Postruminal Casein Infusion and Exogenous Glucagon-Like Peptide 2 Administration Differentially Stimulate Pancreatic α-Amylase and Small Intestinal α-Glucosidase Activity in Cattle.

BACKGROUND: Increasing luminal carbohydrate flow decreases pancreatic α-amylase activity but can increase jejunal maltase activity, suggesting that regulation of carbohydrase activity is perhaps uncoordinated in response to luminal carbohydrate flow. Increasing luminal casein flow increases pancreatic α-amylase activity in cattle, and exogenous glucagon-like peptide 2 (GLP-2) has been shown to increase small intestinal α-glucosidase activity in nonruminants. OBJECTIVES: The objective was to evaluate the effects of postruminal casein infusion, exogenous GLP-2, or their combination on endogenous pancreatic and small intestinal carbohydrase activity in cattle postruminally infused with starch. METHODS: Holstein steers [n = 24; 250 ± 23 kg body weight (BW)] received a continuous abomasal infusion of 3.94 g raw corn starch/kg of BW combined with either 0 or 1.30 g casein/kg of BW. Steers received subcutaneous injections in 2 equal portions daily of excipient (0.5% bovine serum albumin) or 100 µg GLP-2/kg of BW per day. At the end of the 7-d treatment period, steers were slaughtered for tissue collection. Data were analyzed using the MIXED procedure of SAS version 9.4 (SAS Institute Inc.). RESULTS: Postruminal casein infusion increased (P ≤ 0.03) pancreatic mass by 12.6%, total pancreatic α-amylase activity by 50%, and postruminal starch disappearance from 96.7% to 99.3%. Exogenous GLP-2 increased (P < 0.01) total small intestinal and mucosal mass by 1.2 kg and 896 g, respectively. Relative to control, GLP-2 and casein + GLP-2 increased (P = 0.04) total small intestinal α-glucosidase activity by 83.5%. Total small intestinal maltase, isomaltase, and glucoamylase activity was 90%, 100%, and 66.7% greater for GLP-2 and casein + GLP-2 steers compared with control. CONCLUSIONS: Casein increased pancreatic α-amylase activity, GLP-2 increased small intestinal α-glucosidase activity, and the combination of casein and GLP-2 increased both pancreatic α-amylase activity and small intestinal α-glucosidase activity. This novel approach provides an in vivo model to evaluate effects of increasing endogenous carbohydrase activity on small intestinal starch digestion.

Duodenal Infusions of Starch with Casein or Glutamic Acid Influence Pancreatic and Small Intestinal Carbohydrase Activities in Cattle.

BACKGROUND: Small intestinal starch digestion in ruminants is potentially limited by inadequate production of carbohydrases. Previous research has demonstrated that small intestinal starch digestion can be improved by postruminal supply of casein or glutamic acid. However, the mechanisms by which casein and glutamic acid increase starch digestion are not well understood. OBJECTIVES: The objective of this experiment was to evaluate the effects of duodenal infusions of starch with casein or glutamic acid on postruminal carbohydrase activities in cattle. METHODS: Twenty-two steers [mean body weight (BW) = 179 ± 4.23 kg] were surgically fitted with duodenal and ileal cannulas and limit-fed a soybean hull-based diet containing small amounts of starch. Raw cornstarch (1.61 ± 0.0869 kg/d) was infused into the duodenum alone (control), or with 118 ± 7.21 g glutamic acid/d, or 428 ± 19.4 g casein/d. Treatments were infused continuously for 58 d and then steers were killed for tissue collection. Activities of pancreatic (α-amylase) and intestinal (maltase, isomaltase, glucoamylase, sucrase) carbohydrases were determined. Data were analyzed as a randomized complete block (replicate group) design using the GLM procedure of SAS to determine effects of infusion treatment. RESULTS: Duodenal casein infusion increased (P < 0.05) pancreatic α-amylase activity by 290%. Duodenal glutamic acid infusion increased (P < 0.03) duodenal maltase activity by 233%. Duodenal casein infusion increased jejunal maltase (P = 0.02) and glucoamylase (P = 0.03) activity per gram protein by 62.9% and 97.4%, respectively. Duodenal casein infusion tended to increase (P = 0.10) isomaltase activity per gram jejunum by 38.5% in the jejunum. Sucrase activity was not detected in any segment of the small intestine. CONCLUSIONS: These results suggest that small intestinal starch digestion can be improved in cattle with increased small intestinal flow of casein through increases in postruminal carbohydrase activities.

Influence of dietary fructose supplementation on visceral organ mass, carbohydrase activity, and mRNA expression of genes involved in small intestinal carbohydrate assimilation in neonatal calves.

The hypothesis of this experiment was that dietary fructose would influence visceral organ mass, carbohydrase activity, and mRNA expression of carbohydrases and nutrient transporters in the small intestine in neonatal calves. Therefore, our objective was to use the neonatal calf as a model to evaluate the effects of postruminal fructose supply on small intestinal carbohydrate assimilation. Ten calves (<7 d of age; 41.2 ± 1.46 kg of body weight) were fed milk replacer at 2.0% of body weight daily (816 ± 90.5 g/d; 272 ± 30.1 g/L; dry-matter basis) in 2 equal portions and assigned to the following dietary treatment groups: (1) milk replacer (control; n = 6) or (2) milk replacer + 2.2 g of fructose/kg of body weight (fructose; n = 4). Calves were fed dietary treatments for 28 d, with jugular blood sampled every 7 d before and after the morning feeding. Calves were slaughtered, and visceral weights were recorded. Postruminal carbohydrase activities were assayed. Quantitative real-time PCR was conducted for small intestinal mRNA expression of nutrient transporters [solute carrier family 2 member 5 (GLUT5), solute carrier family 2 member 2 (GLUT2), and solute carrier family 5 member 1 (SGLT1)], carbohydrases (lactase, maltase-glucoamylase, and sucrase-isomaltase), and ketohexokinase (KHK). Data were analyzed using MIXED procedures in SAS version 9.4 (SAS Institute Inc, Cary, NC). Dietary fructose supplementation decreased serum glucose concentration. Small intestinal mass was greater in calves supplemented with fructose. Dietary fructose supplementation did not influence pancreatic α-amylase, small intestinal isomaltase, or maltase activities. Sucrase activity was undetected in the small intestine. Dietary fructose supplementation increased small intestinal glucoamylase activity per gram of tissue by 30% and increased maltase-glucoamylase mRNA expression by 6.8-fold. Dietary fructose supplementation did not influence mRNA expression of GLUT5, SGLT1, GLUT2, or KHK. Dietary fructose supplementation increased small intestinal lactase mRNA expression by 3.1-fold. Sucrase-isomaltase mRNA expression in the small intestine decreased 5.1-fold with dietary fructose supplementation. Dietary fructose supplementation does not induce sucrase activity in neonatal calves; however, sucrase-isomaltase may be transcriptionally regulated by dietary fructose in neonatal calves. More research is needed to compare glucose and fructose at isocaloric intakes to examine effects of dietary fructose at equal metabolizable energy intake.

