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Mitochondria of bovine liver contain acyl CoA synthetases necessary for the uptake of propionate, butyrate, and valerate whereas acetate is bound only weakly. Purification of these enzymes separated a distinct propionyl CoA synthetase highly specific for propionate and acrylate and a butyrate-activating fraction with broad substrate specificity for short and medium chain fatty acids. Evidence from kinetic studies and sucrose density centrifugation suggested that this latter fraction was composed of two enzymes, a butyryl CoA synthetase and a valeryl CoA synthetase. The apparent molecular weights of the propionyl, butyryl, and valeryl CoA synthetases were 72,000, 67,000, and 65,000. The Michaelis-Menten constants of propionyl CoA synthetase for propionate, adenosine 5'-triphosphate, and coenzyme A were 1.3 x 10(-3)M, 1.3 x 10(-3)M, and 6.3 x 10(-4)M. Enzyme activity is regulated by the concentration of propionate in portal blood. Relative to propionyl, butyryl, or valeryl CoA synthetases little acetyl CoA synthetase could be demonstrated. In ruminants hepatic metabolism is such that use of acetate as an energy source is minimum. This ensures that an alternative energy source to glucose, as acetate units, will reach the extrahepatic tissues. Separation of a distinct propionyl CoA synthetase regulated by the concentration of propionate in portal blood is significant because a primary role of ruminant liver is to synthesize glucose from ruminally derived propionate.
Mixed ruminal bacteria (400 mg cells/liter) were incubated in artificial media containing ammonia, sodium carbonate, macrominerals, vitamins, sulfide, microminerals, acetate, propionate, and butyrate. When mixed carbohydrates (equal parts glucose, maltose, sucrose, cellobiose, and soluble starch) were added at 155 mg/liter per h for 10 h, average bacterial growth rate was slow, and dry weight yield was greater than 23%. Additions of isobutyrate, valerate, isovalerate, and 2 methylbutyrate had little influence on synthesis of bacterial dry weight, deoxyribonucleic acid, ribonucleic acid, or carbohydrate. When a timothy hay inoculum was used, isovalerate and 2 methyl-butyrate increased protein synthesis by 11.2 and 16.4%, but isobutyrate and valerate alone were without effect. All four acids combined increased bacterial protein by 18.7%. Responses with an inoculum of 60% concentrate and mixed hay were smaller and not statistically different from control incubations. Low concentrations of Trypticase (less than 250 mg/liter) improved efficiency of microbial protein synthesis from organic matter, but more was associated with decreased efficiency and utilization of extracellular ammonia.
When diets containing 33 and 18 ppm monensin sodium were fed for ad libitum intake to dairy goats, milk fat content was reduced by 15 and 5%. Milk protein content was increased 10% when 33 ppm sodium monensin was fed with diet at restricted intake. Milk yield was not affected. Both ad libitum and restricted consumption of diet containing 33 ppm monensin sodium reduced ratios of ruminal acetate:propionate. These resulted from increased propionate concentration with ad libitum consumption and from reduced acetate with restricted feeding. Diets containing 18 ppm monensin sodium resulted in slightly higher concentrations of both propionate and acetate. Monensin sodium did not reduce feed intake significantly.
To determine responses to abomasal protein infusion and ruminal acetate: propionate ratios, four lactating Toggenburg goats fed hourly a 70% roughage and 30% concentrate diet were used in a Latin-square design with a factorial arrangement of treatments. Either acetate or propionate was infused ruminally and casein or saline infused abomasally. Estimated net energy and volume of the infusates were similar for all treatments. To examine the effects of treatments on glucose metabolism, 2-carbon-14 propionate was infused ruminally and 6-hydrogen-3 glucose was infused intravenously for 9 and 5 h, respectively. Although glucose concentration in plasma was higher and propionate turnover greater with propionate treatment, percentage of glucose derived from propionate, amount of propionate coverted to glucose, and glucose turnover remained unchanged. No differences in glucose metabolism due to the abomasal casein infusion were evident. To determine the effects of treatment on insulin, glucagon, growth hormone, and prolactin in plasma, samples were collected at 10-min intervals for 3 h at 0400 and 1600 h. No diurnal variation or consistent peaks were observed for any of the hormones nor were treatment effects on plasma concentrations of insulin, growth hormone, or prolactin evident. Glucagon concentration was higher with casein treatment; however, no relationship existed between glucagon in plasma and glucogenic parameters measured.
