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Lipid metabolism of rumen ciliates and bacteria. II. Uptake of fatty acids and lipid analysis of Isotricha intestinalis and rumen bacteria with further information on Entodinium simplex.

The total lipid and free fatty acid contents of Isotricha intestinalis, Entodinium simplex, and the rumen bacterial flora of the respective protozoa were determined. Warburg manometric data showed that the sodium salts of tributyrin, oleic, and acetic acids stimulated gas production in I. intestinalis, whereas tributyrin was stimulatory with E. simplex and less active with oleic and acetic acids. Rumen bacteria provided fatty acids produced lower manometric gaseous increases when compared with the protozoa. Volatile fatty acids were produced by I. intestinalis and rumen bacteria with tributyrin, but not with tripalmitin. Sodium oleate gave little volatile fatty acid response with I. intestinalis or rumen bacteria. Washed suspensions of I. intestinalis and rumen bacteria concentrated C(14)-labeled oleic, palmitic, stearic, and linoleic acids within the cells during short incubation periods. Autoradiographs demonstrated the conversion of C(14)-labeled oleic, palmitic, stearic, linoleic, and acetic acids in the rumen protozoa and bacterial cells.

Animals↗

Elevated intra-rumen pressure and secondary rumen contractions in sheep (Ovis aries).

1. Sheep rumens were insufflated with nitrogen to 5, 10, 15, and 20 cm water pressure and sustained at each pressure for 10 min. 2. Measurements included rumen motility, reticulorumen myoelectric activity, eructation frequency and volume, changes in tracheal pressure and rumen contraction amplitude. 3. As intra-rumen pressure increased, contractions designated as special secondary contractions appeared. 4. At a pressure of approximately 15 cm water, most of the special secondary contractions became regular secondaries; therefore, the special secondaries were called pro-secondary contractions. 5. Increased intra-rumen pressure was associated with respiratory distress. The recovery phase following, rumen insufflation was accompanied by hyperpnea.

Animals↗

Microbially corrected amino acid composition of rumen-undegraded feed protein and amino acid degradability in the rumen of feeds enclosed in nylon bags.

1. In the previous work (Varvikko & Lindberg, 1985), 15N-labelled rapeseed (Brassica napus), barley, ryegrass (Lolium perenne) and barley straw were incubated in the rumen in nylon bags for 5, 12 and 24 h and microbial nitrogen in the residues was quantified using the feed 15N-dilution method. In the present study, residual amino acids (AA) of these feeds were analysed, and microbially corrected AA of feed origin (feed AA) were estimated as the difference between total residual AA and respective microbial AA, assuming a constant AA composition for the microbial protein. 2. In barley and barley-straw residues, and also in ryegrass incubated in the rumen for 24 h, very large enrichment by microbial N and AA-N was found. The microbial enrichment was rather small in rapeseed residues and ryegrass incubated for 5 or 12 h. During the rumen incubation, feed N and AA-N (g/kg feed dry matter (DM] decreased very clearly in all the feeds, and feed and incubation time effects were always statistically significant (P less than 0.001). 3. The slow degradation of essential (E) feed AA compared with the respective non-essential (NE) AA degradation increased the proportion of feed EAA (g/kg determined feed AA) in barley and barley-straw residues. In rapeseed and ryegrass, residual feed EAA:NEAA remained very similar to the original. Branched-chain (Br) AA tended to increase proportionally in all the feed residues, suggesting these AA to be, on average, more resistant against microbial degradation in the rumen than other AA. Similarly, lysine was clearly increased in barley residues. A rumen degradation faster than the average rate caused decreased residual feed glutamic acid in rapeseed; methionine, alanine and glycine in barley; arginine and alanine in ryegrass; and methionine, asparagine and tyrosine in barley straw. Feed and incubation time effects were significant (P less than 0.05-0.001) for feed AA (g/kg determined feed AA) grouped as EAA, BrAA or NEAA, and for most individual AA, as well as for feed AA disappearance (%) and relative amounts (%) of feed AA in the respective residual AA. 4. According to present findings, AA composition of the rumen-degraded vegetable feed residues may markedly differ, either quantitatively or qualitatively (or both), from their original AA composition. When determining the feed AA composition of nylon-bag residues, the microbial error may be very large with starchy or fibrous feeds of low protein content. The microbial AA do not, however, considerably confuse the AA determination of protein-rich feeds.

