arrow Citing Articles via Google Scholar
Google Scholar
Right arrow Articles by Scholljegerdes, E. J.
Right arrow Articles by Rule, D. C.
Right arrow Search for Related Content
PubMed
Right arrow PubMed Citation
Right arrow Articles by Scholljegerdes, E. J.
Right arrow Articles by Rule, D. C.
J. Anim. Sci. 2004. 82:3577-3588
© 2004 American Society of Animal Science


ANIMAL NUTRITION

Influence of supplemental cracked high-linoleate or high-oleate safflower seeds on site and extent of digestion in beef cattle1

E. J. Scholljegerdes, B. W. Hess2, G. E. Moss, D. L. Hixon and D. C. Rule

Department of Animal Science, University of Wyoming, Laramie 82071-3684


    Abstract
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Our objectives were to evaluate ruminal fermentation patterns, apparent ruminal biohydrogenation, and site and extent of nutrient disappearance in cattle fed supplemental cracked safflower seeds differing in 18 C fatty acid profile. Nine Angus x Gelbvieh heifers (641 ± 9.6 kg) fitted with ruminal and duodenal cannulas were used in a triplicated 3 x 3 Latin square. Cattle were fed (OM basis) 9.1 kg of bromegrass hay and either 1) 1.8 kg of corn and 0.20 kg of soybean meal (Control); 2) 0.13 kg of soybean meal and 1.5 kg of cracked high-linoleate (67.2% 18:2) safflower seeds (Linoleate); or 3) 1.5 kg of cracked high-oleate (72.7% 18:1) safflower seeds (Oleate). Safflower seed supplements were formulated to provide similar quantities of N and TDN and 5% dietary fat. Single degree of freedom orthogonal contrasts (Control vs. Linoleate and Oleate; Linoleate vs. Oleate) were used to evaluate treatment effects. True ruminal OM and ruminal NDF disappearances (percentage of intake) were greater (P ≤0.02) for Control than Linoleate and Oleate. True ruminal N degradability (% of intake) was not different (P = 0.38) among treatments. Apparent ruminal biohydrogenation of dietary 18:2 was greatest (Linoleate vs. Oleate, P < 0.001) for Linoleate, whereas biohydrogenation of dietary 18:1 was greatest (Linoleate vs. Oleate, P = 0.02) for Oleate. Duodenal flow of 18:0 was least (P < 0.001) for Control but did not differ (P = 0.92) between Oleate and Linoleate. Total flow of unsaturated fatty acid to the duodenum was greatest (P < 0.001) in cattle fed safflower seeds, and was greater with Linoleate (P < 0.001) than with Oleate. Duodenal flow of 18:1 and 18:2 increased (P < 0.001) in Oleate and Linoleate, respectively. Duodenal flow of 18:1trans-11 was greater (P < 0.001) in cattle fed safflower seeds and in Linoleate than in Oleate. Postruminal disappearance of saturated fatty acids was greatest (P < 0.001) for Control; however, postruminal disappearance of total unsaturated fatty acids was greater (P = 0.002) for Linoleate vs. Oleate. Supplemental high-linoleate or high-oleate safflower seeds to cattle fed forage-based diets may negatively affect ruminal OM and fiber disappearance but not N disappearance. Provision of supplemental fat in the form of safflower seeds that are high in linoleic acid increased intestinal supply and postruminal disappearance of unsaturated fatty acids, indicating that the fatty acids apparently available for metabolism are affected by dietary fat source.

Key Words: Beef Cattle • Digestion • Fatty Acids • Safflower • Supplementation


    Introduction
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
There has been a resurgence of interest in feeding supplemental fat to cattle because of the benefits that dietary fat can exert on reproduction (Staples et al., 1998Go) and on the quality of ruminant-derived food products (Bauman et al., 2000Go). Our previous results (Bottger et al., 2002Go) suggested that lactating beef cows fed high-linoleate safflower seeds were more capable of maintaining body condition, whereas cows fed high-oleate safflower seeds had greater total milk fat. Cattle fed supplemental oilseeds have greater concentrations of CLA in meat (Wood et al., 1999Go) and milk (Griinari and Bauman, 1999Go), which may constitute a health advantage to consumers of these food products (Bauman et al., 2000Go). Griinari and Bauman (1999)Go reported that milk fat 18:1trans-11 is the major predictor of milk fat 18:2cis-9,trans-11 CLA concentration. Duodenal flow of 18:1trans-11 was greater for dairy cattle fed 60% forage diets supplemented with high-linoleate or high-oleate sunflower oil than for nonsupplemented controls (Kalscheur et al., 1997Go). It may be inappropriate to extrapolate results of research conducted with dairy cows to beef cows because beef cows are typically fed greater proportions of forage. Ruminal cellulolytic bacteria are primarily responsible for biohydrogenation (Harfoot and Hazelwood, 1988Go); therefore, duodenal flow of biohydrogenation products and intermediates could be very different in ruminants consuming markedly different levels of forage (Kucuk et al., 2001Go). Our hypothesis was that supplemental fat high in linoleate or oleate would alter unsaturated fatty acid supply to the small intestine with minimal effects on site and extent of digestion in cattle consuming high-forage diets. Therefore, our objectives were to evaluate site and extent of nutrient disappearance, ruminal biohydrogenation, and ruminal fermentation patterns in cattle fed bromegrass hay and supplemental high-linoleate or high-oleate cracked safflower seeds.


    Materials and Methods
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Animals
Nine Angus x Gelbvieh heifers (4 yr old; 641 ± 9.6 kg) cannulated in the rumen and proximal duodenum were used in a triplicated 3 x 3 Latin square experiment in accordance with an approved University of Wyoming Animal Care and Use Committee protocol. Heifers were housed in individual pens (2 m x 3.3 m) in a temperature-controlled room (20°C) under continuous lighting.

Diets
Heifers were fed (OM basis) 9.1 kg of chopped (2.54 cm) bromegrass hay (1.3% N, 58% NDF) and one of three supplements: 1) 1.8 kg of cracked corn and 0.20 kg of soybean meal (Control); 2) 0.13 kg of soybean meal and 1.5 kg of cracked high-linoleate (postexperiment analysis = 67.2% 18:2 and 3.0% N on an OM basis) safflower seeds (Linoleate); or 3) 1.5 kg of cracked high-oleate (postexperiment analysis = 72.7% 18:1 and 1.9% N on an OM basis) safflower seeds (Oleate). Safflower seeds were cracked with a roller mill to increase safflower seed DM availability within the rumen (Lammoglia et al., 1999Go). Dietary forage and safflower seeds were provided at the levels previously reported by our laboratory (Bottger et al., 2002Go). Dietary TDN contribution from safflower seeds was estimated using IVOMD (Bottger et al., 2002Go), assuming 177% TDN (NRC, 1996Go) for the fat within the safflower seeds. The Control supplement was provided at a level that should have provided quantities of dietary TDN and N similar to that of the safflower seed supplements (Table 1Go). Half the respective diets were fed twice daily at 0600 and 1800. Heifers had ad libitum access to fresh water and trace mineral salt (Champions Choice, Akzo Salt Inc., Georgetown, SC; guaranteed analysis [percentage of DM]: NaCl, 95 to 99; Co, Cu, I, Mn, Zn, and Fe, less than 1).