Isolated caprine hepatocytes were incubated with fatty acids of various chain lengths. Short-chain fatty acids effects on rates of gluconeogenesis and oxidation from [2-14C]propionate were determined. Additions of glucose (2.5 mM) had no effect on hepatic [2-14C]propionate metabolism in the presence and absence of amino acids. A complete mixture of amino acids increased label incorporation from [2-14C]propionate into [14C]glucose by 22%. Butyrate inhibited [2-14C]propionate metabolism and increased the apparent Michaelis constant for [2-14C]propionate incorporation into [14C]glucose from 2.4 +/- 1.5 to 5.6 +/- .9 mM. Butyrate's effects on propionate were similar in the presence and absence of L-carnitine (1 mM). Isobutyrate, 2-methylbutyrate, and valerate (1.25 mM) had no effect on [14C]glucose production but decreased 14CO2 production to 57, 61, and 54% of the control [2-14C]propionate (1.25 mM). This inhibition on 14CO2 production was not competitive. Isovalerate had no effect on either [2-14C]propionate incorporation into glucose or CO2. An increase in ratio of [14C]glucose to 14CO2 from [2-14C]propionate demonstrated that short-chain fatty acids other than butyrate do not inhibit gluconeogenesis from propionate. In addition, fatty acids that generate a net synthesis of intracellular oxaloacetate may partition propionate carbons toward gluconeogenic rather than oxidative pathways in goat hepatocytes.
Net absorption of L-lactate and VFA from the portal drained viscera of first lactation Holstein cows was measured at 4, 8, (four cows), 12, 16, and 20 wk (two cows) of lactation. Chronic indwelling catheters were installed 7 to 14 d postpartum in appropriate vessels to measure blood flow and net nutrient absorption. Cows were fed a completely mixed, 60:40 (dry basis) corn silage:supplement diet and milked every 12 h. Average metabolizable energy intake was 2.8X maintenance and mean milk production was 24.0 kg. Net absorption of lactate, any of the VFA, or their total was not affected by week postpartum. Net absorption of L-lactate, VFA, and alpha-amino N accounted in sum for 53.6% of metabolizable energy intake; contributions of each component to energy in absorbed nutrients were acetate and propionate, 29.5% each; alpha-amino N, 23.4%; L-lactate, 5.4%; n-butyrate, 5.3%; 2-methylbutyrate, 2.8%, and i-butyrate, i-valerate, and n-valerate, 1.2 to 1.6% each. Comparison of paired samples of blood and plasma showed that blood cells contribute to the transport of acetate, propionate, i-butyrate, and 2-methyl-butyrate but not of n-butyrate, i-valerate, or n-valerate.
Net flux of VFA, D-beta-hydroxybutyrate, nonesterified fatty acids, and blood gasses across portal-drained viscera and liver was measured in four lactating Holstein cows fed a 60:40 corn silage: concentrate diet ad libitum and milked at 12-h intervals. Twelve consecutive hourly measurements of net flux (venous-arterial concentration difference times blood flow) were obtained during wk 4 and 8 postpartum for each cow. Milk yield and DM intake averaged 32.2 and 15.6 kg/d. On a net basis, hepatic tissues produced acetate and removed 63 to 101% of other VFA absorbed by portal-drained viscera. Hepatic and portal-drained visceral tissues produced 60 and 40%, respectively, of D-beta-hydroxybutyrate produced by splanchnic tissues. Hepatic tissues removed 9.3% of nonesterified fatty acids in portal vein and hepatic arterial blood. Oxygen use was greater by liver than for portal-drained viscera (3062 vs. 2394 mmol/h). Net portal-drained visceral flux of VFA, D-beta-hydroxybutyrate, alpha-amino nitrogen, L-lactate, and oxygen together accounted for 84.9% of calculated metabolizable energy intake. Net hepatic removal of propionate, L-lactate, and alpha-amino nitrogen maximally accounted for 55.1, 17.4, and 16.5% of carbon in glucose produced by hepatic tissues.