Amino Acids↗

Influence of rumen ammonia concentration on the rumen degradation rates of barley and maize.

Four rumen-cannulated steers were given barley and maize diets supplemented with graded levels of an ammonium acetate solution. Animals were fed hourly from automatic feeders and water consumption was controlled to achieve steady-state conditions in the rumen. Dacron bags containing rolled barley or ground barley were incubated in the rumen of barley-fed steers, while ground maize and autoclaved maize were incubated in the rumen of maize-fed steers. Fractional degradation rates of dry matter were estimated for each cereal substrate incubated using a single-pol exponential decay model. No differences in degradation rate due to the method of feed processing were detected; however, barley was degraded at a faster rate than maize. Furthermore, the minimum rumen ammonia-nitrogen concentration required to maximize the degradation rate of barley (125 mg/l) was greater than that required to maximize the degradation rate of maize (61 mg/l). These results indicate that the optimal NH3-N concentration required to maximize the rate of grain digestion in the rumen is influenced by the chemical or structural characteristics of the grain.

Acetates↗

Influence of post-rumen supply of nutrients on rumen digesta load and voluntary intake of a roughage by sheep.

The effects of post-rumen supply of nutrients on the rumen digesta load and voluntary consumption of roughage by Border Leicester X Merino ram lambs was investigated. A chopped wheaten hay was offered to the ram lambs (n 24), whose nutrient intake was altered by infusing into the abomasum a liquid supplement, containing reconstituted cow's milk, sodium caseinate, minerals and vitamins, for a period of at least 30 d. Either two or three lambs were allocated by live weight to each of ten rates of nutrient supplementation. The lambs were slaughtered at a target live weight of 31 kg. Voluntary intake of hay was estimated over the last 7 d of feeding. The amounts of digesta and organic matter (OM) in the rumen were measured by emptying after slaughter. The particle size distribution of the digesta was measured by wet sieving and the fractional outflow rate of particulate matter by reference to the marker lignin. Growth rate of the lambs increased linearly as total energy infused increased. The linear increase in growth rate indicated that roughage was not substituted for infused nutrients on a direct energy basis. OM intake from hay declined linearly as the amount of nutrients infused increased. The decline in intake was associated with a decline in both total digesta and the amount of rumen OM. Both the fractional digestion rate of OM and the fractional outflow rate of particulate matter from the rumen were unaffected by the amount of nutrients infused. The results indicated that the rumen digesta load of young sheep fed on a single roughage is directly related to their energy deficit, that is the difference between the capacity of the animals to use energy and the energy available to them for metabolism.

Animals↗

Absorption of volatile fatty acids from the rumen of lactating dairy cows as influenced by volatile fatty acid concentration, pH and rumen liquid volume.

The effect of rumen liquid volume, pH and concentration of volatile fatty acids (VFA) on the rates of absorption of acetic, propionic and butyric acids from the rumen was examined in lactating dairy cows. Experimental solutions introduced into the emptied, washed rumen comprised two different volumes (10 or 30 l), four levels of pH (4.5, 5.4, 6.3, 7.2) and three levels of individual VFA concentrations (20, 50 or 100 mM-acetic, propionic or butyric acid). All solutions contained a total of 170 mM-VFA and an osmotic value of 400 mOsmol/l. Absorption rates were calculated from the disappearance of VFA from the rumen corrected for passage with liquid phase to the omasum. An increase in initial fluid pH caused a reduction in fractional absorption rates of propionic and butyric acids. Increasing the initial pH from 4.5 to 7.2 reduced fractional absorption rates of acetic, propionic and butyric acids from 0.35, 0.67 and 0.85 to 0.21, 0.35 and 0.28/h respectively. The fractional absorption rates of all VFA were reduced (P < 0.05) by an increase in initial rumen volume. The fractional absorption rate of acetic acid was lower (P < 0.05) at an initial concentration of 20 mM than of 50 mM. The fractional absorption rate of propionic acid tended (P < 0.10) to decrease as the level of concentration increased while fractional absorption rate of butyric acid was not affected by butyric acid concentration. These results indicate that relative concentrations of VFA in rumen fluid might not represent relative production rates and that attempts to estimate individual VFA production from substrate digestion must take account of pH and VFA concentration.

