N. le Floc'h

The effect of sanitary status degradation and dietary tryptophan content on growth rate and tryptophan metabolism in weaning pigs

Health degradation modifies Trp metabolism through induction of Trp catabolism. This could limit the amount of Trp available for growth. The aims of the present experiment were to investigate the effects of a low grade inflammation and dietary Trp on growth and Trp metabolism. Eighty weaned pigs were assigned to 4 experimental treatments according to a 2 x 2 factorial arrangement: 2 sanitary statuses x 2 dietary Trp contents. The Trp content was deficient (low-Trp: 2.4 and 1.9 g of Trp/kg of the phase I and phase II diets, respectively) or adequate (high-Trp: 2.9 and 2.4 g of Trp/kg of the phase I and phase II diets, respectively). A low grade inflammatory response was induced by housing pigs in unsanitary environment, whereas control pigs were housed in good sanitary conditions. Pigs were not fed ad libitum to avoid feed refusals. Growth performance was calculated 3, 5, and 7 wk after weaning. Blood was sampled 12, 33, and 47 d after weaning for the determination of plasma concentrations of Trp and related metabolites, kynurenine and pyridoxal-5-phosphate. The interaction between sanitary status and dietary Trp was not statistically significant in all measured criteria. Pigs kept in poor sanitary conditions grew slower (P < 0.001) during the entire experimental period and had greater plasma concentrations of haptoglobin (P < 0.001) than pigs housed in good sanitary conditions. Pigs housed in poor sanitary conditions had also decreased Trp plasma concentrations (P < 0.001), but plasma kynurenine concentrations were not affected. Our results indicated that a moderate inflammatory response was obtained by degrading the sanitary quality of environment. Additionally, poor sanitary conditions modified Trp metabolism, indicating that the amount of Trp available for growth and other metabolic functions might be reduced.

The standardized ileal digestible valine-to-lysine requirement ratio is at least seventy percent in postweaned piglets.

To reduce the impact of animal production on the environment, the CP content of the diet can be reduced by limiting the excess supply of AA. Improving the balance between AA relative to the requirement of the animal implies that we need to have accurate knowledge of the requirement of individual AA. The purpose of this study was to determine the Val requirement in postweaned piglets (12 to 25 kg) because Val is considered to be potentially limiting to performance after Lys, Met (and Cys), Thr, and Trp. The first experiment was carried out to identify a diet limiting in Lys supply. In this experiment, piglets were offered 1 of 3 diets: a low-CP diet containing low or adequate Lys concentrations [providing 1.0 and 1.2% standardized ileal digestible (SID) Lys, respectively] or a normal-CP diet with 1.2% SID Lys. Average daily gain of piglets receiving the diet containing 1.0% SID Lys was significantly less than that of piglets receiving diets containing 1.2% SID Lys at low or normal CP (486 vs. 522 g/d, respectively; P < 0.01). In Exp. 2, four diets with 1.0% SID Lys were used in a 2 x 2 factorial design, in which diets contained 57 or 70% SID Val:Lys in combination with 50 or 60% SID Ile:Lys. Independent of the Ile supply, feed intake and daily BW gain were, respectively, 15 and 20% less in piglets receiving diets providing 57% SID Val:Lys compared with piglets receiving 70% SID Val:Lys (P < 0.001). The Ile content of the diet did not affect feed intake or daily BW gain (P > 0.10). Experiment 3 was conducted to evaluate the response of piglets to an increasing Val supply provided by 2 sources of l-Val differing in the degree of purity. Increasing the Val supply from 58 to 66% SID Val:Lys resulted in a linear increase in both feed intake and daily gain by 24 and 30%, respectively (P < 0.001). No difference was observed between both sources of l-Val (P > 0.10). Experiment 4 was a dose-response study using 5 concentrations of Val supply (ranging from 60 to 80% SID Val:Lys). The estimated SID Val:Lys requirements for maximizing ADFI, ADG, and G:F were, respectively, 74, 70, and 68% using a linear-plateau model, and 81, 75, and 72% using a curvilinear-plateau model. Plasma Val, plasma alpha-ketoisovaleric acid, Ile, and Leu concentrations after an overnight fast increased with increasing Val supply (P < 0.001). The results of these experiments indicated that the SID Val:Lys was at least 70%, which was slightly greater than the current NRC recommendation.