Influence of flake density and starch retrogradation on in vitro gas production kinetics, digestibility, and ruminal fermentation characteristics of steam-flaked corn.

Previous research has demonstrated that decreasing starch availability of steam-flaked corn by increasing flake density or increasing the degree of starch retrogradation influences in vitro gas production kinetics. However, it is unclear if increasing flake density or the degree of starch retrogradation influences end-products of in vitro ruminal fermentation (methane, volatile fatty acids, ammonia). The objective of this experiment was to evaluate the effects of increasing flake density and the degree of starch retrogradation on in vitro gas production kinetics, digestibility, and ruminal fermentation characteristics of steam-flaked corn. Three ruminally-cannulated steers were fed a high-concentrate diet and sampled for ruminal digesta for an in vitro fermentation experiment with a 5×2 factorial arrangement of treatments. Steam-flaked corn was produced to flake densities of 257, 296, 335, 373, and 412 g/L by adjusting the rolls of a steam-flaker. Samples were stored for 3 d at either 23°C to produce fresh steam-flaked corn or at 55°C in heat-sealed foil bags to produce retrograded steam-flaked corn. In vitro fermentation vessels were incubated for 24 h and then assessed for fermentation parameters including dry matter digestibility, volatile fatty acid concentrations, and total gas and methane production. Increasing the degree of starch retrogradation decreased (P < 0.01) the rate of gas production across all flake densities of steam-flaked corn but did not decrease the extent of gas production. In vitro methane production, dry matter digestibility, and microbial biomass concentration were not influenced by increasing flake density or starch retrogradation. Increasing the degree of starch retrogradation decreased (P = 0.03) the molar propionate proportion and increased (P < 0.06) the molar proportions of butyrate, isobutyrate, and isovalerate and the acetate:propionate ratio. Enzymatic starch availability of steam-flaked corn was positively correlated with mean propionate proportion (r2 = 0.93) and negatively correlated with the mean butyrate proportion (r2 = 0.89). Results from the current study demonstrate that increasing the degree of starch retrogradation of steam-flaked corn decreased the rate of in vitro gas production and altered volatile fatty acid profiles in the ruminal fermentation media.

Serotonin receptor-mediated vasorelaxation occurs primarily through 5-HT4 activation in bovine lateral saphenous vein.

To better understand mechanisms of serotonin- (5-HT) mediated vasorelaxation, isolated lateral saphenous veins from cattle were assessed for vasoactivity using myography in response to increasing concentrations of 5-HT or selective 5-HT receptor agonists. Vessels were pre-contracted with 1 × 10-4 M phenylephrine and exposed to increasing concentrations of 5-HT or 5-HT receptor agonists that were selective for 5-HT1B, 5-HT2B, 5-HT4, and 5-HT7. Vasoactive response data were normalized as a percentage of the maximum contractile response induced by the phenylephrine pre-contraction. At 1 × 10-7 M 5-HT, a relaxation was observed with an 88.7% decrease (p < 0.01) from the phenylephrine maximum. At 1 × 10-4 M 5-HT, a contraction was observed with a 165% increase (p < 0.01) from the phenylephrine maximum. Increasing concentrations of agonists selective for 5-HT2B, 5-HT4, or 5-HT7 resulted in a 27%, 92%, or 44% (p < 0.01) decrease from the phenylephrine maximum, respectively. Of these 5-HT receptor agonists, the selective 5-HT4 receptor agonist resulted in the greatest potency (-log EC50) value (6.30) compared with 5-HT2B and 5-HT7 receptor agonists (4.21 and 4.66, respectively). To confirm the involvement of 5-HT4 in 5-HT-mediated vasorelaxation, blood vessels were exposed to either DMSO (solvent control) or a selective 5-HT4 antagonist (1 × 10-5 M) for 5-min prior to the phenylephrine pre-contraction and 5-HT additions. Antagonism of the 5-HT4 receptor attenuated the vasorelaxation caused by 5-HT. Approximately 94% of the vasorelaxation occurring in response to 5-HT could be accounted for through 5-HT4, providing strong evidence that 5-HT-mediated vasorelaxation occurs through 5-HT4 activation in bovine peripheral vasculature.

Influence of postruminal casein infusion and exogenous glucagon-like peptide 2 administration on the jejunal mucosal transcriptome in cattle.

We previously demonstrated that postruminal casein infusion and exogenous glucagon-like peptide 2 (GLP-2) administration independently stimulated growth and carbohydrase activity of the pancreas and jejunal mucosa in cattle. The objective of the current study was to profile the jejunal mucosal transcriptome of cattle using next-generation RNA sequencing in response to postruminal casein infusion and exogenous GLP-2. Twenty-four Holstein steers [250 ± 23.1 kg body weight (BW)] received a continuous abomasal infusion of 3.94 g raw corn starch/kg of BW combined with either 0 or 1.30 g casein/kg of BW for 7 d. Steers received subcutaneous injections at 0800 and 2000 h to provide either 0 or 100 µg GLP-2/kg of BW per day. At the end of the 7-d treatment period, steers were slaughtered for collection of the jejunal mucosa. Total RNA was extracted from jejunal mucosal tissue, strand-specific cDNA libraries were prepared, and RNA sequencing was conducted to generate 150-bp paired-end reads at a depth of 40 M reads per sample. Differentially expressed genes (DEG), KEGG pathway enrichment, and gene ontology enrichment were determined based on the FDR-corrected P-value (padj). Exogenous GLP-2 administration upregulated (padj < 0.05) 667 genes and downregulated 1,101 genes of the jejunal mucosa. Sphingolipid metabolism, bile secretion, adherens junction, and galactose metabolism were among the top KEGG pathways enriched with upregulated DEG (padj < 0.05) in response to exogenous GLP-2 administration. The top gene ontologies enriched with upregulated DEG (padj < 0.05) in response to exogenous GLP-2 administration included nutrient metabolic processes, brush border and bicellular tight junction assembly, and enzyme and transporter activities. Exogenous GLP-2 administration increased or tended to increase (padj < 0.10) brush border carbohydrase (MGAM, LCT, TREH), hexose transporter (SLC5A1, SLC2A2), and associated transcription factor (HNF1, GATA4, KAT2B) mRNA expression of the jejunal mucosa. Gene ontologies and KEGG pathways that were downregulated (padj < 0.05) in response to exogenous GLP-2 were related to genetic information processing. Postruminal casein infusion downregulated (padj < 0.05) 7 jejunal mucosal genes that collectively did not result in enriched KEGG pathways or gene ontologies. This study highlights some of the transcriptional mechanisms associated with increased growth, starch assimilation capacity, and barrier function of the jejunal mucosa in response to exogenous GLP-2 administration.