View this table:
[in this window]
[in a new window]
 
Table 1. Ingredient and analyzed composition of diets fed to heifersa
 
Sampling
As a marker of digesta flow, boluses (No. 10 lock ring gelatin capsules, Torpac Inc, Fairfield, NJ) containing 5 g of Cr2O3 were dosed intraruminally at each feeding. Each 17-d experimental period included 14 d of adaptation to the new dietary treatment. Beginning at 0400 on d 15 of each sampling period, duodenal (200 mL) and fecal (50 mL) samples were taken every 4 h. On d 16, collection times were advanced 2 h so that samples were collected to represent every 2 h in a 24-h period. Fecal samples were dried in a 55°C forced-air oven, ground (Wiley mill, 1-mm screen, Thomas Hill and Sons, Philadelphia, PA), and composited (equal dry weight basis) within heifer for each period. Duodenal digesta samples were composited (equal volume) within heifer for each period. Duodenal digesta samples were then lyophilized (Genesis SQ 25 Super ES freeze dryer, The VirTis Co., Gardiner, NY) and ground (Wiley mill, 1-mm screen, Thomas Hill and Sons).

Immediately before the 0600 feeding on d 17 of each period (0 h), approximately 200 mL of whole ruminal contents was collected from the center of the ruminal mat (dorsal to the cranial pillar). Ruminal samples were then collected at 3, 6, 9, 12, 15, 18, and 21 h. Ruminal pH was immediately measured on whole ruminal contents using a combination electrode (Orion Research Inc., Boston, MA), and 10 mL of ruminal fluid was strained through eight layers of cheesecloth. The resulting ruminal fluid was acidified with 0.1 mL of 7.2 N H2SO4 and immediately frozen. The remaining unstrained sample of whole ruminal contents was placed in a blender (Hamilton Beach/Proctor Silex, Washington, NC) with an equal volume of 0.9% NaCl (wt/vol) solution and homogenized for 1.0 min to dislodge particulate-associated bacteria. The homogenized sample was then strained through eight layers of cheesecloth for subsequent bacterial isolation by differential centrifugation (Merchen and Satter, 1983Go). The resulting bacterial isolate was lyophilized (Genesis SQ 25 Super ES freeze dryer, The VirTis Co.) and ground with a mortar and pestle for subsequent laboratory analyses.

Laboratory Analyses
Feed, duodenal, microbial, and fecal samples were analyzed for DM and ash (AOAC, 1990Go). Nitrogen content of feed, microbial, duodenal, and fecal samples was determined using a Leco FP-528 N analyzer (Leco Corp., Henderson, NV). Neutral detergent fiber content of feed, feces, and duodenal digesta was determined using an Ankom 200 fiber analyzer (Ankom Technology, Fairport, NY). Chromium concentration in duodenal digesta and feces was determined (Hill and Anderson, 1958Go) by atomic absorption spectrophotometry (model 210 VGP AASpectr., Buck Scientific, E. Norwalk, CT) with an air-plus-acetylene flame.

Acidified ruminal fluid samples were centrifuged at 10,000 x g for 10 min, and a 2.5-mL aliquot of the resulting supernatant was added to 0.5 mL of 25% metaphosphoric acid containing 2 g/L of 2-ethyl-butyric acid (Goetsch and Galyean, 1983Go). These samples were then centrifuged for 10 min at 10,000 x g, and the supernatant fluid was analyzed for concentrations of VFA using a Hewlett-Packard 5890 GLC (Hewlett-Packard, Avondale, PA) equipped with a 15 m x 0.533 mm (i.d.) column (Nukol, Supelco, Bellefonte, PA) with a ramp temperature of 110 to 150°C at 8°C/min. Helium was used as the carrier gas with a column flow rate of 20 mL/min. Injector and detector temperatures were 250°C. Ruminal NH3 concentration was determined by the phenol- hypochlorite procedure (Broderick and Kang, 1980Go).

Feed, duodenal digesta, and fecal samples were subjected to direct transesterification by incubating samples in a 16-mm x 125-mm screw-cap tubes with 2.0 mL of HCl in methanol (1.09 M HCl; 9.12 mL of HCl/100 mL of methanol) and 2 mL of methanol including 1 mg of tridecanoic acid methyl ester (Sigma, St. Louis, MO), as an internal standard, at 85°C for 1 h. Fatty acid methyl esters were extracted in 2.0 mL of hexane and transferred to GC vials containing a 1-mm bed of anhydrous sodium sulfate. Weight percentages of CLA can be underestimated by using an acid catalyst to prepare fatty acid methylesters (Kramer et al., 1997Go). However, fatty acids in duodenal contents of ruminants are predominately in the nonesterified form because of the extensive fatty acid hydrolysis occurring in the rumen. Alkaline catalysts do not react with nonesterified fatty acids (Hammond, 1993Go); thus, use of acid catalyst is appropriately required. Moreover, treatment effects remained intact when acid and alkaline catalysts were compared (Murrieta et al., 2003Go). Separation of fatty acid methyl esters was achieved by GLC (model 5890 series II, Hewlett-Packard) with a 100-m capillary column (SP-2560, Supelco) and He as a carrier gas at 0.5 mL/min. Oven temperature was maintained at 175°C for 40 min and then ramped to 240°C at 10°C/min to 240°C. Injector and detector temperature was 250°C. Identification of peaks was accomplished using purified standards (Nu-Chek Prep, Elysian, MN; Matreya, Pleasant Gap, PA).

Calculations and Statistical Analyses
Nutrient flows and digestibilities were estimated as previously described by Kucuk et al. (2001)Go and Scholljegerdes et al. (2004)Go. Biohydrogenation of unsaturated fatty acids (C18) was calculated using the equation of Tice et al. (1994)Go. Additionally, the following equations were used to evaluate the percentages of biohydrogenation intermediates at the duodenum:



where D = duodenal flow of the respective fatty acid, and I = intake of the respective fatty acid.

Single degree of freedom orthogonal contrasts were used to compare effects of Control vs. supplemental safflower seeds (Lineolate and Oleate), as well as Linoleate vs. Oleate. All data were analyzed using the Mixed procedure of SAS (SAS Inst., Inc., Cary, NC). The model included animal as the random variable. Additionally, the model used for analysis of time course data included effects of animal, period, treatment, time, and treatment x time. Animal x period x treatment was used to specify variation among animals using the Random statement. Autoregression order one was determined to be the most desirable covariance structure according to the Akaike’s information criterion. No treatment x sampling hour interactions (P = 0.11 to 0.99) were detected for time course data; therefore, only the main effects were reported.