Acetates↗

Effect of defaunation and refaunation of the rumen on rumen fermentation and N-flow in the duodenum of sheep.

In order to confirm earlier fragmentary results, the effect of defaunation and refaunation of the rumen on the fermentation pattern and flow of N-components in the proximal duodenum of two sheep was investigated. Defaunation had no effect on acetic acid as a proportion of the total volatile fatty acids in the rumen, while the proportions of propionic acid increased with a concomitant decrease in butyrate. Refaunation resulted in lower acetic acid and higher butyric acid proportions. The concentration of ammonia N in the rumen was clearly decreased after defaunation, already indicating an effect of the elimination of protozoa on nitrogen metabolism in the rumen. Defaunation also increased significantly the flow of total N, non ammonia N and individual and total amino acids in the proximal duodenum. Defaunation resulted in higher bacterial growth efficiency, significantly in one sheep, but the decrease after refaunation was statistically significant for both sheep. Determination of rumen digestibility of organic matter and acid detergent fibre (ADF) revealed lower values in the absence of the protozoa, while total digestibility was only influenced to a much lower extent. This indicated a shift of digestion from rumen to the lower digestive tract. Finally, earlier work is discussed in the light of the present findings.

Ammonia↗

Influence of dietary niacin on volatile fatty acids in rumen liquid of sheep and rumen dry matter degradability of untreated and ammonia-treated wheat straw.

Three rumen fistulated sheep were fed with roughage (1200 g artificially dried grass) or a concentrate/roughage diet (600 g concentrate, 400 g artificially dried grass per animal and day). The diets were unsupplemented or 0.5, 1 and 2 g niacin per animal and day were added. After a 14-day adaptation period samples of rumen liquid were taken 0, 1, 2, 3, 4 and 6 h after morning feeding. In a second experiment untreated and NH3-treated wheat straw were incubated in nylon bags for 6, 12, 24, 48 and 72 h in the rumen of 5 sheep fed with 1.2 kg artificially dried grass and unsupplemented or supplemented with 1 g niacin per day. Niacin did not significantly influence the total concentration of volatile fatty acids, acetate and branched chain fatty acids in both types of diet (P less than 0.05). 0.5 and 1 g niacin in the roughage and 0.5 g niacin in concentrate/roughage diet increased the propionic acid concentration and reduced the butyric acid concentration in rumen liquid of sheep (P less than 0.05). Niacin resulted in a small decrease (P greater than 0.05) of dry matter degradability in rumen. After 48 h incubation time, the dry matter disappearance of untreated and NH3-treated wheat straw amounted to 35.4 and 46.2% resp. The effects of niacin on rumen fermentation may be useful in avoiding ketogenic situations in dairy cows.

Ammonia↗

Effect of dietary choline on volatile fatty acids in rumen liquid of sheep and rumen dry matter degradability of untreated and ammonia-treated wheat straw.

Three rumen fistulated wethers consumed a roughage (1200 g artificially dried grass) or a concentrate/roughage diet (600 g concentrate, 400 g dried grass per animal and day). The diets were unsupplemented or 0.5, 1 or 2 g choline per animal and day were added. After a 14-day feeding period, samples of rumen liquid were taken 0, 1, 2, 3, 4 and 6 h after morning feeding. In a second experiment, untreated and ammonia-treated wheat straw were incubated in nylon bags for 6, 12, 24, 48 and 72 h in rumen of 5 sheep fed on 1.2 kg dried grass and without or with 1 g choline per day. Choline supplementation decreased the total concentration of volatile fatty acids in rumen liquid in both types of diet (roughage: 106, 83, 89 and 87; concentrate/roughage: 142, 114, 91 and 106 mmol/l for 0, 0.5, 1 and 2 g choline per day resp.). Concentration of acetate and branched chain fatty acids in both diets and butyrate in concentrate/roughage diet was reduced as total acids. Butyrate in roughage diet was increased by added choline. Molar propionate concentration was not significantly influenced by choline (P greater than 0.05). Choline resulted in a little increase (P greater than 0.05) of dry matter degradability in rumen, especially when NH3-treated straw was incubated. The effect of choline on milk fat production of cows fed with diets rich in concentrate is probably not related to changes in rumen fermentation and digestion.

Acetates↗

Medium without rumen fluid for nonselective enumeration and isolation of rumen bacteria.