Contribution of cecal and ruminal VFA to metabolizable energy was investigated in steers with cannulas in both the rumen and cecum. Animals were fed ad libitum so that data would be applicable to the lactating dairy cow. Diets assigned within a 4 x 4 Latin square were: 20% long alfalfa hay and 80% concentrate; 15% pelleted alfalfa, 5% hay and 80% concentrate; 80% hay and 20% concentrate; 60% pellets, 20% hay and 20% concentrate. Intake of DM was unaffected by diet. Cecal fluid pH, osmolality, and concentrations of valerate and isovalerate were unaffected by diet. Concentrations of total VFA, acetate, propionate, butyrate, and lactate in the cecum increased with proportion of grain in the diet. The high grain diets depressed cecal ammonia concentration and acetate to propionate ratio. Acetate production in the cecum was higher with the high grain diets whereas that in the rumen was lower. Production of propionate and butyrate in both the cecum and rumen was unaffected by diet. Cecal VFA provided 8.6% of metabolizable energy intake, on average. Contribution of ruminal VFA to total metabolizable energy was affected by diet, accounting for 72, 51, 74, and 52% of metabolizable energy from the 20% hay, 20% pelleted alfalfa, 80% hay, and 80% pelleted alfalfa, respectively. Cecal VFA were an important source of energy for ad libitum-fed steers; this contribution would undoubtedly increase with increasing feed intake.
Previous reports have demonstrated that butyrate inhibits metabolism of propionate by liver cells isolated from sheep and goats. Our objectives were to examine some possible mechanisms for this inhibition and to test for this inhibition in the bovine animal. Incorporation of label from 2.5 mM [2-(14)C]propionate into glucose (nmol propionate/mg cell DM/h) in the presence of 0, 1.25, and 2.5 mM butyrate was 107, 66, and 62 by goat hepatocytes and 79, 25, and 29 by calf hepatocytes; therefore, butyrate inhibited propionate metabolism at least as effectively in calves as in goats. In goat hepatocytes 1.25 mM butyrate reduced 1.25 mM propionate uptake to 46% of control, and 1.25 mM [2-(14)C] propionate incorporation into glucose to 44% of control. Propionate had no effect on butyrate uptake. Isovalerate and valerate tended to be cleared from the media to a greater extent than butyrate but had no effect on propionate uptake. Therefore, inhibition of propionate conversion to glucose by butyrate is specific and is not due to a general competition among VFA for metabolism. Butyrate inhibits hepatic propionate utilization generally, not specifically propionate conversion to glucose. Butyrate also inhibited propionate utilization by goat liver homogenates, indicating that butyrate inhibits propionate metabolism at a step subsequent to propionate transport across the hepatocyte plasma membrane.
In previous studies ammonium salts of a mixture of isobutyrate, 2-methylbutyrate, isovalerate, and valerate were fed in a corn silage, corn, corn gluten meal, and urea diet to Holstein cows throughout lactation to define the optimum level of ammonium salts of milk production. The objective of this work was to conduct another dose response study using other forage and protein sources and to determine the effects of decreasing VFA intakes as lactation advanced. The concentrate portion of the diet contained 0, .4, .8, 1.2, or 1.6% ammonium salts of VFA. The forage to concentrate ratio was 50:50, 60:40, and 70:30 for the first, middle, and last third of lactation, respectively. The study was conducted at four university locations using 191 Holstein cows. Feeds used included corn silage, alfalfa silage or hay, corn, soybean meal, minerals, and vitamins. Treatment x location interactions were significant for milk yield during early lactation. During mid- and late lactation, supplemental VFA (.8%) improved milk and protein yield. Milk composition was not greatly affected by feeding VFA. In mid-lactation, cows fed .8% ammonium salts of VFA ate more feed than did controls. Feed efficiencies were similar among groups throughout the experiment. Cows fed VFA tended to gain less BW during lactation than did controls. Health and reproduction were not different among groups.
Effects of weaning age on plasma VFA were examined using 16 Holstein heifer calves. Animals entered the study at 6 +/- 3.5 d of age and were fed 1.8 kg milk twice daily to 28 (early weaning) or 56 d (late weaning) and a commercial pelleted calf starter from 0 (early) or 28 (late) d. Blood was sampled once weekly for 14 wk. Total blood concentrations of VFA, acetate, propionate, and butyrate were higher in calves weaned early. Difference between treatments was greatest during wk 5 to 8, after early calves had been weaned. Total VFA and acetate were both highly correlated with grain intake (r = .77), whereas propionate (r = .47) and butyrate (r = .56) were less highly correlated. Data indicate that blood VFA responded rapidly to dry feed intake, and adaptation to high grain diets was complete by 1 to 2 wk postweaning.