    Results and Discussion
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
OM Intake and Disappearance
Total OM intake differed (P < 0.001) across all treatments (data not shown) due to the level at which supplement was fed to equalize intake of TDN (Table 2Go). Decreased duodenal OM flow (P = 0.02) for Control reflected the improved (P = 0.01) true ruminal OM disappearance observed for this treatment because microbial OM flow to the duodenum was not affected (P = 0.19 to 0.21) by treatment. Whitney et al. (2000)Go demonstrated that as dietary soybean oil level increased from 3% to 6%, IVDMD decreased linearly (P < 0.001) compared with a corn–soybean meal-based control, which was similar to the Control supplement fed in the present experiment. Expressing duodenal OM flow on a fatty acid free basis did not account (P = 0.40) for the increased duodenal OM flow (data not shown) for the safflower seed supplements, indicating that the OM of the corn-based Control supplement was more digestible than the supplements containing safflower seeds. Post-ruminal OM disappearance (percentage of intake) did not differ (P = 0.72 to 0.95) across treatments. Cattle fed Control had lower (P < 0.001) fecal OM output, reflecting the greater (P < 0.001) total-tract OM disappearance for this treatment. No differences (P = 0.40 to 0.68) were noted between cattle fed Linoleate and Oleate for fecal OM output or total-tract OM disappearance. In contrast to our results, supplementing lower levels of soybean oil to beef heifers (0.03% of BW) grazing bromegrass pasture did not negatively influence ruminal, postruminal, or total-tract OM digestibility (Brokaw et al., 2001Go). Aldrich et al. (1997)Go reported no differences in ruminal and postruminal OM disappearance, but observed greater digestion of OM in the total tract for nonfat-supplemented steers compared with steers supplemented with whole canola seeds. When supplemental soybean oil was given via ruminal or duodenal infusion to beef heifers consuming grass hay (Krysl et al., 1991Go), true ruminal OM digestibility was not influenced; however, total-tract OM digestibility was lower for the ruminally infused treatments compared with the control and duodenally infused treatments. However, relatively low ruminal OM digestibilities coupled with a relatively large standard error may have precluded detection of a significant difference between treatments in the study by Krysl et al. (1991)Go. Our results agree with those of Elliot et al. (1997)Go, who demonstrated a decrease in ruminal OM digestibility and no difference in postruminal OM digestibility when tallow was supplemented to beef steers consuming a 60% forage diet; however, as saturation and esterification increased, there was a linear decrease in total tract OM digestibility. Thus, decreased total-tract OM disappearance for cattle receiving supplemental lipids may be attributed to decreased ruminal OM disappearance.


View this table:
[in this window]
[in a new window]
 
Table 2. Influence of supplemental high-linoleate or high-oleate safflower seeds on organic matter intake, duodenal flow, and disappearance by heifers consuming bromegrass hay
 
Nitrogen Intake and Disappearance
Despite supplements initially formulated to provide similar quantities of dietary N, N intake by cattle fed the high-linoleate safflower seeds was greater (P < 0.001) because the N content of the seeds throughout the trial were higher than originally estimated from pretrial laboratory analysis (Table 3Go). Nonetheless, total, microbial, and nonmicrobial N flow to the duodenum did not differ (P = 0.27 to 1.0) across treatments. These results are similar to those reported by Brokaw et al. (2001)Go. True ruminal N degraded (g/d) tended (P = 0.12) to be greater for Linoleate and Oleate compared with Control, and tended (P = 0.13) to be greater for Linoleate than for Oleate. Using the average ruminal N degradability value for bromegrass hay reported by Scholljegerdes et al. (2004)Go in addition to average ruminally degraded protein values of the NRC (2001)Go for cracked corn, soybean meal, and safflower seed meal (assuming ruminally degraded protein for whole seeds is equal to the seed meal), we estimated that ruminally degraded N would have ranked Linoleate (+ 16 g/d) >Oleate (+ 5 g/d) >Control. A similar ranking (Linoleate = 46 g, Oleate = 12 g, and Control = –80 g) was noted for postexperimental evaluations using the NRC (1996)Go, with microbial efficiency adjusted to 10% of TDN (Lardy et al., 2004Go). Although the Control diet seemed to be deficient in ruminally degraded protein, estimates of N recycling according to NRC (1985)Go equations indicated that N recycled to the rumen should have been more than sufficient (approximately 65 g) to maintain proper ruminal N status. Postruminal N disappearance did not differ (P = 0.22 to 0.34) across treatments; however, total-tract N disappearance was greater (P < 0.001) for fat-supplemented heifers, as well as between fat supplements with Linoleate being greater (P = 0.04) than Oleate. Brokaw et al. (2001)Go reported that ruminal and total-tract N disappearance did not differ, but lower-tract N disappearance (percentage of intake) decreased when grazing beef heifers were given cracked corn or soybean oil. Microbial efficiency did not differ (P = 0.15 to 0.92) among treatments, which agrees with other research (Murphy et al., 1987Go; Kalscheur et al., 1997Go; Brokaw et al., 2001Go) on the influence of supplemental fat on site and extent of digestion. The numerical increase in microbial efficiency for cattle fed supplemental fat in our experiment was also similar in magnitude to the nonsignificant increase previously reported by Krysl et al. (1991)Go.


View this table:
[in this window]
[in a new window]
 
Table 3. Influence of supplemental high-linoleate or high-oleate safflower seeds on nitrogen intake, duodenal flow, ruminal degradation, and apparent disappearance by heifers consuming bromegrass hay
 
NDF Intake and Disappearance
Neutral detergent fiber intake was greatest (P < 0.001) by cattle fed safflower seeds (Table 4Go) as a result of differences in the NDF content of the supplements, or specifically, the seed coat on the safflower seeds. Whitney et al. (2000)Go noted that 6% added soybean oil had deleterious effects on digestibility of a forage-based diet, whereas 3% dietary soybean oil may have been optimal for maintaining digestion. Our laboratory also has reported that feeding soybean oil at 1.74% of the total diet did not affect NDF digestibility by heifers grazing bromegrass pastures (Brokaw et al., 2001Go). However, in the present experiment, ruminal NDF disappearance was decreased (P = 0.02) with the provision of cracked safflower seeds, and no difference (P = 0.78) was noted between safflower seed types. Murphy et al. (1987)Go indicated that decreased ruminal availability of the lipid in supplemental oilseeds could be an effective way to circumvent the negative effects fat exert on fiber digestion. However, if oil seeds are cracked, thereby increasing ruminal availability (Lammoglia et al., 1999Go), negative effects on ruminal NDF digestibility are more likely if fed above critical levels (Aldrich et al., 1997Go). In addition to providing supplemental oilseeds at a level that was slightly greater than normally assumed not to negatively influence ruminal fermentation (Jenkins, 1993Go), the combination of cracking the oilseeds to increase ruminal availability (Lammoglia et al., 1999Go) and the relatively indigestible oilseed hull (Aldrich et al., 1997Go) may have had additive negative effects on ruminal NDF digestibility. No differences (P = 0.55 to 0.91) were noted among treatments for postruminal NDF disappearance. Likewise, Brokaw et al. (2001)Go reported that the concentration of supplemental soybean oil to beef heifers grazing bromegrass pasture did not influence postruminal NDF digestibility (percentage of intake). Due to the greater ruminal NDF disappearance for Control without an effect on postruminal NDF disappearance, total-tract NDF disappearance was greater (P = 0.003) for Control than for the safflower seed treatments.