Dietary tryptophan helps to preserve tryptophan homeostasis in pigs suffering from lung inflammation1

In pigs, inflammation modifies Trp metabolism and consequently could impact on Trp requirement for growth. In this study, the effects of lung inflammation, induced by the intravenous injection of complete Freunds adjuvant, and dietary Trp content on Trp metabolism and availability were investigated. Two dietary Trp contents, one corresponding to a low-Trp diet (1.5 g of Trp/kg of diet, Basal diet) and the second to an adequate-Trp diet (2 g of Trp/kg of diet, TRP diet), were used. Ten blocks of 4 littermate piglets were selected at 40 d of age. Within each block, piglets were randomly assigned to 1 of the 4 experimental treatments: (1) healthy control and Basal diet, (2) inflammation and Basal diet, (3) inflammation and Basal diet + antioxidant, and (4) inflammation and TRP diet. Inflammation induced an increase in indoleam-ine 2,3 dioxygenase (IDO) activity, an enzyme involved in Trp catabolism, in lung, lymph nodes, heart, and spleen (P < 0.01). Contrary to piglets fed the TRP diet, pigs suffering from inflammation did not maintain their plasma Trp concentrations when they were fed the Basal diet. Furthermore, pigs fed the TRP diet had decreased plasma haptoglobin concentrations, IDO activity, and lung weight than those fed the Basal diet, indicating that the inflammatory response was moderated with the greater Trp supply. Antioxidant addition in the Basal diet decreased the effects of inflammation on plasma Trp concentrations and IDO activity. These results indicated that inflammation increases Trp catabolism and thus may decrease Trp availability for growth.

Estimation of the tryptophan requirement in piglets by meta-analysis

There is no consensus concerning the Trp requirement for piglets expressed relative to Lys on a standardized ileal digestible basis (SID Trp : Lys). A meta-analysis was performed to estimate the SID Trp : Lys ratio that maximizes performance of weaned piglets between 7 and 25 kg of BW. A database comprising 130 experiments on the Trp requirement in piglets was established. The nutritional values of the diets were calculated from the composition of feed ingredients. Among all experiments, 37 experiments were selected to be used in the meta-analysis because they were designed to express the Trp requirement relative to Lys (e.g. Lys was the second-limiting amino acid in the diet) while testing at least three levels of Trp. The linear-plateau (LP), curvilinear-plateau (CLP) and asymptotic (ASY) models were tested to estimate the SID Trp : Lys requirement using average daily gain (ADG), average daily feed intake (ADFI) and gain-to-feed ratio (G : F) as response criteria. A multiplicative trial effect was included in the models on the plateau value, assuming that the experimental conditions affected only this parameter and not the requirement or the shape of the response to Trp. Model choice appeared to have an important impact on the estimated requirement. Using ADG and ADFI as response criteria, the SID Trp : Lys requirement was estimated at 17% with the LP model, at 22% with the CLP model and at 26% with the ASY model. Requirement estimates were slightly lower when G : F was used as response criterion. The Trp requirement was not affected by the composition of the diet (corn v. a mixture of cereals). The CLP model appeared to be the best-adapted model to describe the response curve of a population. This model predicted that increasing the SID Trp : Lys ratio from 17% to 22% resulted in an increase in ADG by 8%.