Colony counts which approximated those in a habitat-simulating, rumen fluid-agar medium (RFM) were obtained in medium 10, a medium identical to the RFM except for the replacement of rumen fluid with 1.5 x 10(-6)m hemin, 0.2% Trypticase, 0.05% yeast extract, and a 6.6 x 10(-2)m volatile fatty acid mixture qualitatively and quantitatively similar to that in rumen fluid. Single deletion of Trypticase, yeast extract, or the volatile fatty acid mixture from medium 10 significantly reduced colony counts. Colony counts were also reduced when medium 10 was modified to contain higher concentrations of Trypticase or volatile fatty acids. Significant differences were found between colony counts obtained from diluted rumen contents of animals fed a cracked corn-urea diet, and the colony counts obtained from animals fed either a cracked corn-soyean oil meal or an alfalfa hay-grain diet. Qualitative differences were found between the predominant bacterial strains isolated from rumen contents of animals fed cracked corn diets and strains isolated from animals fed alfalfa hay-grain. Regardless of differences in the predominant flora associated with diet, medium 10 and the RFM supported growth of similar bacterial populations. The results show that medium 10 is suitable for enumeration and isolation of many predominant rumen bacteria.

Animal Feed↗

Relationship of rumen fluid dilution rate to rumen fermentation and dietary characteristics of beef steers.

Data from seven beef steer trials were compiled and regression analyses used to evaluate relationships among molar proportions of acetate (Ac), propionate (Pr) and butyrate (Bu), total concentration of volatile fatty acids (VFA), rumen ammonia (NH3), rumen pH, rumen fluid dilution rate (FDR), rumen fluid volume (FVOL), body weight (WT), dry matter intake (DMI) and dietary concentration and intake of crude protein (CP and CPI), acid detergent fiber (ADF and ADFI), ash (ASH and ASHI) and metabolizable energy (ME and MEI). Of the six fermentation variables, Pr (negative regression coefficient, beta) and pH (positive beta) were related (P less than .05) to FDR, but only 3 and 12% of the variation in these two variables, respectively, was explained by FDR. When FDR was described by dietary characteristics, ASHI was positively related to FDR (R2 = .16). The best two-variable model for FDR contained DMI (positive beta) and WT (negative beta) and increased R2 to .36. Fluid volume was best described by ME (positive beta; R2 = .20). The two-variable model for FVOL added ASH with a positive partial beta (R2 = .23). When fermentation variables were regressed on dietary characteristics, Ac was best described by ADF (positive beta; R2 = .71). The variable that best described Pr proportion was ADF (negative beta; R2 = .50), and addition of CP (negative beta) and MEI (positive beta) into the Pr model improved R2 to .70. Molar proportion of butyrate was related to CP (positive beta; R2 = .23), and addition of ME (positive beta) to the model improved the R2 to .31. Total VFA concentration was best described by ADFI (positive beta; R2 = .14). An R2 of .29 resulted when ME (positive beta) and CPI (negative beta) were included in the total VFA model. Rumen pH was related to ADF (positive beta; R2 = .45), and addition of CP (positive beta) to the rumen pH model increased R2 to .55. Crude protein concentration was related to ruminal NH3 level (positive beta; R2 = .42), and inclusion of ADFI (positive beta) into the model improved the R2 to .47.

Ammonia↗

Thiomolybdates in rumen contents and rumen cultures.

Examination of direct and (Cu)-difference spectra of i) the aqueous supernatants of in vitro cultures of bovine rumen contents incubated with MoO42- and potential sources of S2- and ii) samples drawn directly from the rumen of animals receiving high Mo diets yielded evidence of the presence of thiomolybdates. Only MoS42- was detected in the soluble phase of in vitro cultures. Although intense and variable background absorbance precluded full characterization of thiomolybdate species in samples drawn directly from the rumen, both spectral data and the biochemical and clinical responses of animals given high Mo diets were consistent with the conclusion that MoS42- rather than MoOS32- was the predominant thiomolybdate species present in the aqueous phase. Addition of Ca2+ either to rumen cultures before incubation or as a supplement to diets high in MoO42- content inhibited the appearance of MoS42- in the aqueous phase. Evidence of the sequestration of MoS42- and MoOS32- by particulate or microbial fractions of rumen contents is considered in relation to the inhibitory action of Mo upon Cu absorption by ruminants.