Levodopa attenuates the feed intake reduction caused by ergot alkaloids in cattle.

Consumption of ergot alkaloids from endophyte-infected tall fescue results in losses to the livestock industry in many countries and a means to mitigate these losses is needed. The objective of this study was to evaluate intra-abomasal infusion of the dopamine precursor, levodopa (L-DOPA), on dopamine metabolism, feed intake, and serum metabolites of steers exposed to ergot alkaloids. Twelve Holstein steers (344.9 ±â€…9.48 kg) fitted with ruminal cannula were housed with a cycle of heat challenge during the daytime (32 °C) and thermoneutral at night (25 °C). The steers received a basal diet of alfalfa cubes containing equal amounts of tall fescue seed composed of a mixture of endophyte-free (E-) or endophyte-infected tall fescue seeds (E+) equivalent to 15 µg ergovaline/kg body weight (BW) for 9 d followed by intra-abomasal infusion of water (L-DOPA-) or levodopa (L-DOPA+; 2 mg/kg BW) for an additional 9 d. Afterward, the steers were pair-fed for 5 d to conduct a glucose tolerance test. The E+ treatment decreased (P = 0.005) prolactin by approximately 50%. However, prolactin increased (P = 0.050) with L-DOPA+. Steers receiving E+ decreased (P < 0.001) dry matter intake (DMI); however, when supplemented with L-DOPA+ the decrease in DMI was less severe (L-DOPA × E, P = 0.003). Also, L-DOPA+ infusion increased eating duration (L-DOPA × E, P = 0.012) when steers were receiving E+. The number of meals, meal duration, and intake rate were not affected (P > 0.05) by E+ or L-DOPA+. The L-DOPA+ infusion increased (P < 0.05) free L-DOPA, free dopamine, total L-DOPA, and total dopamine. Conversely, free epinephrine and free norepinephrine decreased (P < 0.05) with L-DOPA+. Total epinephrine and total norepinephrine were not affected (P > 0.05) by L-DOPA+. Ergot alkaloids did not affect (P > 0.05) circulating free or total L-DOPA, dopamine, or epinephrine. However, free and total norepinephrine decreased (P = 0.046) with E+. Glucose clearance rates at 15 to 30 min after glucose infusion increased with L-DOPA+ (P < 0.001), but not with E+ (P = 0.280). Administration of L-DOPA as an agonist therapy to treat fescue toxicosis provided a moderate increase in DMI and eating time and increased plasma glucose clearance for cattle dosed with E+ seed.

Fescue has become the dominant cool-season perennial grass in the southeastern region of the United States and is also found in other countries. Endophytes from a plant­fungus symbiotic relationship produce toxic alkaloids that have caused significant annual economic losses to the livestock industry. Treatments to alleviate this toxicosis are still demanded. This study evaluates the infusion of the dopamine precursor, levodopa (L-DOPA), to mitigate the toxicosis caused by ergot alkaloids. When L-DOPA was infused, eating duration increased and the decrease in feed intake caused by ergot alkaloids was less severe. Additionally, circulating dopamine and glucose clearance increased with L-DOPA. These results suggest that L-DOPA has the potential to aid in the mitigation of the toxicosis caused by ergot alkaloids.

Disrupted one-carbon metabolism in heifers negatively affects their health and physiology.

The objective of this study was to determine the dose-dependent response of one-carbon metabolite (OCM: methionine, choline, folate, and vitamin B12) supplementation on heifer dry matter intake on fixed gain, organ mass, hematology, cytokine concentration, pancreatic and jejunal enzyme activity, and muscle hydrogen peroxide production. Angus heifers (n = 30; body weight [BW] = 392.6 ±â€…12.6 kg) were individually fed and assigned to one of five treatments: 0XNEG: total mixed ration (TMR) and saline injections at days 0 and 7 of the estrous cycle, 0XPOS: TMR, rumen-protected methionine (MET) fed at 0.08% of the diet dry matter, rumen-protected choline (CHOL) fed at 60 g/d, and saline injections at days 0 and 7, 0.5X: TMR, MET, CHOL, 5-mg B12, and 80-mg folate injections at days 0 and 7, 1X: TMR, MET CHOL, 10-mg vitamin B12, and 160-mg folate at days 0 and 7, and 2X: TMR, MET, CHOL, 20-mg vitamin B12, and 320-mg folate at days 0 and 7. All heifers were estrus synchronized but not bred, and blood samples were collected on days 0, 7, and at slaughter (day 14) during which tissues were collected. By design, heifer ADG did not differ (P = 0.96). Spleen weight and uterine weight were affected cubically (P = 0.03) decreasing from 0XPOS to 0.5X. Ovarian weight decreased linearly (P < 0.01) with increasing folate and B12 injection. Hemoglobin and hematocrit percentage were decreased (P < 0.01) in the 0.5X treatment compared with all other treatments. Plasma glucose, histotroph protein, and pancreatic α-amylase were decreased (P ≤ 0.04) in the 0.5X treatment. Heifers on the 2X treatment had greater pancreatic α-amylase compared with 0XNEG and 0.5X treatment. Interleukin-6 in plasma tended (P = 0.08) to be greater in the 0XPOS heifers compared with all other treatments. Lastly, 0XPOS-treated heifers had reduced (P ≤ 0.07) hydrogen peroxide production in muscle compared with 0XNEG heifers. These data imply that while certain doses of OCM do not improve whole animal physiology, OCM supplementation doses that disrupt one-carbon metabolism, such as that of the 0.5X treatment, can induce a negative systemic response that results in negative effects in both the dam and the conceptus during early gestation. Therefore, it is necessary to simultaneously establish an optimal OCM dose that increases circulating concentrations for use by the dam and the conceptus, while avoiding potential negative side effects of a disruptive OCM, to evaluate the long-term impacts of OCM supplementation of offspring programming.

