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[Protein digestibility and the absorption of amino acids in various segments of the digestive tract of pigs. 4. Digestibility of crude protein and amino acids and rate of passage through the duodenum and ileum and the total digestive tract of growing pigs].

The crude protein digestibility and the amino acid absorption of six female pigs (average live weight 61 kg) with duodenal and ileocecal re-entrant cannulae, which were fed with a wheat + wheat gluten + lysine ration and an N-free ration, were determined in various segments of the intestines. Comparative experiments concerning the N-metabolism with pigs without and with cannulae showed that the animals renormalised their metabolism 14 days after narcosis and fistulation of the intestines. The extents of secretion and absorption of the various amino acids vary as it is shown from the values of the apparent and true digestibility resp. rate of passage through various segments of the intestines. While for methionine and glutamic acid absorption exceeds endogenous secretion already in the duodenum, the amino acids with a high endogenous quota (glycine, alanine, threonine, tryptophan) are, even at the terminal ileum, not as well absorbed as the others. Methionine is obviously synthesised on a large scale by the colon flora and excreted in feces. The fractionation of the duodenal and ileum chyme after feeding wheat + wheat gluten + lysine as well as N-free mixture, into the fractions "solid particles", "peptides-free amino acids" and "proteines precipitable by trichlor-acetic acid" supplies information on the degree of protein degradation in various segments of the intestines.

Amino Acids↗

[Protein digestibility and absorption of amino acids in various segments of the digestive tract of pigs. 2. Protein and amino acid balances at the end of the small intestine and of the whole digestive tract (apparent and true protein and amino acid digestibility)].

Growing pigs with ileum and ileocecal re-entrant canulae were given 5 different rations (pig fattening feed 1 and 2, rations with dried skim milk, wheat gluten + lysine and wheat + wheat gluten + lysine). The crude protein and amino acid excretion was quantitatively determined with ileum chyme and feces. The calculation of the amino acid balances (apparent and true digestibility) at the end of the small intestine, at the end of the wholedigestive tract and the isolated colon showed characteristic differences for the individual amino acids. On ileum level the amino acids with relatively high endogenous quotas, threonine, tryptophane, alanine, aspartic acid and glycine showed a distinctly lower, the amino acids arginine, methionine, phenylalanine, glutamic acid, proline and tyrosine a distinctly higher digestibilityhan crude protein. In comparison of the values on the feces level, including the true digestibility, these differences are largely balanced out. At the end of the small intestine the absorption of the amino acids is in the main completed. The disappearance rate of amino acids from the colon shows large differences with reference to the different rations as well as the individual amino acids. They were particularly high for various rations as far as proline, tryptophane, glycine and cystine are concerned. If wheat gluten + lysine rations were given, a net synthesis of methionine in the colon could be proved.

Amino Acids↗

Digestion, absorption and utilization of single-cell protein by the preruminant calf. The true digestibility of milk and bacterial protein and the apparent digestibility and utilization of their constituent amino acids.

Two experiments of Latin square design were made, each with four Friesian bull calves fitted with re-entrant duodenal and ileal cannulas at 4-10 d of age. The calves were used to study the effect of giving milk-substitutes containing 0, 300, 500 and 700 g bacterial protein (Pruteen)/kg total protein on apparent digestibility of nitrogen fractions and amino acids and true digestibility of 3H-labelled milk protein and 35S-labelled bacterial protein in the small intestine. A third experiment of Latin square design with four intact Friesian calves was used to measure apparent digestibility of nutrients throughout the alimentary tract and retention of N, calcium and phosphorus. At the duodenum, volume of outflow, its pH, and outflow of total-N(TN), protein-N (PN) and non-protein-N (NPN) decreased with time after feeding. At the ileum, volume of outflow and TN outflow were unaffected by time after feeding but PN outflow decreased; NPN outflow at the ileum increased to a maximum 6 h after feeding and then declined. Increased inclusion of Pruteen did not affect the volume of outflow at the duodenum or ileum, but duodenal PN outflow increased. At the ileum, pH values were lower and TN, PN and NPN outflows were higher with increasing concentration of Pruteen in the diet. Apparent digestibility in the small intestine tended to decrease with greater amounts of Pruteen, but was only significant for NPN. Apparent digestibility from mouth to ileum significantly decreased for TN and PN as Pruteen inclusion increased. Amino acid concentration in duodenal outflow, with the exception of that of arginine, reflected intake. The total amount of each amino acid in ileal outflow increased and the apparent digestibility of most amino acids decreased with greater amounts of Pruteen in the diet. Apparent digestibility of nucleic acid-N from Pruteen was very high. True digestibility in the small intestine and between mouth and ileum of 3H-labelled milk protein was high and did not differ between dietary treatments. True digestibility of 35S-labelled Pruteen was low for the milk-protein diet and tended to increase linearly as more dietary Pruteen was included. Dry matter concentration in faeces and a high apparent digestibility throughout the whole alimentary tract of carbohydrates did not differ between treatments.(ABSTRACT TRUNCATED AT 400 WORDS)