Response of piglets to the valine content in diet in combination with the supply of other branched-chain amino acids

The branched-chain amino acids (BCAA) valine (Val) and isoleucine (Ile) are considered to be among the next-limiting amino acids for growth in piglets. In earlier studies, we estimated the standardized ileal digestible (SID) Val : Lys (lysine) requirement to be at least 70%, whereas the Ile : Lys requirement may be as low as 50%. Because the BCAA partially share a common route of catabolism, the supply of one BCAA may affect the availability of the other BCAA. Four experiments were conducted to determine the response of 6-week-old piglets to the Val supply in relation to the other BCAA. A deficient supply of Val or Ile typically results in a reduction in average daily feed intake (ADFI). Experiment 1 was designed to determine the effect of a limiting Val supply, independent of the effect on feed intake. In a dose-response study using restrictively fed piglets, nitrogen retention did not increase for an SID Val : Lys supply greater than 64%. In the remaining experiments, piglets were offered feed ad libitum using ADFI, average daily gain (ADG) and gain-to-feed ratio as response criteria. The interaction between the Val and leucine (Leu) was studied in Experiment 2 in a 2 × 2 factorial design (60% and 70% SID Val : Lys, and 111% and 165% SID Leu : Lys). Performance was considerably lower in piglets receiving 60% Val : Lys compared with those receiving 70% Val : Lys and was lowest in piglets receiving the diet with low Val and high Leu content. To further evaluate the interaction between Val and Leu, a dose-response study was carried out in which the response to Val supply was studied in combination with high Leu supply (165% Leu : Lys). Using a curvilinear-plateau model, the average SID Val : Lys requirement was 72%. However, low Val supply (60% SID Val : Lys) reduced performance by 13% to 38%, which was much greater than what we observed in earlier studies. Experiment 4 was carried out to test the hypothesis that the Val requirement is not affected by low Ile supply (50% SID Ile : Lys). Performance was not improved for Val : Lys supplies greater than 65%, which may indicate that Ile (and not Lys) was second-limiting in this study. In conclusion, the first response of piglets to deficient Val supply appears to be a reduction in ADFI, rather than a reduction in ADG or nitrogen retention. A large supply of Leu may not affect the Val requirement per se, but may aggravate the consequences of Val deficiency.

The standardized ileal digestible isoleucine-to-lysine requirement ratio may be less than fifty percent in eleven- to twenty-three-kilogram piglets.

Most studies concerning the Ile requirement in pigs have been carried out using blood products as a protein source, and these have a relatively low Ile content relative to the other branched-chain AA (BCAA). There are indications that an excess supply of one BCAA can affect the utilization of the other BCAA. Little information is available concerning the Ile requirement in pigs when the supply of the other BCAA is moderate (e.g., in cereal- and soybean meal-based diets). The objective of the present study was to evaluate the response of piglets to Ile supplementation under different nutritional conditions. In all experiments, piglets were housed individually and had ad libitum access to feed during a 3-wk period. The first experiment was carried out to study the response of piglets to an increasing Ile supply by using 2 sources of l-Ile differing in degree of purity. Piglets received either a control diet with 48% standardized ileal digestible (SID) Ile:Lys or 1 of 4 other diets containing graded levels of either source of l-Ile to provide 52 or 56% SID Ile:Lys. All diets were formulated to provide 1.00% SID Lys in the diet. Feed intake and growth were not affected by Ile level or Ile source. Experiment 2 was performed to exclude a possible interaction between Ile and Lys supply. In a 2 x 2 factorial arrangement, 2 levels of Lys (1.00 and 1.15% SID Lys) and 2 levels of Ile (48 and 60% SID Ile:Lys) were used. Growth and G:F were 8 and 7% greater in piglets receiving the diet with the greater Lys content, but the Ile:Lys did not affect performance. No interactions were observed between the Lys and Ile supplies. In Exp. 3, a 2 x 2 factorial arrangement was used to test the effect of protein source (spray-dried blood cells or corn gluten meal) and Ile supply (50 or 65% SID Ile:Lys) on performance in piglets. Both protein sources had an elevated BCAA content but differed in Leu and Val contents. Protein source or Ile supply did not affect feed intake, growth, or G:F in the piglets. Plasma concentrations after an overnight fast reflected the difference in AA concentrations of the diets. In conclusion, the results of these experiments indicate that the SID Ile:Lys requirement may be not greater than 50% in piglets receiving cereal- and soybean meal-based diets with a moderate BCAA content. In contrast to other studies, we could not confirm that the Ile requirement was affected by BCAA content of the diet.