Animals↗

Effect of level and origin of rumen degradable nitrogen on rumen microbial growth and nitrogen utilisation efficiency of animals fed maize silage at maintenance.

Isoenergetic maize silage diets, fed at maintenance to 24 suckling cows (Exp. 1) and to 3 (Exp. 2) or 4 (Exp. 3) rumen fistulated sheep, were compared with a urea and controlled release NPN diet (Exps. 1 and 2) and with a protein-N supplemented diet (Exp. 3). Supplementation increased blood urea concentrations (44.7 +/- 22.3 vs. 97.6 +/- 23.7 mg urea-N x L(-1)) (Exp. 1) and renal urea excretion (2.5 +/- 1.1 vs. 7.6 +/- 1.8 g urea-N x d(-1)) (Exps. 2 and 3), whereas blood allantoin concentrations (286.7 +/- 77.0 micromol x L(-1)) (Exp. 1) and renal excretion of purine derivatives (357.6 +/- 90.7 mg purine-N x d(-1)) (Exps. 2 and 3) were not affected, indicating additional N supplementation did not improve rumen microbial growth. However, some deficiency of rumen degradable N might occur in non supplemented diets as suggested by the reduced rumen NH3-N and RNA concentrations (868 +/- 270 vs. 1466 +/- 466 mg RNA x kg(-1) rumen contents).

Animals↗

Hydrolysis of 14C-labelled proteins by rumen micro-organisms and by proteolytic enzymes prepared from rumen bacteria.

Proteins were labelled with 14C in a limited reductive methylation using [14C]formaldehyde and sodium borohydride. The rate of hydrolysis of purified proteins was little (less than 10%) affected by methylation and the 14C-labelled digestion products were not incorporated into microbial protein during a 5 h incubation with rumen fluid in vitro. It was therefore concluded that proteins labelled with 14C in this way are valid substrates for study with rumen micro-organisms. The patterns of digestion of 14C-labelled fish meal, linseed meal and groundnut-protein meal by rumen micro-organisms in vitro were similar to those found in vivo. The rates of hydrolysis of a number of 14C-labelled proteins, including glycoprotein II and lectin from kidney beans (Phaseolus vulgaris), were determined with mixed rumen micro-organisms and with proteases extracted from rumen bacteria. Different soluble proteins were digested at quite different rates, with casein being most readily hydrolysed. Proteins modified by performic acid oxidation, by cross-linking using 1,6-di-iso-cyanatohexane or by diazotization were labelled with 14C. Performic acid treatment generally increased the susceptibility of proteins to digestion, so that the rates of hydrolysis of performic acid-treated proteins were more comparable than those of the unmodified proteins. Cross-linking resulted in a decreased rate of hydrolysis except with the insoluble proteins, hide powder azure and elastin congo red. Diazotization had little effect on the rate of hydrolysis of lactoglobulin and albumin, but inhibited casein hydrolysis and stimulated the breakdown of gamma-globulin.

Animals↗

The nutritive value of rumen micro-organisms in ruminants. 1. Large-scale isolation and chemical composition of rumen micro-organisms.

A method is described whereby a large quantity of rumen microbial dry matter of high purity was isolated from whole rumen contents obtained from abattoirs, by means of a continuous process of one filtration through four sieves followed by three differential centrifugations. The contents of ash, carbohydrate, lipid, nitrogen, RNA, DNA and individual amino acids of the three centrifugal fractions are given and compared with values summarized from more than sixty published reports on the chemical composition of rumen micro-organisms isolated from both whole rumen contents and pure cultures. The amino acid composition of isolated rumen micro-organisms, in particular that of the bacteria, was found to be remarkably constant.

Amino Acids↗

Relations between passage rates of rumen fluid and particulate matter and foam production in rumen contents of cattle fed on different diets ad lib.