Absorption↗

[Digestion of pea and soya proteins in the preruminant calf. II. Apparent digestibility at the end of the ileum and the digestive tube].

Three milk substitutes (control, pea and soya-bean) were given to 6 preruminant calves fitted with re-entrant canulas in the terminal ileum. In the control diet, protein was almost entirely provided by skim milk powder. In the pea diet, a pregelatinized dehulled pea flour provided 33.5% of the protein, the remainder being supplied by skim milk powder. In the soya-bean powder diet 73.2% of the protein were provided by a soya-bean isolate and the remainder by whey powder. The apparent digestibility of total nitrogen was significantly lower with the pea and soya-bean diets. than with the control diet (0.92, 0.91 and 0.95 at the end of the ileum, 0.92, 0.94 and 0.97 at the end of the whole digestive tract, respectively). Also the ileal digestibility of most amino acids decreased with the pea and soya-bean diets; the differences were greatest for cystine (-0.6; non significant) with the pea diet and for threonine (-0.7; P less than 0.01) with the soya-bean diet. Irrespective of the diet the protein escaping digestion in the small intestine appeared to be mainly from endogenous and bacterial origin. Although small amounts of partially degraded dietary protein could be present in ileal digesta with the pea and soya-bean diets, their true digestibility was probably very high.

Amino Acids↗

Dietary fiber viscosity and amino acid digestibility, proteolytic digestive enzyme activity and digestive organ weights in growing rats.

The effect of dietary fiber viscosity on apparent ileal nitrogen and amino acid digestibility, proteolytic enzyme activity and digestive organ weights was investigated. Eighteen growing rats were fed for 21 d purified casein-based diets containing carboxymethylcellulose (50 g/kg) of low (20 cP), medium (800 cP) and high (2000 cP) viscosity (LV, MV and HV treatment groups, respectively). Dietary fiber viscosity did not significantly affect apparent ileal (terminal 15 cm of the ileum) nitrogen or amino acid digestibility, trypsin or chymotrypsin activity in the small intestinal contents and pancreatic tissue, aminopeptidase-N activity in the small intestinal contents and tissue, or the weights of the stomach, pancreas, small or large intestines. Intragastric pepsin activity in LV rats was significantly higher than in MV or HV rats (P < 0.01), but fiber viscosity did not affect pepsin activity in the stomach tissue. The intragastric pH of the HV and MV rats was significantly higher than that for the LV rats (P < 0.01). The stomach contents (dry matter) of MV and HV rats were greater than in LV rats (P < 0.05). Delayed passage rate of the more viscous digesta may have resulted in greater absorption of amino acids, because the HV rats had a higher estimated true ileal digestibility than the LV animals for threonine, serine, aspartic acid, glutamic acid, histidine, tyrosine and phenylalanine.

Amino Acids↗

Stunting syndrome in broilers: effect of age and exogenous amylase and protease on performance, development of the digestive tract, digestive enzyme activity, and apparent digestibility.