View this table:
[in this window]
[in a new window]
 
Table 4. Influence of supplemental high-linoleate or high-oleate safflower seeds on neutral detergent fiber intake, duodenal flow, and disappearance by heifers consuming brome-grass hay
 
Ruminal pH, NH3N, and VFA
Ruminal pH was greater (P = 0.05) for Linoleate than for Oleate, but pH was not different (P = 0.79) between Control and safflower-supplemented animals (Table 5Go). Nonetheless, pH values for all treatments were within the range considered acceptable for fiber digestion (Ørskov and Ryle, 1990Go). Ruminal ammonia concentrations did not differ (P = 0.23 to 0.32) across all treatments ranging from 4.1 to 5.5 mM, and were above the range (0.60 to 1.59 mM NH3) considered by Satter and Slyter (1974)Go to be sufficient for microbial N production.


View this table:
[in this window]
[in a new window]
 
Table 5. Ruminal pH, NH3, and VFA in heifers consuming bromegrass hay and high-oleate or high-linoleate safflower seeds
 
Diet did not affect (P = 0.90 to 0.95) ruminal concentrations of total VFA. Even though ruminal NDF digestion was greater for Control than safflower seed treatments, ruminal molar proportions of acetate did not differ (P = 0.63 to 0.80) across treatments. However, ruminal molar proportions of butyrate were lower (P < 0.001) for the diets supplemented with lipids. The higher proportion of butyrate for Control may have contributed to the lack of differences observed for acetate because butyrate has the potential to be formed from acetate (Van Soest, 1994Go). Also, ruminal fermentation of corn can result in greater ruminal molar proportions of butyrate (Hess et al., 1996Go). Ruminal molar proportions for all of the VFA reported in Table 5Go did not differ between Linoleate and Oleate safflower seed treatments. This was expected because fatty acid composition does not influence VFA production (Palmquist, 1991Go). Ruminal molar proportions of propionate were greater (P = 0.002) for cattle fed safflower seeds than Control. Krysl et al. (1991)Go attributed greater ruminal molar proportion of propionate in beef heifers infused with soybean oil in the rumen to decreased fiber digestion; however, Chalupa et al. (1986)Go suggested that increased molar proportions of propionate may be attributed to bacterial metabolism of glycerol. Supplemental lipids have also decreased methanogenesis (Jounay, 1994Go), which will increase proportions of propionate (Van Soest, 1994Go).

The acetate:propionate ratio reflected the observed difference in ruminal NDF disappearance, in that the ratio tended (P = 0.06) to be greatest for Control compared with Linoleate and Oleate. A decrease in the acetate:propionate ratio with supplemental fat has been noted by several researchers (Jenkins et al., 1989Go; Palmquist, 1991Go; Whitney et al., 2000Go). Ruminal molar proportions of isobutyrate and valerate were not different (P = 0.39 to 0.96) among all treatments. The trend for increased (P = 0.11) ruminal molar proportions of isovalerate for the Control treatment likely had little biological importance (Gunter et al., 1990Go).

Fatty Acid Intake and Disappearance
Fatty acid intake was greater (P < 0.001) by heifers fed safflower seeds, and heifers fed Oleate consumed 4.9 g/d more (P < 0.001) fatty acid than heifers fed Linoleate (Table 6Go). Intake of 18:1 was greatest (P < 0.001) for Oleate, and intake of 18:2 was greatest (P < 0.001) for Linoleate, reflecting differences in fatty acid composition of the safflower seeds (Table 1Go). The primary source of dietary 18:3 in the diets used for this experiment originated from the forage; however, there were minor differences in 18:3 content between the three supplements. Forage intake was held constant across treatment, and the minor differences noted for duodenal 18:3 caused the treatments to differ (P < 0.001), but this difference was small enough that the biological significance (46.0 to 46.7 g/d) was questionable.


View this table:
[in this window]
[in a new window]
 
Table 6. Long-chain fatty acid intake by heifers consuming bromegrass hay and high-oleate or high-linoleate safflower seeds
 
Dietary lipids undergo rapid and extensive hydrolysis in the rumen to form glycerol and unesterified fatty acids (Jenkins, 1993Go). Accumulation of unsaturated fatty acids in the rumen can be toxic to ruminal bacteria. Consequently, ruminal bacteria hydrogenate double bonds of unsaturated fatty acids in an effort to alleviate toxicity (Harfoot and Hazelwood, 1988Go). Thus, ruminal biohydrogenation of dietary unsaturated C18 fatty acids expressed as a percentage of intake was greater (P = 0.007) in heifers fed safflower seeds (Table 7Go). Biohydrogenation of dietary 18:3 tended (P = 0.07) to differ between Control and lipid-supplemented heifers, but did not differ (P = 0.68) between the Linoleate and Oleate treatments. Additionally, ruminal biohydrogenation of dietary unsaturated C18 fatty acids was higher (P < 0.001) for Linoleate than Oleate. Ruminal biohydrogenation of dietary 18:1 was greater (P < 0.001) for diets with safflower seed than Control, and Oleate was greater (P = 0.02) than Linoleate. Kalscheur et al. (1997)Go also noted increased biohydrogenation of dietary 18:1 in dairy cattle fed either high-oleate or high-linoleate sunflower oil. Biohydrogenation of 18:2 is initiated by the isomerization of the cis-12 double bond, which produces cis-9,trans-11 CLA (Harfoot and Hazelwood, 1988Go). The cis-9 double bond of the conjugated diene is then hydrogenated resulting in formation of 18:1trans-11. Complete biohydrogenation of dietary unsaturated C18 fatty acid results in ruminal accumulation of 18:0; however, Harfoot et al. (1973)Go reported that increased ruminal concentrations of 18:2 increased the extent of biohydrogenation of 18:2. Furthermore, Bauchart et al. (1990)Go showed that as linoleic acid intake increased (4.1 to 46.6 g/d) biohydrogenation of 18:2 concomitantly increased (55 to 91%, respectively). Kalscheur et al. (1997)Go also showed an increase in biohydrogenation of 18:2 when dairy cows where fed high-linoleate sun-flower oil. Harfoot et al. (1973)Go stated that ruminal levels of 18:2 is the predominant factor affecting biohydrogenation of 18:2, and as intake of 18:2 increases, the accumulation of biohydrogenation intermediates increase with time. In our study, the proportion of biohydrogenation intermediates reaching the small intestine was greater (P ≤0.002) for Linoleate than Oleate. Our results are comparable to those of Gould (2000)Go, who noted more extensive ruminal biohydrogenation and a concomitant increase in production of biohydrogenation intermediates in lambs fed high-forage diets with 6% supplemental soybean oil.