In vivo threonine oxidation in growing pigs fed on diets with graded levels of threonine

Threonine oxidation to glycine was investigated in vivo in twelve growing pigs (27.4 kg live weight) fed on one of the following three diets with graded levels of threonine supply: a low-threonine diet (LT), a control well-balanced diet (C) or a high-threonine diet (HT), during 10 h constant infusion of L-[1-13C]threonine and [2-3H]glycine in the cranial vena cava and [1-14C]glycine in the portal vein. 13C-threonine and glycine enrichments and [3H]glycine and [14C]glycine specific radioactivities (SR) were determined at plateau in peripheral venous plasma, liver and pancreas. Glycine production rates calculated from plasma [2-3H]glycine or [1-14C]glycine SR gave similar values suggesting that [1-14C]glycine SR could be used in order to estimate whole-body glycine flux. The high pancreas [1-13C]glycine enrichment provided evidence that the pancreas may be, with the liver, a major site of threonine oxidation to glycine. Moreover, the present findings suggest that threonine transport into the liver could be the limiting step of threonine oxidation in this tissue when dietary threonine supply is low. Total threonine oxidation to glycine, calculated from plasma values of enrichment and specific radioactivity, was low and constant when the estimated absorbed threonine was lower than 4 g/d and increased for higher amounts of absorbed threonine.

Review: divergent selection for residual feed intake in the growing pig.

This review summarizes the results from the INRA (Institut National de la Recherche Agronomique) divergent selection experiment on residual feed intake (RFI) in growing Large White pigs during nine generations of selection. It discusses the remaining challenges and perspectives for the improvement of feed efficiency in growing pigs. The impacts on growing pigs raised under standard conditions and in alternative situations such as heat stress, inflammatory challenges or lactation have been studied. After nine generations of selection, the divergent selection for RFI led to highly significant (P<0.001) line differences for RFI (−165 g/day in the low RFI (LRFI) line compared with high RFI line) and daily feed intake (−270 g/day). Low responses were observed on growth rate (−12.8 g/day, P<0.05) and body composition (+0.9 mm backfat thickness, P=0.57; −2.64% lean meat content, P<0.001) with a marked response on feed conversion ratio (−0.32 kg feed/kg gain, P<0.001). Reduced ultimate pH and increased lightness of the meat (P<0.001) were observed in LRFI pigs with minor impact on the sensory quality of the meat. These changes in meat quality were associated with changes of the muscular energy metabolism. Reduced maintenance energy requirements (−10% after five generations of selection) and activity (−21% of time standing after six generations of selection) of LRFI pigs greatly contributed to the gain in energy efficiency. However, the impact of selection for RFI on the protein metabolism of the pig remains unclear. Digestibility of energy and nutrients was not affected by selection, neither for pigs fed conventional diets nor for pigs fed high-fibre diets. A significant improvement of digestive efficiency could likely be achieved by selecting pigs on fibre diets. No convincing genetic or blood biomarker has been identified for explaining the differences in RFI, suggesting that pigs have various ways to achieve an efficient use of feed. No deleterious impact of the selection on the sow reproduction performance was observed. The resource allocation theory states that low RFI may reduce the ability to cope with stressors, via the reduction of a buffer compartment dedicated to responses to stress. None of the experiments focussed on the response of pigs to stress or challenges could confirm this theory. Understanding the relationships between RFI and responses to stress and energy demanding processes, as such immunity and lactation, remains a major challenge for a better understanding of the underlying biological mechanisms of the trait and to reconcile the experimental results with the resource allocation theory.