The feeding of one-carbon metabolites (including methionine and B vitamins) has been shown to improve fetal growth and milk production in species such as mice, sheep, and dairy cattle. Extending this to beef cattle around the time of breeding is a growing area of research. Our group previously determined that one-carbon metabolite supplementation to beef heifers altered the abundance of circulating methionine-folate cycle intermediates in a dose-dependent manner. Therefore, we aimed to determine a whole-body response to one-carbon metabolite supplementation in heifers by measuring the effects on specific physiological systems as well as a total systemic response. We determined that treatments that negatively altered the methionine-folate cycle yielded a fundamental negative whole-body response to supplementation.

Duration of ergovaline exposure influences serotonin-mediated vasoactivity of bovine mesenteric vasculature.

Ergovaline (ERV), produced in toxic endophyte-infected tall fescue, causes potent vasoconstriction of bovine peripheral and visceral vasculature. Ergovaline acts as both an agonist and an antagonist in bovine gut blood vessels through serotonin (5-HT) receptors and it appears that the type of action could be influenced by the extent of ERV exposure. Because it was unclear how the duration of ERV exposure influences 5-HT-mediated vasoactivity, experiments were designed to evaluate how simultaneous or prior ERV exposure influenced 5-HT-mediated vasoactivity of mesenteric artery (MA) and vein (MV) segments from Holstein steers (N = 10). Vessels were incubated in Krebs-Henseleit buffer containing 0, 0.01, or 0.1 µM ERV for 24 h prior to the 5-HT dose-response or exposed to fixed concentrations of 0, 0.01, or 0.1 µM ERV simultaneously during the 5-HT dose-response. Vessels were suspended in chambers of a multimyograph containing Krebs-Henseleit buffer and equilibrated to 1 g tension for 90 min. Vessels were exposed to increasing concentrations of 5-HT (5 × 10-8 M to 1 × 10-4 M) every 15 min and contractile responses were normalized as a percentage of the maximum contractile response induced by 120 mM KCl reference addition. Two-way analysis of variance was used to separately analyze data for each vessel type and duration of exposure using the MIXED procedure of SAS. When 5-HT concentration increased from 5 × 10-8 to 1 × 10-6 M, simultaneous addition of 0.1 µM ERV increased (P < 0.01) the contractile response of MV compared with additions of 0 and 0.01 µM ERV. At 1 × 10-4 M 5-HT, the simultaneous presence of 0.01 and 0.1 µM ERV decreased (P < 0.01) the contractile response of both MA and MV compared with 0 µM ERV addition. As 5-HT concentrations increased, the contractile response increased (P < 0.01) in both MA and MV with no previous ERV exposure, but decreased in MA and MV with 24 h prior exposure to 0.01 and 0.1 µM ERV. These data demonstrate that the duration of ERV exposure influences 5-HT-mediated vasoconstriction and likely vasorelaxation in bovine mesenteric vasculature. If ERV and 5-HT exposure occur simultaneously, ERV can act as a partial agonist of 5-HT-mediated vasoconstriction. If 5-HT exposure occurs after blood vessels have had prior ERV exposure, it appears that 5-HT may induce vasorelaxation of blood vessels. More research is needed to identify cellular and molecular mechanisms involved with 5-HT-mediated vasoactivity.

Consumption of ergot alkaloids found in endophyte-infected tall fescue can lead to symptoms of fescue toxicosis, such as vasoconstriction, in ruminant livestock species. Ergovaline is one of the primary ergot alkaloids responsible for causing vasoconstriction when toxic varieties of fescue are consumed. It has been previously shown that ergovaline causes vasoconstriction by interacting with vascular serotonin receptors in cattle and sheep. Depending on when ergovaline exposure occurs, ergovaline can function as an agonist (stimulant) or antagonist (inhibitor) of vascular activity. However, it is unclear how the duration of ergovaline exposure affects vasoconstriction caused by serotonin. Experiments were conducted using the bovine mesenteric artery and mesenteric vein that were exposed to either 0, 0.01, or 0.1 µM ergovaline for 24-h prior to serotonin additions or simultaneously with serotonin additions. Maximum contractile response data were recorded using a multimyograph system and normalized as a percentage of the contractile response produced by the reference compound, KCl. The results of these experiments demonstrated that the duration of ergovaline exposure influences serotonin-mediated vasoconstriction and possibly vasorelaxation in bovine mesenteric vasculature. If ergovaline and serotonin exposure occur simultaneously, ergovaline can act as an agonist or antagonist of serotonin-mediated vasoconstriction. If serotonin exposure occurs after prior ergovaline exposure, serotonin can induce vasorelaxation of blood vessels. Understanding how complex interactions between ergovaline and serotonin occur and affect vascular function will aid in the development of strategies to mitigate sustained vasoconstriction caused during fescue toxicosis.

Influence of Anti-Coccidial Compounds and Phytogenic Saponin Extracts on In Vitro and In Vivo Ruminal Fermentation and Methane Production of Cattle.

Four experiments were conducted to evaluate sources of anti-coccidial compounds and phytogenic saponin extracts on in vitro and in vivo ruminal fermentation and CH4 production at multiple inclusion levels. In experiment 1, eight steers were fed either a finishing diet or a finishing diet supplemented with 0.5 mg/kg BW decoquinate (DCQ) and 3.33 mg/kg BW Yucca schidigera extract (YSE), and respiratory gas exchange was measured. In experiment 2, four ruminally-cannulated steers were fed the same treatments as experiment 1, and ruminal fermentation was evaluated. Anti-coccidial sources (experiment 3; monensin, DCQ, amprolium) and saponin sources (experiment 4; YSE, Quillaja saponaria extract) and levels were evaluated for effects on in vitro ruminal fermentation and CH4 production. DCQ + YSE supplementation did not influence (p ≥ 0.24) in vivo respiratory gas consumption/production, in situ DM degradation, or liquid passage kinetics. Ruminal propionate proportion tended to increase (p = 0.09) with DCQ + YSE. Monensin decreased (p ≤ 0.04) in vitro acetate:propionate and CH4 production; saponin supplementation linearly increased (p < 0.01) propionate proportion but did not influence (p ≥ 0.38) in vitro CH4 production. Saponins and non-antibiotic anti-coccidials did not influence in vitro or in vivo CH4 production with finishing diets.