View this table:
[in this window]
[in a new window]
 
Table 7. Ruminal biohydrogenation of long-chain fatty acids in heifers consuming bromegrass hay and high-oleate or high-linoleate safflower seeds
 
Duodenal flow of 18:0 was greatest (P < 0.001) for the safflower seed treatments, whereas type of supplemental safflower seed had no influence (P = 0.92) on duodenal flow of 18:0. Although ruminal biohydrogenation of dietary 18:1 and 18:2 was more extensive for Oleate and Linoleate, duodenal flow of 18:1 and 18:2 was greatest (P < 0.001) for Oleate and Linoleate, respectively. Duodenal flow of 18:1trans-11 was three- to sixfold greater (P < 0.001) for cattle fed safflower seeds than Control, with heifers fed Linoleate having twice (P <0.001) the duodenal flow of 18:1trans-11 than Oleate heifers. Kalscheur et al. (1997)Go suggested that 18:1 should not be a source of 18:1trans-11; however, when these researchers evaluated 18:1trans-11 flow to the small intestine in dairy cattle fed high-linoleate or high-oleate sunflower oil, they observed similar duodenal flow of 18:1trans-11 for both treatments. Kalscheur et al. (1997)Go attributed this result to sufficient amounts of PUFA in the high-oleate sunflower oil diet to provide adequate precursors for the production of 18:1trans-11. In the current study, this was not the case because duodenal flow of 18:1trans-11 with Linoleate was twice that with Oleate supplements. Kemp et al. (1975)Go indicated that a minor cellulolytic bacterial species, Fusocillus spp., can isomerize 18:1cis-9 to 18:1trans-11. There may have been sufficient Fusocillus spp. present in the rumen of cattle fed Oleate to promote the accumulation of significant quantities of 18:1trans-11. Increased duodenal flow of 18:1trans-11 for both safflower treatments is noteworthy because 18:1trans-11 is the predominant precursor for 18:2cis-9,trans-11 production within animal tissues, including the mammary gland (Bauman et al., 2000Go). Duodenal flow of 18:2cis-9,trans-11 tended (P = 0.06) to be higher in heifers supplemented with oilseeds compared with Control. In contrast, heifers fed Control had greater (P = 0.03) 18:2trans-10,cis-12 flowing to the duodenum than did heifers fed the oil-seeds. Greater duodenal flow of 18:2trans-10,cis-12 for Control may be related to this CLA isomer, constituting a greater proportion of the CLA isomers when cereal grains are included in the diet (Kucuk et al., 2001Go). Compared with Control, total saturated and unsaturated fatty acid flow to the duodenum was greater (P < 0.001) in heifers fed safflower seeds; duodenal flow of these fatty acids did not differ (P = 0.44) between Linoleate and Oleate (Table 8Go). Kalscheur et al. (1997)Go noted that duodenal fatty acid flow in dairy cattle increased equally whether diets were supplemented with high-linoleate or high-oleate sunflower oil. Greater flow of fatty acid to the duodenum in cattle fed oilseeds was expected because negligible amounts of dietary long-chain fatty acid disappear from the rumen (Jenkins, 1993Go).


View this table:
[in this window]
[in a new window]
 
Table 8. Duodenal long-chain fatty acid flow in heifers consuming bromegrass hay and high-oleate or high-linoleate safflower seeds
 
Postruminal disappearance of saturated fatty acid was greatest (P < 0.001) for Control (Table 9Go). Postruminal disappearance of monounsaturated fatty acids and PUFA did not differ (P = 0.60 to 0.79) between Control and heifers fed safflower seeds, and the Linoleate treatment had greater postruminal disappearance of mono-unsaturated fatty acids and PUFA (P ≤0.001) than the Oleate treatment. Kalscheur et al. (1997)Go noted that when lactating dairy cows were fed either high-linoleate or high-oleate sunflower oil, postruminal digestibility of individual and total fatty acids did not differ. Postruminal 18:1trans-11 disappearance was not different (P = 0.48 to P = 0.54) across all treatments and averaged 92 ± 1.0%, indicating that 18:1trans-11 is readily digested. Efforts to increase the flow of 18:1trans-11 could be a useful way to increase 18:2cis-9,trans-11 content within ruminant tissues because 18:1trans-11 is the predominant precursor for 18:2cis-9,trans-11 production within adipose and mammary tissues (Bauman et al., 2000Go). Although postruminal disappearance of 18:1trans-11 did not differ across dietary treatments, heifers fed high-linoleate safflower seeds would have had the greatest quantity of 18:1trans-11 available for metabolism because intestinal supply of this fatty acid was greatest in these animals. Postruminal disappearance of total fatty acid was not influenced (P = 0.98) by supplementing safflower seeds but was greater (P = 0.05) for Linoleate than for Oleate. This may have been due to a greater proportion of the total fatty acids presented to the small intestine being comprised of PUFA for the Linoleate treatment compared with the Oleate treatment. Andrews and Lewis (1970)Go indicated that unsaturated fatty acids are more digestible than saturated fatty acids.


View this table:
[in this window]
[in a new window]
 
Table 9. Postruminal long-chain fatty acid disappearance in heifers consuming bromegrass hay and high-oleate or high-linoleate safflower seeds
 
Our results for postruminal disappearance are comparable to those of Wu et al. (1991)Go, who observed decreased postruminal digestibility of saturated fatty acid and total fatty acid in dairy cows fed 60% forage diets with 6% fat from an animal-vegetable blend. Andrews and Lewis (1970)Go suggested that postruminal digestibility of unsaturated fatty acids is greater than saturated fatty acids because unsaturated fatty acids are more hydrophilic and more easily form micelles, and therefore are more readily absorbed. Previous research conducted in our laboratory (Gould, 2000Go; Kucuk et al., 2001Go) indicated that the CLA isomers are between 90 and 100% digestible in the small intestine of wethers fed a forage-based diet and supplemental soybean oil. Caution must be exercised, however, when interpreting results of apparent postruminal fatty acid disappearance because of the contribution of endogenous fatty acids (Kucuk et al., 2001Go).