Day-old male, meat-type chicks raised in brooder batteries were infected by orally administering an inoculum prepared from intestines of broiler chicks infected with stunting syndrome (SS). Naive controls were kept in a parallel room. The chicks were fed a commercial starter diet supplemented with two levels of enzyme preparations to 14 d of age. The experiment was continued to the age of 6 wk in order to estimate compensatory feed intake and growth. In a parallel study, digestibility of the feed was determined from 1 to 3 wk of age with control or inoculated chicks. The enzymes amylase and proteases were produced by Bacillus subtilis and Penicillium emersonii. Enzyme supplementation had no effect on feed intake, growth, or feed utilization, or on digestibility of fat, starch, protein, or energy. Because enzyme supplementation did not consistently affect performance of chicks and no interactions were observed between enzyme supplementation and infection status, data are presented for effects of infection only. Inoculation of SS-infective material reduced performance to 4 wk. Compensatory growth and feed intake were observed from the age of 4 wk onward. At the age of 6 wk the slight retardation of the inoculated chicks was not significant. On Week 1, retention of fat, starch, protein, and energy was significantly depressed in the inoculated chicks. At the age of 2 wk, retention of starch was not depressed, and at the age of 3 wk, the only consistent depression was that observed for fat. The proventriculus weight and content were consistently higher in inoculated chicks, as were the small intestine and intestinal content. The pH of the gizzard content was higher, and that of the small intestine content was lower, in the inoculated birds than in their control counterparts. Stunting syndrome infection was accompanied by a significant depression of trypsin activity in the pancreas at the age of 1 and 2 wk. At these periods, amylase and chymotrypsin were not affected. At 6 wk of age, the activities of amylase, trypsin, and chymotrypsin in the pancreas were higher in the inoculated than in the control birds. In the intestinal chime, amylase, trypsin, and chymotrypsin activities were lower in the inoculated birds on Week 1 and 2 (NS for amylase on Week 1). On Week 6, the activity of all enzymes assayed was higher in the inoculated birds (NS for amylase). It is suggested that the main factors depressing feed intake and growth in SS-infected birds are most probably beyond those of digestion.

Aging↗

[Protein and amino acid metabolism in the digestive tract of growing young bulls. 4. Apparent digestibility of NH3-free raw protein reaching the duodenum and critical evaluation of apparently digestible raw protein].

The experimental determination of the apparent digestibility (AD) of the NH3-free crude protein at the duodenum (NH3 fr. CPD) showed a value of 65.7% at a standard error of +/- 3.6% for the section of the digestive tract between proximal duodenum and end of rectum. This standard error can only be insignificantly diminished due to the existing relation between AD of NH3 fr. CPD in dependence on the CP concentration in the ration (y = 56.4 + 0.598 x +/- 3.17; r = 0.501xx) so that this relation has no practical significance. The criticism of AD CP is correct from a point of view of digestion physiology. There is only agreement between AD PC and AD NH3 fr. CPD in a CP concentration range between 12.5 and 14.0% CP in the DM, which, however, applies to a large range of the production sphere in the rearing of young bulls. It excludes, however, the low live weight range with high live weight gains.

Amino Acids↗

Digestive development of the early-weaned pig. 1. Effect of continuous nutrient supply on the development of the digestive tract and on changes in digestive enzyme activity during the first week post-weaning.

Gastric intubation was adopted to examine the effect of continuous nutrient supply on digestive development of the pig during the immediate post-weaning period. The 14 d-weaned animals were slaughtered at 3, 5 and 7 d post-weaning (3W, 5W and 7W respectively) and the suckled animals were slaughtered at 14 and 22 d of age (14SR and 22SR respectively). The weight of the pancreas (g/kg bodyweight) was significantly greater (P less than 0.05) in the 5W and 7W groups, as was the weight of large intestine (g/kg) in all weaned groups (P less than 0.01) compared with sow-reared pigs. The stomach weight (g/kg) tended to be greater in the weaned groups. Weaning, in conjunction with a continuous nutrient supply, did not significantly alter the time-related changes in the weight of the small intestine (SI) or the SI mucosa, although both variables tended to be lowest in the 3W group. However, there was a 20% reduction in the protein content of the mucosa within the first 3 d post-weaning (P less than 0.01) which persisted during the 7 d experimental period. Lactase, (beta-galactosidase; EC 3.2.1.23) activity (mumol/g protein and mol/d) of the 7W group was reduced to approximately 40% of the 22SR value. Hence, continuous nutrient supply may have delayed, but did not prevent, the loss of lactase activity at weaning. The activity of sucrase (sucrose-alpha-glucosidase; EC 3.2.1.48) was significantly higher in 22SR compared with 14SR animals. Sucrase activity in weaned pigs was intermediate to the values for sow-reared pigs whereas maltase (alpha-glucosidase; EC 3.2.1.20) and glucoamylase (glucan 1,4-alpha-glucosidase; EC 3.2.1.3) were significantly increased in relation to their sow-reared counterparts. Continuous nutrient supply did not prevent the reduction in villous height and the crypt hypertrophy associated with weaning. The results of the present study suggest that there may be some degree of interaction between nutrient intake and gut development during the immediate post-weaning period but that there is also a component of the adaptive response which is independent of nutrient intake. They confirm the rapid substrate induction of the brush-border glucoamylases and indicate the importance of considering total as well as specific enzyme activity for satisfactory interpretation of changes in digestive function.