Meta-analysis of the response of growing pigs to the isoleucine concentration in the diet

The efficiency of nitrogen utilization will be highest when the amino acid (AA) supply approaches the requirement of the animal. With the availability of different crystalline AA, it is theoretically possible to formulate low-protein diets for growing pigs in which seven AA are co-limiting for performance. In such a diet, the concentration of Lys, Met, Met + Cys, Thr, Trp and Val and a seventh AA would exactly match the requirement. To determine the extent to which low-protein diets can be used, it is important to have reliable information about the requirements for these AA. Isoleucine is often considered the seventh-limiting AA in diets for growing pigs; however, information about the Ile requirement is limited and sometimes conflicting. The purpose of this study was to carry out a meta-analysis of the available literature information to determine the Ile requirement in growing pigs. A total of 46 Ile dose-response experiments were identified that used at least four concentrations of Ile in the diet. Because of differences in experimental design, both the Ile concentration and the response criteria were standardized. In 13 dose-response experiments, there was no indication of a response to an increasing Ile concentration. For the other 33 experiments, a response to the increasing Ile concentration was observed and the Ile requirement estimates ranged from 53% to 114% of that of the National Research Council (1998). An Ile concentration below the requirement resulted in important reductions in both feed intake and growth. A 10% reduction in the Ile concentration (below the requirement) resulted in a 15% reduction in feed intake and a 21% reduction in daily gain. The use of blood products in the diet was the main factor determining whether a response to the Ile concentration was observed or not. Blood meal and blood cells are protein sources with a very low Ile concentration, but with high or very high concentrations of Leu, Val, Phe and His. Some of these AA compete with Ile for catabolic pathways or transport across the blood-brain barrier, thereby potentially increasing the requirement for Ile. In diets without blood products, the Ile requirement appears to be lower than the currently recommended requirement. On the basis of the outcome of this study, we recommend a Ile : Lys requirement ratio of at least 50% on a standardized ileal digestible basis.

Reduced feed intake of lactating primiparous sows is associated with increased insulin resistance during the peripartum period and is not modified through supplementation with dietary tryptophan.

The aim of this experiment was to investigate whether insulin resistance is related to the dietary concentration of Trp and the ADFI of primiparous sows having similar body conditions. Twenty-four primiparous sows were catheterized on d 97 of pregnancy. Blood samples were collected during 3 tests: after the ingestion of 1.5 kg of feed (meal test), after the intravenous infusion of 0.5 g of glucose/kg of BW (glucose tolerance test), and during an euglycemic hyperinsulinemic clamp with an infusion rate of 100 ng of insulin x kg of BW(-1) x min(-1). Both tests were performed at 4 stages at approximately d 103 and 110 of pregnancy and at d 3 and 10 of lactation. Sows were fed a diet containing 0.16 or 0.26% of total Trp (suboptimal vs. slight excessive Trp supply according to recommendations for lactating sows) from d 104 of pregnancy after the first clamp until weaning. The dietary treatment did not result in differences in ADFI, BW, and backfat changes, and growth of piglets during lactation. Plasma Trp concentration was greater for the sows allocated to the slight excessive Trp diet than for the sows allocated to the suboptimal Trp diet (P < 0.05). Plasma glucose, NEFA, and urea profiles during the meal tests were not affected by the dietary treatment. At d 3 of lactation, the insulin concentration at 105 (P = 0.03) and 120 min (P = 0.04) after meal intake was less for the sows allocated to the slight excessive Trp diet than for the sows allocated to the suboptimal Trp diet. On d 10 of lactation, the glucose half life (P = 0.03) and the time needed to reach 25% of the area under the insulin curve (P = 0.04) during the tolerance test were less for the sows allocated to the slight excessive Trp diet than for the sows allocated to the suboptimal Trp diet. The glucose infusion rate during euglycemic hyperinsulinemic clamps was similar in the 2 Trp groups of sows. Irrespective of the dietary treatment, the ADFI of the sows was negatively related to the glucose half life during the glucose tolerance test and positively related to the glucose infusion rate during the clamp (P < 0.05). This relationship observed with the tests performed during early lactation was already found with the tests performed during late pregnancy (P < 0.02). Present findings indicate that a dietary Trp supply of 0.26% does not increase feed intake in lactating primiparous sows. This result indicates that the interest in a Trp supplementation during the peripartum period can be questioned. Irrespective of the dietary treatment, the reasons why sows with similar rearing conditions develop different rates of insulin resistance during pregnancy remain to be elucidated.