Overall, supplementing beef heifers with high-fat safflower seeds can boost the quantity of saturated and unsaturated fatty acids that reach the small intestine. However, with regard to overall energy available at the small intestine, increased postruminal disappearance of fat did not compensate for the differences in intake and ruminal OM and NDF disappearance between treatments. We estimated dietary TDN as the sum of digestible fatty acid free OM and digestible fatty acids times 2.25. Our estimates of total TDN intake using results from this experiment indicated that the Control diet was greater (P < 0.001) than the diets supplemented with fat, and Linoleate tended (P = 0.12) to be greater than Oleate (7,923, 7,199, and 6,846 g/d, respectively). Preexperiment estimates of total TDN intake were 6,858, 6,570, and 6,581 g/d for Control, Linoleate, and Oleate, respectively. Comparisons of preexperiment dietary TDN to our estimates using the experimental results revealed that the deviation in expected vs. observed dietary TDN was within the normal range reported in the literature (Moore et al., 1999Go). Moreover, we anticipated TDN intake to be slightly greater for Control than for the oilseed diets; however, the TDN value of our diets containing safflower seeds may have been decreased because of the unexpected decrease in ruminal fiber and OM disappearance, which resulted in decreased disappearance of OM from the total tract for these treatments. Alternatively, the greater than expected magnitude of difference in TDN intake between the Control and the safflower seed diets could be due to positive associative effects for the Control treatment. Positive associative effects have been noted in many studies where researchers have fed grain-based supplements at less than 0.3% of BW (Gunter, 1996Go; Caton and Dhuyvetter, 1997Go). Corn was fed to the Control heifers at 0.28% of BW in the current experiment. Decreased postruminal disappearance of fatty acids for Oleate also resulted in a decrease in estimated total TDN intake compared with the Linoleate treatment. Thus, it may be erroneous to ascribe the same TDN value to all varieties of safflower seeds.

As humans become more cognizant of health issues and demand meat that is healthier, the development of ruminant-derived food products as functional foods will be an important goal for the livestock industry. Additionally, certain fatty acids have been shown to modulate various immune and inflammatory responses (Hwang, 2000Go), as well as contribute to improved reproductive success (Staples et al., 1998Go). Therefore, it is becoming more important to understand ruminal metabolism of dietary fatty acids, which will allow nutritionists to manipulate the diet to optimize intestinal supply of specific fatty acids that may have beneficial attributes. Heifers consuming safflower seeds high in linoleic acid compared with safflower seeds high in oleic acid had greater ruminal biohydrogenation of 18C fatty acids and, therefore, had greater duodenal flow of 18:1trans-11. We also observed an increase in duodenal flow of the 18:2cis-9,trans-11 isomer in cattle fed safflower seeds; however, duodenal flow of 18:2trans-10,cis-12, the isomer implicated to decrease fat deposition (Park et al., 1999Go), was not detected in heifers fed Oleate. In an experiment conducted by Bolte et al. (2002)Go, lambs fed high-linoleate safflower seeds had higher muscle and adipose tissue levels of 18:1trans-11 and 18:2cis-9,trans-11 compared with lambs fed high-oleate safflower seeds or a low-fat control. The CLA status of the consumer is increased because 18:1trans-11 can be converted to 18:2cis-9,trans-11 by {Delta}9 desaturase (Turpeinen et al., 2002Go). Therefore, consumers of beef produced from cattle fed safflower seeds in diets comparable to those reported herein would be expected to have increased CLA status. Results of the current study are also suggestive of increased 18:1trans-11 and 18:2cis-9,trans-11 available to the calf from the milk of lactating beef cow. Dairy cows fed high-oleate or high-linoleate sunflower seeds had increased milk output of both of these fatty acids (Kalscheur et al., 1997Go). The beneficial effects of CLA observed in laboratory animals (McGuire and McGuire, 2000Go) may occur in beef calves as well. Thus, determining the effects of oilseed supplementation on fatty acid output via the mammary gland of beef cows and subsequent fatty acid status of the nursing calf would be of great importance.


    Implications
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Despite more extensive ruminal biohydrogenation, postruminal supply of unsaturated fatty acids was enhanced by the provision of supplemental safflower seeds. Differences in postruminal fatty acid disappearance attributable to diet indicate that fatty acids apparently available for metabolism are affected by dietary fat source. Thus, supplementing oilseeds with varying fatty acid compositions to beef cattle consuming forage-based diets may be an effective means of altering tissue and milk fatty acid composition. However, consideration must also be given to livestock production in response to altering dietary energy through the provision of oilseeds with varying fatty acid composition.


    Footnotes
 
1 Research was supported by Grant No. 99-02390 from USDA-NRI. Back

2 Correspondence: P.O. Box 3684 (phone: 307-766-5173; fax: 307-766-2355; e-mail: brethess{at}uwyo.edu).

Received for publication April 15, 2004. Accepted for publication August 19, 2004.


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


Aldrich, C. G., N. R. Merchen, J. K. Drackley, S. S. Gonzalez, G. C. Fahey, Jr., and L. L. Berger. 1997. The effects of chemical treatment of whole canola seed on lipid and protein digestion by steers. J. Anim. Sci. 75:502–511.[Abstract/Free Full Text]

Andrews, R. J., and D. Lewis. 1970. The utilization of dietary fat by ruminants. I. The digestibility of some commercially available fats. J. Agric. Sci. 75:47–54.

AOAC. 1990. Official Methods of Analysis. 15th ed. Assoc. Off. Anal. Chem., Arlington, VA.

Bauchart, D., F. Legay-Carmier, and M. Doreau. 1990. Relationship between intake and duodenal flows of linoleic acid in dairy cows fed lipid-supplemented diets. Reprod. Nutr. Dev. 30(Suppl. 2):188. (Abstr.)

Bauman, D. E., L. H. Baumgard, B. A. Corl, and J. M. Griinari. 2000. Biosynthesis of conjugated linoleic acid in ruminants. Proc. Amer. Soc. Anim. Sci., 1999. Available: http://www.asas.org/jas/symposia/proceedings/0937.pdf. Accessed March 27, 2001.

Bolte, M. R., B. W. Hess, W. J. Means, G. E. Moss, and D. C. Rule. 2002. Feeding lambs high-oleate or high-linoleate safflower seeds differentially influences carcass fatty acid composition. J. Anim. Sci. 80:609–616.[Abstract/Free Full Text]

Bottger, J. D., B. W. Hess, B. M. Alexander, D. L. Hixon, L. F. Woodard, R. N. Funston, D. M. Hallford, and G. E. Moss. 2002. Effects of supplementation with high linoleic or oleic cracked safflower seeds on postpartum reproduction and calf performance of primiparous beef heifers. J. Anim. Sci. 80:2023–2030.[Abstract/Free Full Text]

Broderick, G. A., and J. H. Kang. 1980. Automated simultaneous determinations of ammonia and total amino acids in ruminal fluid and in vitro media. J. Dairy Sci. 63:64–75.