Animal Nutritional Physiological Phenomena↗

Refining in vitro digestibility assays: fractionation of digestible and indigestible peptides.

Typically, in vitro methods used for estimating the amount of ileal digestible AA do not exhaustively digest samples, and arbitrary methods for separating digestible from indigestible protein are used. This may lead to over- or underestimation of digestibility coefficients. A method that exhaustively digests proteins using pepsin and pancreatin was developed, and the first objective of this research was to confirm that exhaustive digestion was indeed appropriate and to determine the fractionation method for separating digestible from indigestible proteins. For this, three homoarginine-labeled animal proteins were prepared. Samples were subsequently digested in vivo and in vitro to determine which fraction should be considered indigestible, and in vitro followed by in vivo to determine whether the extent of digestion in vivo was improved by predigestion. In vivo, soluble but unabsorbed peptides were smaller than 1 kDa, suggesting that the size of soluble peptides is not what prevents their absorption. Thus, all in vitro-soluble proteins should be considered digestible. In vitro, 88 +/- 3% of the soluble peptides were smaller than 1 kDa, with the remainder between 1 and 5 kDa, suggesting that in vitro digestion is less complete. Predigested samples were digested in vivo to the same size distribution as the nonpredigested samples. The second objective was to test whether in vitro digestibility assays based on these principles equaled in vivo digestibility. For this, digestibility data for 25 animal proteins were compared. Results showed a lack of correlation between lysine digestibility coefficients; however, across samples, the extent of digestion did not differ for lysine (P = 0.71), threonine (P = 0.26), methionine (P = 0.18), or valine (P = 0.66), whereas in vitro digestibility coefficients were lower for (the less water-soluble) histidine (P = 0.05), isoleucine (P < 0.01), leucine (P < 0.01), and phenylalanine (P = 0.05). In conclusion, in vitro digestibility assays should exhaustively digest proteins to mimic in vivo digestibility. All in vitro-soluble peptides could be considered digestible, because in vivo, no large soluble peptides were observed whose size prevented them from being absorbed. However, an in vitro assay based on these principles lacked precision for highly water-soluble AA, and underestimated digestibility for other AA. Better solubilization of the digesta and more replicates may improve the in vitro assay further.

Amino Acids↗

In vitro starch digestion correlates well with rate and extent of starch digestion in broiler chickens.

Current feed evaluation systems for poultry are based on digested components (fat, protein and nitrogen-free extracts). Digestible starch is the most important energy source in broiler chicken feeds and is part of the nitrogen-free extract fraction. Digestible starch may be predicted using an in vitro method that mimics digestive processes in the gastrointestinal tract of broiler chickens. An experiment was designed to use this method for predicting site, rate and extent of starch digestion in broiler chickens. In vitro starch digestion was studied in 12 experimental diets differing in starch sources. These diets were also used in a digestibility trial with broiler chickens. Correlations between in vitro and in vivo starch digestion were calculated. Starch digestion after 2 h incubation correlated well with in vivo starch digestion in the first half of the small intestine (r = 0.94). A 4-h incubation period resulted in a good correlation between in vitro starch digestion and ileal starch digestion (r = 0.96). In vitro starch digestion rate (h(-1)) correlated well with in vivo starch digestion rate (r = 0.87). In vitro starch digestion of individual starch sources was additive. It appeared that legume seeds and waxy corn contained two starch fractions, which were digested at different rates. We conclude that starch digestion rate in broiler chickens is well predicted by the in vitro method.

Animal Feed↗

Pelleted beet pulp substituted for high-moisture corn: 2. Effects on digestion and ruminal digestion kinetics in lactating dairy cows.