In vitro gas production kinetics are influenced by grain processing, flake density, starch retrogradation, and Aspergillus oryzae fermentation extract containing α-amylase activity.

Grain processing such as particle size, flake density, or starch retrogradation can influence ruminal degradability characteristics; however, it is unclear how exogenous α-amylase supplementation interacts with different processed grains. Four experiments were conducted to compare the effects of Aspergillus oryzae fermentation extract (Amaize; Alltech Biotechnology Inc., Nicholasville, KY) supplementation on in vitro gas production kinetics of grain substrates with different processing methods that are common in the feedlot industry. In experiment 1, corn processing (dry-rolled, high-moisture, steam-flaked) and Amaize supplementation (0 or 15 U α-amylase activity/100 mL) were evaluated in a 3 × 2 factorial arrangement of treatments. The rate of gas production for dry-rolled corn was higher (P < 0.001) with Amaize supplementation. In experiment 2, flake density (296, 322, 348, 373, and 399 g/L) and starch retrogradation (storage in heat-sealed foil bags for 3 d at 23 or 55°C) were evaluated in a 5 × 2 factorial arrangement of treatments. There was a flake density × starch retrogradation interaction (P < 0.01) for the rate of gas production because the decrease in the rate of gas production with starch retrogradation was greater at lighter flake densities compared with heavier flake densities. In experiment 3, Amaize supplementation was evaluated across flake densities of nonretrograded steam-flaked corn (stored at 23°C) used in experiment 2. There was a flake density × Amaize interaction (P < 0.01) for the rate of gas production where Amaize supplementation resulted in a lower rate of gas production at lighter flake densities (296, 322, and 348 g/L) but a higher rate of gas production at heavier flake densities (373 and 399 g/L). In experiment 4, Amaize supplementation was evaluated across flake densities of retrograded steam-flaked corn (stored at 55°C) used in experiment 2. Gas production was lower after 24 h with Amaize supplementation for retrograded flakes produced to a density of 322 and 399 g/L while Amaize supplementation did not influence gas production at 24 h at other flake densities. There was a flake density × Amaize interaction for the rate of gas production because Amaize supplementation resulted in a faster (P < 0.01) rate of gas production for all flake densities except retrograded flakes produced to a density of 296 g/L. Enzymatic starch availability was positively correlated with the rate of gas production. These data demonstrate that supplementation of 15 U/100 mL of Amaize resulted in greater rates of gas production for dry-rolled corn, corn steam-flaked to heavier densities, and retrograded steam-flaked corn.

Grain processing has been used for decades to improve digestibility of finishing cattle diets, leading to improved growth and feed efficiency. Grain processing methods that result in changes in particle size, flake density, or starch retrogradation have all been shown to affect the degradability characteristics of nutrients in the rumen. Supplementation of feed additives containing exogenous enzyme activity could have the potential to improve digestibility, growth performance, and feed efficiency of livestock. However, it is unknown how supplementation of exogenous α-amylase activity influences degradability characteristics of different processed grains. The objectives of this study were to compare the effects of Aspergillus oryzae fermentation extract supplementation on in vitro gas production kinetics of grain substrates with different processing methods that are common in the feedlot industry. Enzymatic starch availability of steam-flaked corn, but not dry-rolled or high-moisture corn, was reflective of the rate of in vitro gas production. Increasing flake density and increasing starch retrogradation decreased the rate of in vitro gas production. Supplementation of A. oryzae fermentation extract resulted in increased rates of gas production for dry-rolled corn, corn steam-flaked at heavier densities, and retrograded steam-flaked corn.

Timing of maternal nutrient restriction during mid- to late-gestation influences net umbilical uptake of glucose and amino acids in adolescent sheep.

Previous research demonstrated that maternal nutrient restriction during mid- to late-gestation influenced net umbilical uptakes of glucose and amino acids in sheep. However, it is unclear how the timing and duration of nutrient restriction during mid- to late-gestation influences net uterine, uteroplacental, and fetal flux of glucose and amino acids. On day 50 of gestation, 41 adolescent ewe lambs carrying singletons were randomly assigned to one of six dietary treatments: 1) 100% of nutrient requirements from days 50 to 90 of gestation (CON; n = 7); 2) 60% of nutrient requirements (RES; n = 7) from days 50 to 90 of gestation; 3) 100% of nutrient requirements from days 50 to 130 of gestation (CON-CON; n = 6); 4) 100% of nutrient requirements from days 50 to 90 of gestation and 60% of nutrient requirements from days 90 to 130 of gestation (CON-RES; n = 7); 5) 60% of nutrient requirements from days 50 to 90 of gestation and 100% of nutrient requirements from days 90 to 130 of gestation (RES-CON; n = 7); or 6) 60% of nutrient requirements from days 50 to 130 of gestation (RES-RES; n = 7). On day 90 (n = 14) and day 130 (n = 27), intraoperative procedures were performed to evaluate uteroplacental blood flows, collect blood samples, and then ewes were euthanized. Net uterine, uteroplacental, and umbilical fluxes of glucose and amino acids were calculated by multiplying blood flow by the arterial-venous concentration difference. Data from days 90 and 130 were analyzed separately using ANOVA in SAS. Maternal nutrient restriction during mid-gestation increased (P = 0.04) net umbilical glucose uptake but, maternal nutrient restriction during late-gestation decreased (P = 0.02) net umbilical glucose uptake. Net umbilical essential amino acid uptake decreased (P = 0.03) with nutrient restriction during mid-gestation; however, net umbilical uptakes of Phe (P = 0.02), Thr (P = 0.05), Met (P = 0.09), and His (P = 0.08) increased or tended to increase after nutrient restriction during late-gestation. These data demonstrate that net umbilical glucose and amino acid uptakes were influenced by the timing of nutrient restriction during mid- to late-gestation. Elevated net umbilical glucose uptake after mid-gestational nutrient restriction was sustained throughout late-gestation, independent of late-gestational feeding level. Long-term adaptations in umbilical glucose uptake may have implications for prenatal and postnatal growth and development of the offspring.