This study investigated the effects of amounts of RDP and branched-chain VFA on milk production and DMI by 32 early lactation Holstein cows fed diets based on corn silage and corn. All supplemental dietary protein was supplied by animal protein by-products and urea. Hydrolyzed feather meal and ring-dried blood meal served as sources of supplemental protein and were fed in a 3:1 ratio on a N basis. The experimental design was a completely randomized design with a 2 x 2 factorial arrangement of treatments. Main factors were percentage of RDP (8.0 vs. 9.5% of dietary DM) and amount of branched-chain VFA in the diet (0 vs. 90 g/d per cow). Urea was used to adjust the amount of degradable CP. Individual DMI, milk production, and milk composition were monitored during wk 5 to 19 of lactation. Ruminal fluid and blood were collected to examine the treatment effects on ruminal VFA patterns and plasma urea N concentrations. The DMI, total milk production, and milk component yield were unaffected by treatments. The molar percentages of isobutryate, isovalerate, and n-valerate increased when branched-chain VFA were fed, and concentrations of urea N in plasma increased with higher percentages of RDP. A combination of feather meal and blood meal can be used as supplemental protein to support high milk production (> 37 kg/d) in early lactation. No production benefits were observed by increased dietary RDP or branched-chain VFA.
The objectives of this study were, first, to determine the relationship between hepatic glucose release and milk production and, second, to determine the relationship between net hepatic uptake of gluconeogenic precursors and milk production. Nine multiparous ewes were individually penned and fed an alfalfa hay-based diet for ad libitum intake. Catheters were surgically placed in the portal vein, a branch of the hepatic vein, a mesenteric vein, and the abdominal aorta. Metabolite fluxes across the portal-drained viscera and liver were subsequently measured at 1, 3, 6, and 10 wk after parturition. Net hepatic glucose release, net hepatic lactate uptake, and net hepatic propionate uptake increased with increased milk production. Hepatic oxygen consumption increased with increased net hepatic glucose release. Net hepatic glucose release increased with increased hepatic propionate uptake and tended to increase with increases in metabolized amino acid and lactate uptakes. The observed increases in oxygen consumption by the portal-drained viscera with increased milk production were probably the result of increased nutrient flux. Increased hepatic oxygen consumption with increased milk production was probably due to increased glucose and urea synthesis.
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The ability of Salmonella typhimurium to invade the intestinal mucosal cells is an important step in pathogenesis. This invasion process requires genes encoded on the Salmonella pathogenicity island 1 (SPI1). Two transcriptional activators, HilA and InvF, encoded in SPII regulate the expression of invasion genes in response to environmental stimuli such as osmolarity, oxygen tension, and pH. During its pathogenic life cycle, Salmonella typhimurium is also exposed to short-chain fatty acids (SCFA), especially acetate, propionate, and butyrate, in the intestinal lumen, as well as the SCFA used as food preservatives. The effects of SCFA on the expression of hilA and invF-lacZY transcriptional fusions were examined to determine the potential role of SCFA in the pathogenesis of Salmonella typhimurium. Growth rates were reduced by increasing SCFA concentrations at pH 6 but not at pH 7. At pH 7, hilA and invF expression was induced by acetate but not by propionate or butyrate, while at pH 6, all SCFA induced hilA and invF expression at 1 h. In general, hilA and invF expression levels when compared to respective control responses were higher at 1 h than at 4 and 8 h in the presence of most SCFA concentrations at pH 6. However, expression levels at 4 and 8 h were either similar or higher than the 1-h responses for the hilA-lacZY fusion strain in the presence of acetate while exposure to 20 mM propionate yielded similar levels of expression at 1, 4, and 8 h. The pH-dependent manner of induction suggests that entry of SCFA into the cell was necessary for induction. We speculate that SCFA may serve as an environmental signal that triggers the expression of invasion genes in the gastrointestinal tract.
Twelve crossbred wethers, four per diet, were allotted at random as follows: (1) purified control diet; (2) purified diet plus a weekly oral dose of 1 mg sodium selenite per sheep; (3) purified diet plus a weekly oral dose of 1000 l.U. vitamin E. The wethers were maintained on the diets for six months. Weight gains during the initial 140 days of the experiment were significantly greater in selenium (Se)-supplemental sheep than in the others. The rumem liquor from the untreated sheep contained lower proportions (molar %) of acetic acid than from the Se or vitamin E-supplemented sheep and had higher butyric acid concentration than the Se-supplemented sheep. Concentrations of acetate, butyrate and valerate were higher in vitamin E than in Se-supplemented sheep. In addition, the concentration and molar percent of isovaleric acid was significantly higher in Se-supplemented sheep than in the other sheep.