1. A group of six cattle, three of which had a non-bloating history (group A) and had been ruminally cannulated for the previous 2 years, and three with a history of being bloat-prone (group B) and which had been ruminally cannulated only 3 months before the study, were fed ad lib. on chopped lucerne (Medicago sativa) hay, lucerne pellets, or a 100 g chopped hay and 900 g rolled barley grain/kg diet over three periods of 30 d each. Flow of rumen digesta, by reference to CoEDTA and chromium-mordanted fibres, and foam production from samples of rumen contents were measured. 2. Samples of rumen contents (50 ml) from group A produced foam heights of 150 and 60 mm, 2 and 4 h after feeding respectively, compared with 240 and 150 mm for group B (P less than 0.05). 3. The fractional passage rate of the 1-2 mm particles mordanted with Cr did not differ (P greater than 0.05) between groups. 4. The fractional outflow rates (FOR) for CoEDTA 0-2 h and 2-7 h after feed was offered were 0.205 and 0.160/h for group A and 0.093 and 0.086/h for group B respectively (P less than 0.05). 5. Rumen-fluid FOR 0-2 h and 2-7 h after provision of feed were significantly (P less than 0.05) inversely correlated (r -0.74 and -0.85 respectively) with the amount of foam produced from rumen contents at these times.

Animal Feed↗

Studies of method of conserving grass herbage and frequency of feeding in cattle. 2. Eating behaviour, rumen motility and rumen fill.

The influence of the method of conserving grass herbage and the frequency of feeding on eating behaviour, rumen motility and rumen fill was studied in growing steers. Silage and hay were offered to twelve rumen-cannulated Friesian steers (average initial live weight (LW) 128 kg) at a restricted level of intake (20 g dry matter (DM)/kg LW) either once or eight times daily. With once daily feeding, the daily intake of hay was consumed in a single large meal which lasted about 2 h, while silage was eaten in many small meals throughout the day. The proportion of the day spent ruminating was higher (0.39) for steers offered hay once daily than for those offered silage (0.28) or hay eight times per day (0.29). There was little effect of conservation method on frequency of rumen contraction, but contractile intensity (integration of frequency and amplitude) increased during the second half of the 24 h feeding cycle in steers offered hay either once daily or eight times daily, while the same effect was only observed when silage was offered once daily. Total daily contractile activity (the integration of the frequency and duration of myoelectric spike bursts) was significantly (P less than 0.05) higher in steers offered hay, compared with silage, once daily. The mean weight of organic matter in the reticulo-rumen of steers offered hay was significantly higher than for silage both with once (1593 v. 1326 g organic matter (OM)/100 kg LW, P less than 0.01) and eight (1367 v. 1160 g OM/100 kg LW, P less than 0.05) times daily feeding. The proportion of the particulate pool which was present as small (less than 1.2 mm) particles was always greater than 0.60.

Animals↗

Responses in the voluntary intake of hay or silage by lactating cows to intraruminal infusions of sodium acetate or sodium propionate, the tonicity of rumen fluid or rumen distension.

Rumen-fistulated lactating cows were individually fed on hay or silage and intakes were monitored during 3 h treatment periods and for 2 h after. Each experiment used five, six or seven animals and the treatments were applied in a Latin Square design. Sodium acetate infusions of 1.8-11.0 mol in 4.5 litres water caused a dose-related depression in hay intake, the extent being 82 g dry matter (DM)/mol infused (P < 0.01). Sodium acetate infusions of 6.0-15.0 mol in 4.5 litres water caused a dose-related depression in silage intake of 118 g DM/mol infused. Rumen fluid pH for both diets was unaffected by treatment. Acetate and Na concentrations were increased and significantly negatively correlated with intake of both diets. Infusions of 2-8 mol sodium propionate caused a dose-related depression of hay intake which was significant when cow and day effects were accounted for. Sodium propionate infusions of 4-8 mol significantly depressed silage intake by 140 g DM/mol infused (P < 0.001). Rumen fluid pH was unaffected by treatment while propionate and Na concentrations were elevated and significantly negatively correlated with intake for both diets. Inflation of a rubber balloon in the rumen with 12.5-20 litres warm water resulted in a dose-dependent depression in hay intake of 66 g DM/l distension (P < 0.05). There was significant overeating during the 2 h following the 20 litre treatment. With silage, 15-25 litres of balloon distension for 3 h resulted in a dose-dependent depression in intake of 28 g DM/l distension (P < 0.001). There was no significant overeating during the 2 h following distension. When given in physiological amounts, at the lower end of the range used in these experiments, acetate, propionate and distension of the rumen did not significantly affect hay intakes. However, in each case the linear relationship between intake depression and level of treatment suggested that these factors could contribute to the control of feed intake.

Acetates↗