Maternal undernutrition during gestation can lead to decreased fetal growth, decreased uteroplacental blood flow, and changes in nutrient supply to the fetus. However, it is unclear how the timing (mid-gestation vs. late-gestation) and duration (40 d vs. 80 d) of nutrient restriction influence nutrient supply to the fetus during mid- to late-gestation. Pregnant ewe lambs fed a pelleted diet to meet 100% of nutritional requirements or 60% of nutritional requirements during mid-gestation alone (days 50 to 90) or during mid- and late-gestation (days 50 to 130). At the end of mid-gestation and late-gestation, the net nutrient supply between the maternal, placental, and fetal compartments was measured. The results indicated that the timing of nutrient restriction influenced the net nutrient supply to the fetus but, the duration of nutrient restriction did not. Nutrient restriction during mid-gestation increased glucose to the fetus but nutrient restriction during late-gestation decreased glucose to the fetus. The opposite response occurred for fetal essential amino acid supply where nutrient restriction during mid-gestation decreased essential amino acid supply to the fetus but increased for several essential amino acids during late-gestational nutrient restriction. The timing of maternal undernutrition during mid- to late-gestation can affect the amount of nutrients delivered to the fetus and thus, could potentially impact postnatal growth and development.

Influence of form of selenium supplementation and tall fescue endophyte toxicity on growth performance, serum parameters, and tissue masses of grazing beef steers.

To test the hypothesis that average daily gain (ADG) and clinical parameters of steers grazing novel non-toxic (NTE) or toxic KY-31 (TE) endophyte-infected tall fescue would be improved by ad libitum intake of vitamin-mineral mixes (V-M) that contain 27 ppm Se as a 1:1 blend of SELPLEX:sodium selenite (MIX) vs. sodium selenite (ISe), 32 fescue-naïve beef steers partially depleted of Se were randomly assigned to ad libitum consumption ISe vs. MIX for 35 days and fed enough of a NTE/alfalfa/grain diet to achieve 0.57 kg BW gain/day. Then, within Se-form treatments, two steers were randomly assigned to each of four NTE (ISe = 316 ± 31 kg BW, MIX = 315 ± 22 kg BW) or TE (ISe = 316 ± 37 kg BW, MIX = 314 ± 39 kg BW) paddocks for 84 days and had ad libitum access to their respective V-M. The MIXED procedure of SAS was used to assess effects of day, Se-form (ISe, MIX) and endophyte (NTE, TE) treatments, and their interactions. Whole blood Se decreased (P < 0.01) 31% from days 0 to 84 and was 6.2% greater (P < 0.01) for MIX steers. Serum prolactin decreased (P < 0.01) 18% for NTE and 48% for TE steers from days 0 to 84 and was 17% greater (P = 0.01) for MIX vs. ISe for TE steers. Serum alkaline phosphatase activity decreased (P < 0.02) 27% from days 0 to 84 and was 15% greater (P < 0.02) for MIX steers. Serum urea nitrogen increased (P < 0.02) 8.2% from days 0 to 84 for TE but not NTE steers. Average daily gain was less (P < 0.01) for steers grazing TE (-0.18 kg/day) compared with NTE (0.09 kg/d). Although there was increased serum alkaline phosphatase activity and increased serum prolactin for TE + MIX steers compared with TE + ISe steers, MIX supplementation was unable to increase serum prolactin concentrations or ADG to the same levels as steers grazing NTE. Longer adaptation to MIX supplementation ad libitum may be necessary for maximal Se assimilation to restore serum prolactin levels in steers grazing TE.

Corn processing, flake density, and starch retrogradation influence ruminal solubility of starch, fiber, protein, and minerals.

Five ruminally cannulated steers (body weight = 390 ± 7.86 kg) were used in three experiments to evaluate effects of corn processing, flake density, and starch retrogradation on in situ ruminal degradation. In experiment 1, corn was left whole or processed with no screen, ground through a 6-mm screen, or ground through a 1-mm screen. In experiment 2, we produced steam-flaked corn at four densities: 309, 335, 360, and 386 g/L. These four flake densities were sifted for 20 s through a 4-mm screen to produce two particle sizes within each flake density: sifted flakes (>4 mm) and sifted fines (<4 mm). In experiment 3, sifted flakes (335 g/L) were stored for 3-d at either 23 °C (starch availability = 55%) or 55 °C to induce starch retrogradation (starch availability = 41%). All samples for each of the three experiments were weighed into nylon bags and ruminally incubated for 0-h to estimate the soluble fraction. The residue remaining was analyzed for nutrient composition. In experiment 1, whole shelled corn had lesser (P < 0.01) ruminal solubility of all nutrients measured compared with ground corn. Corn ground with a screen (6 and 1 mm) had greater (P < 0.01) ruminal solubility of all nutrients measured compared with corn ground with no screen. Corn ground through a 1-mm screen had greater (P < 0.03) ruminal solubility of DM, total starch, CP, ADF, AHF, P, Mg, K, S, Zn, Fe, and Mn compared with corn ground through a 6-mm screen. In experiment 2, increasing flake density linearly decreased (P < 0.02) the soluble fraction of DM, total starch, CP, ADF, AHF, P, K, S, and Zn of sifted flakes. The soluble DM fraction of sifted fines tended to decrease (P = 0.06) linearly with increasing flake density. Total starch, CP, NDF, and Zn soluble fractions of sifted fines were not influenced by flake density. In experiment 3, storage of sifted flakes in heat-sealed foil bags at 55 °C for 3-d decreased (P < 0.04) the soluble fractions of DM, total starch, CP, NDF, P, Mg, K, S, and Fe. With each increase in the degree of corn processing, there was an increase in the solubility of nutrients. Increasing flake density can decrease ruminal solubility of flakes; however, the soluble fraction of sifted fines is not influenced as much by changes in flake density. Inducing starch retrogradation decreases ruminal solubility of starch, nonstarch OM, and minerals.

Grain processing has been used for decades to improve digestibility of finishing cattle diets, leading to improved growth performance and feed efficiency. The soluble fraction of a feed can be defined as the fraction that disappears immediately in the rumen and its measurement can be useful for understanding kinetic properties of feed digestion. Grain processing methods that result in changes in particle size, flake density, or starch retrogradation have been shown to affect the soluble fraction of dry matter in the rumen. However, it is unknown how the solubility of different nutrients are affected by these changes. The objective of this experiment was to characterize how corn processing, flake density, particle size, and starch retrogradation influence the soluble fraction of starch, protein, fiber, and minerals. With each increase in the degree of corn processing, there was an increase in the solubility of nutrients. Increasing flake density can decrease ruminal solubility of flakes; however, the soluble fraction of sifted fines is not influenced as much by changes in flake density. Inducing starch retrogradation decreases ruminal solubility of starch, nonstarch OM, and minerals. Understanding the factors influencing ruminal solubility of processed corn is important when modeling digestion in beef cattle.