Brokaw, L., B. W. Hess, and D. C. Rule. 2001. Supplemental soybean oil or corn for beef heifers grazing summer pasture: Effects on forage intake, ruminal fermentation, and site and extent of digestion. J. Anim. Sci. 79:2704–2712.[Abstract/Free Full Text]

Caton, J. S., and D. V. Dhuyvetter. 1997. Influence of energy supplements on grazing ruminants: Requirements and responses. J. Anim. Sci. 75:533–542.[Abstract/Free Full Text]

Chalupa, W., B. Vecchiatelli, A. Esler, D. S. Kronfeld, D. Sklan, and D. L. Palmquist. 1986. Ruminal fermentation in vivo as influenced by long-chain fatty acids. J. Dairy Sci. 69:1293–1301.

Elliot, J. P., J. K. Drackley, C. G. Aldrich, and N. R. Merchen. 1997. Effects of saturation and esterification of fat sources on site and extent of digestion in steers: Ruminal fermentation and digestion of organic matter, fiber, and nitrogen. J. Anim. Sci. 75:2803–2812.[Abstract/Free Full Text]

Goetsch, A. L., and M. L. Galyean. 1983. Influence of feeding frequency on passage of fluid and particulate markers in steers fed a concentrate diet. Can. J. Anim. Sci. 63:727–730.

Gould, J. 2000. Supplemental ethanol for ruminants consuming forage-based diets. M.S. Thesis, Univ. of Wyoming, Laramie.

Griinari, J. M., and D. E. Bauman. 1999. Biosynthesis of conjugated linoleic acid and its incorporation into meat and milk in ruminants. Pages 180–200 in Advances in Conjugated Linoleic Acid Research. M. P. Yurawecz, M. M. Mossoba, J. K., G. Kramer, G. Nelson, and M. W. Pariza, ed. AOCS Press, Champaign, IL.

Gunter, S. A. 1996. Comparison of forage quality and cattle performance with different supplementation programs. Proc. 3rd Grazing Livest. Nutr. Conf. Proc. West. Sec. Amer. Soc. Anim. Sci. 47(Suppl. 1):30–39.

Gunter, S. A., L. J. Krysl, M. B. Judkins, J. T. Broesder, and R. K. Barton. 1990. Influence of branched-chain fatty acid supplementation on voluntary intake, site and extent of digestion, ruminal fermentation, digesta kinetics and microbial protein synthesis in beef heifers consuming grass hay. J. Anim. Sci. 68:2885–2892.[Abstract]

Hammond, E. W. 1993. Gas liquid chromatography. Pages 65–111 in Chromatography for Analysis of Lipids. E. W. Hammond, ed. CRC Press, Boca Raton, FL.

Harfoot, C. G., and G. P. Hazelwood. 1988. Lipid metabolism in the rumen. Pages 285–322 in The Rumen Microbial Ecosystem. P. N. Hobson, ed. Elsevier Applied Science, New York, NY.

Harfoot, C. G., R. C. Nobel, and J. H. Moore. 1973. Factors influencing the extent of biohydrogenation of linoleic acid by rumen micro-organisms in vitro. J. Sci. Food Agric. 24:961–970.[Medline]

Hess, B. W., L. J. Krysl, M. B. Judkins, D. W. Holcombe, J. D. Hess, D. R. Hanks, and S. A. Huber. 1996. Supplemental cracked corn or wheat bran for steers grazing endophytefree fescue pasture: Effects on live weight gain, nutrient quality, forage intake, particulate and fluid kinetics, ruminal fermentation, and digestion. J. Anim. Sci. 74:1116–1125.[Abstract]

Hill, F. W., and D. L. Anderson. 1958. Comparison of metabolizable energy and productive energy determinations with growing chicks. J. Nutr. 64:1405–1415.

Hwang, D. 2000. Fatty acids and immune responses: A new perspective in searching for clues to mechanism. Annu. Rev. Nutr. 20:431–456.[Medline]

Jenkins, T. C. 1993. Lipid metabolism in the rumen. J. Dairy Sci. 76:3851–3863.[Abstract/Free Full Text]

Jenkins, T. C., T. Gimenez, and D. L. Cross. 1989. Influence of phospholipids on ruminal fermentation in vitro and on nutrient digestion and serum lipids in sheep. J. Anim. Sci. 67:529–537.

Jounay, J. P. 1994. Manipulation of microbial activity in the rumen. Arch. Anim. Nutr. 46:133–153.

Kalscheur, K. F., B. B. Teter, L. S. Piperova, and R. A. Erdman. 1997. Effect of fat source on duodenal flow of trans-C18:1 fatty acids and milk fat production in dairy cows. J. Dairy Sci. 80:2115–2126.[Abstract]

Kemp, P., R. W. White, and D. J. Lander. 1975. The hydrogenation of unsaturated fatty acids by five bacterial isolates from the sheep rumen, including new species. J. Gen. Microb. 90:100–114.[Medline]

Kramer, J. K. G., V. Fellner, M. E. R. Dugan, F. D. Sauer, M. M. Massoba, and M. P. Yurawecz. 1997. Evaluating acid and base catalysts via the methylation of milk and rumen fatty acids with special emphasis on conjugated dienes and total trans fatty acids. Lipids 32:1219–1228.[Medline]

Krysl, L. J., M. B. Judkins, and V. R. Bohman. 1991. Influence of ruminal or duodenal soybean oil infusion on intake, ruminal fermentation, site and extent of digestion, and microbial protein synthesis in beef heifers consuming grass hay. J. Anim. Sci. 69:2585–2590.[Abstract]

Kucuk, O., B. W. Hess, P. A. Ludden, and D. C. Rule. 2001. Effect of forage:concentrate ratio on ruminal digestion and duodenal flow of fatty acids in ewes. J. Anim. Sci. 79:2233–2240.[Abstract/Free Full Text]

Lammoglia, M. A., R. A. Bellows, E. E. Grings, and J. W. Bergman. 1999. Effects of prepartum supplementary fat and muscle hypertrophy genotype on cold tolerance in newborn calves. J. Anim. Sci. 77:2227–2233.[Abstract/Free Full Text]

Lardy, G. P., D. C. Adams, T. J. Klopfenstein, and H. H. Patterson. 2004. Building beef cow nutritional programs with the 1996 NRC beef cattle requirements model. J. Anim. Sci. 84(Suppl. E):E83–E92.

McGuire, M. A., and M. K. McGuire. 2000. Conjugated linoleic acid (CLA): A ruminant fatty acid with beneficial effects on human health. Proc. Am. Soc. Anim. Sci., 1999. Available: www.asas.org/jas/symposia/proceedings/0938pdf. Accessed December 10, 2000.