The effects of increasing concentrations of dried, pelleted beet pulp substituted for high-moisture corn on digestion and ruminal digestion kinetics were evaluated using eight ruminally and duodenally cannulated multiparous Holstein cows in a duplicated 4 x 4 Latin square design with 21-d periods. Cows were 79 +/- 17 (mean +/- SD) d in milk at the beginning of the experiment. Experimental diets with 40% forage (corn silage and alfalfa silage) and 60% concentrate contained 0, 6.1, 12.1, or 24.3% beet pulp substituted for high-moisture corn on a dry matter basis. Diet concentrations of neutral detergent fiber (NDF) and starch were 24.3 and 34.6% (0% beet pulp), 26.2 and 30.5% (6% beet pulp), 28.0 and 26.5% (12% beet pulp), and 31.6 and 18.4% (24% beet pulp), respectively. Ruminal dry matter pool decreased and NDF turnover rate increased as dietary beet pulp content increased. Potentially digestible NDF was digested more extensively and at a faster rate in the rumen with increasing beet pulp, resulting in increased total tract NDF digestibility. Passage rates of potentially digestible NDF and of indigestible NDF were not affected by treatment. True ruminal digestibility of starch decreased with increasing beet pulp substitution. This was caused by a linear increase in starch passage rate, possibly because of increasing ruminal fill, and a linear decrease in digestion rate of starch in the rumen, possibly because of reduced amylolytic enzyme activity for lower-starch diets. Although true ruminal starch digestibility decreased when more beet pulp was fed, whole tract starch digestibility was not affected because of compensatory digestion of starch in the intestines. Due to more thorough digestion of fiber in diets containing more beet pulp, whole-tract digestibility of organic matter increased linearly, and intake of digestible organic matter was not affected. Partially replacing high-moisture corn with beet pulp in low-forage diets increased fiber digestibility without reducing whole-tract starch digestibility.

Animal Feed↗

Acute and acclimated digestive responses of the cockle Cerastoderma edule (L.) to changes in the food quality and quantity. II. Enzymatic, cellular and tissular responses of the digestive gland.

Cockles Cerastoderma edule were fed two different concentrations of two diets with different qualities which were achieved by mixing different proportions of ashed silt particles with cells of the microalgae Tetraselmis suecica. After 3 days (acute response) and 11 days (acclimated response) of exposure to the diets, we analysed the digestive activity of the digestive gland using cyto-histological and enzymatic techniques. We measured (i) the volumetric fraction of digestive and basophilic cells in digestive tubules, (ii) the diverticular radius and the thickness of digestive epithelia, (iii) the stereological parameters characterizing the lysosomal system and, (iv) dry weight, soluble protein content and specific and total amylase, cellulase, laminarinase, and protease activities of the digestive gland. In the conditions of the present study, specific cellulase and laminarinase activities in the digestive gland of cockles were correlated with the volumetric fraction of basophilic cells (r=0.672 and 0.642, respectively), whereas the specific protease was highly correlated (r=0.866) with lysosomal volume density. The implications of these correlations are discussed in relation to the feeding and absorptive parameters reported in the preceding publication. In the acute response, adjustments of the synthesis of constituents of the lysosomal/proteolytic system of the digestive cells seemed to be the only mechanism operating at the digestive level to respond to the changes in food availability. Lysosomal volume density increased with rising ingestion rate of organic matter, however, the occurrence of a limit in this short-term tissular response would account for the recorded trade-off between absorption efficiency and ingestion rate with different food qualities. With regard to acclimation, food quality determined the nature of the response of the digestive gland. With low quality diets, a time-dependent capability of the digestive gland for intensifying lysosomal/proteolytic production explains the increase of food absorption rates that result from higher filtration and ingestion rates. With high quality food, digestive acclimation differed with food particle concentration: with low rations, in spite of constant morphometrical and stereological parameters, the significant changes in the absorptive balance of biochemical components suggests the existence of an increased production of lysosomes that promotes an accelerated turn-over rate of the digestive epithelia. With high food concentrations, this response was coupled with increased activities of cellulase and laminarinase enzymes, probably as a consequence of higher rates of enzyme secretions from basophilic cells.

Journal Article↗

Digestive tract morphology and digestion in the wombats (Marsupialia: Vombatidae).