Prenatal and Postnatal Nutrition Influence Pancreatic and Intestinal Carbohydrase Activities of Ruminants.

In ruminant livestock species, nutrition can play an important role in the long-term programming of gastrointestinal function. Pancreatic and small intestinal digestive enzymes are important for postruminal digestion of carbohydrates and protein. Carbohydrases have been shown to respond to changes in the level of feed intake and the dietary inclusion of specific nutrients, including arginine, butyrate, folic acid, fructose, and leucine. Understanding how diet influences enzyme development and activity during prenatal and postnatal life could lead to the development of dietary strategies to optimize offspring growth and development to increase digestive efficiency of ruminant livestock species. More research is needed to understand how changes in fetal or neonatal carbohydrase activities in response to nutrition influence long-term growth performance and efficiency in ruminant livestock species to optimize nutritional strategies.

Influence of air equilibration time, sampling techniques, and storage temperature on enzymatic starch availability of steam-flaked corn.

Measuring enzymatic starch availability is commonly used as a quality control method to ensure steam-flaked corn manufacturing consistency in commercial cattle feeding operations. However, starch availability estimates can be variable. We conducted five experiments to evaluate factors influencing starch availability estimates of steam-flaked corn. In Exp. 1, sample handling methods were evaluated. Sifted flakes were immediately placed into a plastic bag, air equilibrated for 240 min, oven-dried, or freeze-dried. Directly oven-drying samples at 55°C decreased (P < 0.01) starch availability compared to other sample handling methods. In Exp. 2, sifted flakes were air equilibrated for 0, 15, 30, 60, 120, or 240 min. Air equilibration time did not influence (P ≥ 0.54) starch availability. In Exp. 3, samples were evaluated for effects of sifting through a 4-mm screen (flakes + fines vs. sifted flakes) and air equilibration time (0 vs. 240 min). Both sifting steam-flaked corn samples and air equilibration for 240 min increased starch availability (P < 0.01 and P = 0.02, respectively). In Exp. 4, we evaluated the effects of air equilibration time (0 vs. 240 min) on the two sifted portions (sifted flakes vs. sifted fines). There was an air equilibration time × sifted portion interaction for starch availability because air equilibration time increased (P < 0.01) starch availability of sifted fines but did not influence starch availability of sifted flakes. Concentrations of crude protein, soluble crude protein, neutral and acid detergent fiber, ether extract, and acid-hydrolyzed fat, Ca, P, K, Mg, S, Fe, Zn, Mg, and Cu were greater (P < 0.01) for sifted fines compared to sifted flakes. Starch availability and total starch concentration were greater (P < 0.01) for sifted flakes compared to sifted fines. In Exp. 5, effects of air equilibration time (0 vs. 240 min) and storage temperature (23°C vs. 55°ºC) on flakes + fines were evaluated. Storage of flakes + fines in heat-sealed foil bags at 55°C for 3-d decreased (P < 0.01) starch availability by 40.7%. Sifted flakes contained less moisture, greater total starch concentrations, and greater starch availability than sifted fines. Moisture, sifting, air equilibration time, and storage temperature influence starch availability of steam-flaked corn. Adoption of the strategies discussed in the current study will lead to more consistent estimates of starch availability.

Flake density and starch retrogradation influence in situ ruminal degradability characteristics of steam-flaked corn and predicted starch digestibility and energetic efficiency.

Five ruminally cannulated steers (body weight = 390 ± 7.86 kg) were used in two experiments to evaluate the effects of flake density and starch retrogradation on in situ ruminal degradation of steam-flaked corn. In experiment 1, sifted flakes with flake densities of 257, 296, 335, 373, and 412 g/L (enzymatic starch availabilities: 87%, 76%, 66%, 43%, and 49%, respectively) were evaluated in a randomized complete block design experiment. In experiment 2, the experimental design was a randomized complete block design with a 3 × 2 factorial arrangement of treatments. Three steam-flaked corn fractions corresponding to different particle sizes were used: flakes + fines (not sifted; >4 and <4 mm), sifted flakes (>4 mm), and sifted fines (<4 mm). Particle size fractions were stored for 3 d at either 23 °C or 55 °C (starch availabilities averaged across particle sizes: 53.3% and 25.5%, respectively) in heat-sealed foil bags. Samples were ruminally incubated for 0, 3, 6, 12, 24, 48, 72, or 96 h. Degradation data were modeled to obtain the rate and extent of degradation and passage rate was set to 6% per hour. In experiment 1, the rate of degradation decreased linearly (P < 0.01) and in situ ruminal dry matter (DM) degradability decreased linearly (P < 0.01) from 78.9% to 57.3% as flake density increased from 257 to 412 g/L. In experiment 2, storage of steam-flaked corn samples at 55 °C for 3 d decreased (P < 0.01) the rate of degradation by 37.6% across all particle sizes. Storing samples at 55 °C for 3 d decreased (P < 0.01) in situ ruminal DM degradability of flakes + fines, sifted flakes, and sifted fines by 20.9%, 22.6%, and 14.7%, respectively. Using data from experiment 1 and 2, enzymatic starch availability of sifted flakes was positively correlated (R2 = 0.97; P < 0.01) with in situ ruminal DM degradability. The results demonstrate that decreased starch availability resulting from either starch retrogradation or increased flake density is associated with decreased ruminal digestibility. Decreases in starch availability and in situ ruminal degradability may indicate that increasing flake density or starch retrogradation could potentially alter the site of digestion in cattle. Using prediction equations, decreases in ruminal starch digestibility of steam-flaked corn caused by increasing flake density or increasing starch retrogradation could increase energetic efficiency, depending on the rate of passage and if small intestinal starch digestibility is maintained.

Effects of Nutrient Restriction During Midgestation to Late Gestation on Maternal and Fetal Postruminal Carbohydrase Activities in Sheep.