Merchen, N. R., and L. D. Satter. 1983. Digestion of nitrogen by lambs fed alfalfa conserved as baled hay or as low moisture silage. J. Anim. Sci. 56:943–951.[Abstract/Free Full Text]

Moore, J. E., M. H. Brant, W. E. Kunkle, and D. I. Hopkins. 1999. Effects of supplementation on voluntary forage intake, diet digestibility, and animal performance. J. Anim. Sci. 77(Suppl. 2):122–135.

Murphy, M., P. Udén, D. L. Palmquist, and H. Wiktorsson. 1987. Rumen and total diet digestibilities in lactating cows fed diets containing full-fat rapeseed. J. Dairy Sci. 70:1572–1582.

Murrieta, C. M., B. W. Hess, and D. C. Rule. 2003. Comparison of acidic and alkaline catalysts for preparation of fatty acid methyl esters from ovine muscle with emphasis on conjugated linoleic acid. Meat Sci. 65:523–529.

NRC. 1985. Page 61 in Ruminant Nitrogen Usage. Natl. Acad. Press, Washington, DC.

NRC. 1996. Nutrient Requirements of Beef Cattle. 7th rev. ed. Natl. Acad. Press, Washington, DC.

NRC. 2001. Pages 290–299 in Nutrient Requirements of Dairy Cattle. 7th rev. ed. Natl. Acad. Press, Washington, DC.

Ørskov, E. R., and R. Ryle. 1990. Energy Nutrition in Ruminants. Elsevier Science Publishers, New York.

Palmquist, D. L. 1991. Influence of source and amount of dietary fat on digestibility in lactating cows. J. Dairy Sci. 74:2601–2609.[Abstract]

Park, Y., J. M. Storkson, K. J. Albright, W. Liu, and M. W. Pariza. 1999. Evidence that the trans-10,cis-12 isomer of conjugated linoleic acid induces body composition changes in mice. Lipids 34:235–241.[Medline]

Satter, L. D., and L. L. Slyter. 1974. Effect of ammonia concentration on rumen microbial protein production in vitro. Br. J. Nutr. 32:199–208.[Medline]

Scholljegerdes, E. J., P. A. Ludden, and B. W. Hess. 2004. Site and extent of digestion and amino acid flow to the small intestine in beef cattle consuming limited amounts of forage. J. Anim. Sci. 82:1146–1156.[Abstract/Free Full Text]

Staples, C. R., J. M. Burke, and W. W. Thatcher. 1998. Influence of supplemental fat on reproductive tissues and performance of lactating cows. J. Dairy Sci. 81:856–871.[Abstract]

Tice, E. M., M. L. Eastridge, and J. L. Firkins. 1994. Raw soybeans and roasted soybeans of different particle sizes. 2. Fatty acid utilization by lactating dairy cows. J. Dairy Sci. 77:166–180.[Abstract]

Turpeinen, A. M., M. Mutanen, A. Aro, I. Salminen, S. Basu, D. L. Palmquist, and J. M. Griinari. 2002. Bioconversion of vaccenic acid to conjugated linoleic acid in humans. Am. J. Clin. Nutr. 76:504–510.[Abstract/Free Full Text]

Van Soest, P. J. 1994. Nutritional Ecology of the Ruminant. Cornell Univ. Press, Ithaca, NY.

Whitney, M. B., B. W. Hess, L. A. Burgwald-Balstad, J. L. Sayer, C. M. Tsopito, C. T. Talbott, and D. M. Hallford. 2000. Effects of supplemental soybean oil level on in vitro digestion and performance of prepubertal beef heifers. J. Anim. Sci. 78:504–514.[Abstract/Free Full Text]

Wood, J. D., M. Esner, A. V. Fisher, G. R. Nute, R. I. Richardson, and P. R. Sheard. 1999. Manipulating meat quality and composition. Proc. Nutr. Soc. 58:363–370.[Medline]

Wu, Z., O. A. Ohajuruka, and D. L. Palmquist. 1991. Ruminal synthesis, biohydrogenation, and digestibility of fatty acids by dairy cows. J. Dairy Sci. 74:3025–3034.[Abstract]


This article has been cited by other articles:


Home page
J ANIM SCIHome page
E. Pavan and S. K. Duckett
Corn oil or corn grain supplementation to steers grazing endophyte-free tall fescue. I. Effects on in vivo digestibility, performance, and carcass quality
J Anim Sci, November 1, 2008; 86(11): 3215 - 3223.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
E. Scholljegerdes and S. Kronberg
Influence of level of supplemental whole flaxseed on forage intake and site and extent of digestion in beef heifers consuming native grass hay
J Anim Sci, September 1, 2008; 86(9): 2310 - 2320.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
J. P. Banta, D. L. Lalman, C. R. Krehbiel, and R. P. Wettemann
Whole soybean supplementation and cow age class: Effects on intake, digestion, performance, and reproduction of beef cows
J Anim Sci, August 1, 2008; 86(8): 1868 - 1878.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
B. W. Hess, G. E. Moss, and D. C. Rule
A decade of developments in the area of fat supplementation research with beef cattle and sheep
J Anim Sci, April 1, 2008; 86(14_suppl): E188 - E204.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
J. C. Marini, D. G. Fox, and M. R. Murphy
Nitrogen transactions along the gastrointestinal tract of cattle: A meta-analytical approach
J Anim Sci, March 1, 2008; 86(3): 660 - 679.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
E. J. Scholljegerdes, S. L. Lake, T. R. Weston, D. C. Rule, G. E. Moss, T. M. Nett, and B. W. Hess
Fatty acid composition of plasma, medial basal hypothalamus, and uterine tissue in primiparous beef cows fed high-linoleate safflower seeds
J Anim Sci, June 1, 2007; 85(6): 1555 - 1564.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
E. Pavan, S. K. Duckett, and J. G. Andrae
Corn oil supplementation to steers grazing endophyte-free tall fescue. I. Effects on in vivo digestibility, performance, and carcass traits
J Anim Sci, May 1, 2007; 85(5): 1330 - 1339.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
S. L. Lake, T. R. Weston, E. J. Scholljegerdes, C. M. Murrieta, B. M. Alexander, D. C. Rule, G. E. Moss, and B. W. Hess
Effects of postpartum dietary fat and body condition score at parturition on plasma, adipose tissue, and milk fatty acid composition of lactating beef cows
J Anim Sci, March 1, 2007; 85(3): 717 - 730.
[Abstract] [Full Text] [PDF]


Home page
J ANIM SCIHome page
C. M. Murrieta, B. W. Hess, E. J. Scholljegerdes, T. E. Engle, K. L. Hossner, G. E. Moss, and D. C. Rule
Evaluation of milk somatic cells as a source of mRNA for study of lipogenesis in the mammary gland of lactating beef cows supplemented with dietary high-linoleate safflower seeds
J Anim Sci, September 1, 2006; 84(9): 2399 - 2405.
[Abstract] [Full Text] [PDF]


Home page