Wombats consume grasses and sedges which are often highly fibrous. The morphology of the digestive tract and the sequence of digestion were studied in two species of wombats from contrasting habitats: Vombatus ursinus from mesic habitats and Lasiorhinus latifrons from xeric regions. Studies were performed on wild wombats consuming their natural winter diets, and on captive wombats fed a high-fibre pelleted straw diet. Vombatus had a shorter digestive tract (9.2 vs 12.5 times body length) of greater capacity (wet contents 17.9 vs 13.7% body weight) than Lasiorhinus. The most capacious region of the digestive tract was the proximal colon (62-79% of contents). The proportional length and surface area of the proximal colon were greater in Vombatus, but those of the distal colon were greater in Lasiorhinus. These digestive morphologies may reflect adaptations for greater capacity and longer retention of digesta in Vombatus, but greater absorption and lower faecal water loss in Lasiorhinus. Apparent digestion along the digestive tract was estimated by reference to lignin. The proximal colon was the principal site of fibre and dry matter digestion, whereas nitrogen was mainly digested in the small intestine. Depot fats in captive wombats were highly unsaturated and reflected those in the diet. Therefore, lipids, proteins and soluble carbohydrates in the plant cell contents were digested and absorbed in the stomach and small intestine. Conversely, dietary fibre was probably retained and digested by microbial fermentation along the proximal colon.

Animal Nutritional Physiological Phenomena↗

Mesophilic and thermophilic temperature co-phase anaerobic digestion compared with single-stage mesophilic- and thermophilic digestion of sewage sludge.

The performance of thermophilic and mesophilic temperature co-phase anaerobic digestions for sewage sludge, using the exchange process of the digesting sludge between spatially separated mesophilic and thermophilic digesters, was examined, and compared to single-stage mesophilic and thermophilic anaerobic digestions. The reduction of volatile solids from the temperature co-phase anaerobic digestion system was dependent on the sludge exchange rate, but was 50.7-58.8%, which was much higher than 46.8% of single-stage thermophilic digestion, as well as 43.5% of the mesophilic digestion. The specific methane yield was 424-468 mL CH(4) per gram volatile solids removed, which was as good as that of single-stage mesophilic anaerobic digestion. The process stability and the effluent quality in terms of volatile fatty acids and soluble chemical oxygen demand of the temperature co-phase anaerobic digestion system were considerably better than those of the single-stage mesophilic anaerobic processes. The destruction of total coliform in the temperature co-phase system was 98.5-99.6%, which was similar to the single-stage thermophilic digestion. The higher performances on the volatile solid and pathogen reduction, and stable operation of the temperature co-phase anaerobic system might be attributable to the well-functioned thermophilic digester, sharing nutrients and intermediates for anaerobic microorganisms, and selection of higher substrate affinity anaerobic microorganisms in the co-phase system, as a result of the sludge exchange between the mesophilic and thermophilic digesters.

Bacteria, Anaerobic↗

Influence of two plant extracts on broilers performance, digestibility, and digestive organ size.

A 42-d trial was conducted to study the influence of 2 plant extracts on performance, digestibility, and digestive organ weights in broilers. The feeding program consisted of a starter diet until 21 d and a finisher diet until 42 d. There were 4 treatment groups: control; 10 ppm avilamycin (AB); 200 ppm essential oil extract (EOE) from oregano, cinnamon, and pepper; and 5,000 ppm Labiatae extract (LE) from sage, thyme, and rosemary. No differences in feed intake or feed conversion were observed. From 14 to 21 d of age, broilers fed the LE diet grew faster than the broilers fed the control or EOE feeds (68.8 vs. 63.9 and 61.6 g/d, respectively). Antibiotic and plant extract supplementation improved apparent whole-tract and ileal digestibility of the nutrients. For starter feed, LE supplementation improved apparent fecal digestibility of DM (P < 0.01), and all additives increased ether extract digestibility (P < 0.001). However, no effect was detected for CP digestibility (P > 0.1). At the ileal level, the AB, EOE, and LE supplementation of the starter feed increased DM and starch (P < 0.01) digestibility but not CP digestibility (P > 0.1). All additives improved apparent fecal digestibility of DM and CP of the finisher diet. No differences were observed for proventriculus, gizzard, liver, pancreas, or large or small intestine weight. In the present study, both plant extracts improved the digestibility of the feeds for broilers. The effect of different additives on digestibility improved the performance slightly, but this effect was not statistically significant.

Animals↗