To examine the effects of nutrient restriction during midgestation to late gestation on maternal and fetal digestive enzyme activities, 41 singleton ewes (48.3 ±â€…0.6 kg of BW) were randomly assigned to dietary treatments: 100% (control; CON; n = 20) or 60% of nutrient requirements (restricted; RES; n = 21) from day 50 until day 90 (midgestation). At day 90, 14 ewes (CON, n = 7; RES, n = 7) were euthanized. The remaining ewes were subjected to treatments of nutrient restriction or remained on a control diet from day 90 until day 130 (late gestation): CON-CON (n = 6), CON-RES (n = 7), RES-CON (n = 7), and RES-RES (n = 7) and were euthanized on day 130. The fetal and maternal pancreas and small intestines were weighed, subsampled, and assayed for digestive enzyme activity. One unit (U) of enzyme activity is equal to 1 µmol of product produced per minute for amylase, glucoamylase, lactase, and trypsin and 0.5 µmol of product produced per minute for maltase and isomaltase. Nutrient restriction during midgestation and late gestation decreased (P < 0.05) maternal pancreatic and small intestinal mass but did not affect fetal pancreatic or small intestinal mass. Maternal nutrient restriction during late gestation decreased (P = 0.03) fetal pancreatic trypsin content (U/pancreas) and tended to decrease (P < 0.08) fetal pancreatic trypsin concentration (U/g), specific activity (U/g protein), and content relative to BW (U/kg of BW). Nutrient restriction of gestating ewes decreased the total content of α-amylase (P = 0.04) and tended to decrease total content of trypsin (P = 0.06) and protein (P = 0.06) in the maternal pancreas on day 90. Nutrient restriction during midgestation on day 90 and during late gestation on day 130 decreased (P = 0.04) maternal pancreatic α-amylase-specific activity. Sucrase activity was undetected in the fetal and maternal small intestine. Nutrient restriction during late gestation increased (P = 0.01) maternal small intestinal maltase and lactase concentration and tended to increase (P = 0.06) isomaltase concentration. Realimentation during late gestation after nutrient restriction during midgestation increased lactase concentration (P = 0.04) and specific activity (P = 0.05) in the fetal small intestine. Fetal small intestinal maltase, isomaltase, and glucoamylase did not respond to maternal nutrient restriction. These data indicate that some maternal and fetal digestive enzyme activities may change in response to maternal nutrient restriction.

Influence of maternal nutrient restriction and rumen-protected arginine supplementation on post-ruminal digestive enzyme activity of lamb offspring.

To determine the influence of maternal nutrient restriction and rumen-protected arginine supplementation on post-ruminal digestive enzyme activity in lambs, 31 multiparous, Rambouillet ewes were allocated to one of three dietary treatments at 54 d of gestation. Dietary treatments were 100% of nutrient requirements (control, CON; n = 11), 60% of control (restricted, RES; n = 10), or RES plus 180 mg rumen-protected arginine•kg BW-1•d-1 (RES-ARG; n = 10). Immediately after parturition, lambs were removed from dams and reared independently. Milk-replacer and alfalfa hay + creep feed were offered for ad libitum intake. At day 54 of age, lambs were slaughtered and the pancreas and small intestine were collected. Pancreatic (α-amylase and trypsin) and jejunal (maltase, glucoamylase, sucrase, isomaltase, and lactase) digestive enzyme activities were assayed. Data were analyzed using the GLM procedure of SAS for effects of treatment. Contrast statements were used to determine differences between means for effects of restriction (CON vs. RES and RES-ARG) and rumen-protected arginine supplementation (RES vs. RES-ARG). There was no influence (P ≥ 0.15) of maternal nutrient restriction or rumen-protected arginine supplementation on pancreatic or jejunal protein concentrations. No treatment effects were observed (P ≥ 0.12) for enzymes involved in starch digestion including pancreatic α-amylase and jejunal maltase, glucoamylase, and isomaltase. Sucrase activity was undetected in the jejunum of lambs across all treatments. Maternal nutrient restriction tended to increase (P = 0.08) pancreatic trypsin activity per gram protein in lambs. Lactase activity per gram protein in the jejunum of lambs tended to decrease (P = 0.09) with maternal nutrient restriction. Rumen-protected arginine supplementation to gestating ewes did not influence (P ≥ 0.19) digestive enzyme activities of lamb offspring. These data suggest that maternal nutrient restriction and rumen-protected arginine supplementation have minimal effects on digestive enzyme activity in offspring.

Effects of ractopamine hydrochloride supplementation on feeding behavior, growth performance, and carcass characteristics of finishing steers.

Ractopamine hydrochloride (RAC) is a ß-adrenergic agonist that functions as a repartitioning agent to improve muscling in feedlot cattle. Many studies have investigated the effects of RAC on growth performance and carcass characteristics; however, there is minimal information about the influence of RAC on feeding behavior. Sixty-nine steers (body weight [BW] = 364 ± 3.9 kg) predominately of Angus and Simmental breeding were subjected to a 126-d (n = 46) or 154-d (n = 23) feeding period and randomly assigned to one of two treatment groups: supplementation to provide 0 (CON; n = 34) or 267 ± 4.9 mg/d of RAC (n = 35). Ractopamine was provided as Optaflexx 45 at 0.024% of the diet (dry matter [DM] basis; Elanco Animal Health, Greenfield, IN). Dietary treatments were fed the final 42 d in the feed yard (treatment period). Feeding behavior and growth performance were measured using radio frequency identification tags and the Insentec feeding system. Following the final day of treatment, steers were slaughtered and carcass measurements were recorded. Data were analyzed using MIXED models in SAS. There were no differences in BW, average daily gain (ADG), DM intake (DMI), gain:feed ratio (G:F), or feeding behavior during the pretreatment period (P > 0.44). Ractopamine supplementation increased G:F during the treatment period (P = 0.02) and during the total period (P = 0.03) and tended to increase ADG during the treatment and total period (P ≤ 0.08). DMI was not affected during the treatment or total period (P > 0.67). Eating time per visit, per meal, and per day were decreased (P < 0.02) in steers supplemented with RAC during the treatment period. DMI per minute was increased (P = 0.02) in steers supplemented with RAC. Hot carcass weight, dressing percentage, and 12th rib fat were not influenced by RAC supplementation. Ractopamine supplementation decreased marbling (P = 0.008) and kidney, pelvic, and heart percentage (P = 0.04) and increased longissimus muscle area (P = 0.01). These data demonstrate that RAC supplementation for 42 d improves feed efficiency, increases the rate of DMI without altering DMI, and increases muscling in finishing cattle.