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The Role of Dietary Starch and Non-Starch Polysaccharides in Swine Performance and Nutrient Receptors Gene Expression

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The Role of Dietary Starch and Non-Starch Polysaccharides in Swine Performance and Nutrient Receptors Gene Expression UNIVERSITÀ CATTOLICA DEL SACRO CUORE Sede di Piacenza Scuola di Dottorato per il Sistema Agro-alimentare Doctoral School on the Agro-Food System cycle XXVI S.S.D: AGR15; AGR18; BIO11 THE ROLE OF DIETARY STARCH AND NON-STARCH POLYSACCHARIDES IN SWINE PERFORMANCE AND NUTRIENT RECEPTORS GENE EXPRESSION Candidate: Marcin Rzepus Matr. n.: 3911385 Academic Year 2013/2014 Scuola di Dottorato per il Sistema Agro-alimentare Doctoral School on the Agro-Food System cycle XXVI S.S.D: AGR15; AGR18; BIO11 THE ROLE OF DIETARY STARCH AND NON-STARCH POLYSACCHARIDES IN SWINE PERFORMANCE AND NUTRIENT RECEPTORS GENE EXPRESSION Coordinator: Ch.mo Prof. Antonio Albanese ___________________________ Candidate: Marcin Rzepus Matr. n.: 3911385 Tutors: Prof. Francesco Masoero Dr Maurizio Moschini Dr Eugeni Roura Academic Year 2013/2014 γηράςκω δ' αἰεὶ πολλὰ διδαςκόμενοσ 4 Table of contents Words index 7 1. General introduction 9 2. Literature review 12 2.1. Carbohydrates 12 2.2. Gastrointestinal digestive and absorptive processes 26 2.3. Methods to evaluate digestion 38 2.4. Nutrient perception in gastrointestinal tract 48 2.5. References 66 3. Research projects – Part 1 - Dietary starch and non-starch polysaccharides in swine performance 90 3.1. Monocereal diets with hulled or hulless barley varieties. Performances and carcass quality of Italian heavy pigs 90 3.1.1. Appendix 1 - Diet based on corn can be replaced by naked barley variety in pig production - in vitro approach 107 3.2. Addition of nonstarch polysaccharides degrading enzymes to two hulless barley varieties fed in diets for weaned pigs 133 3.2.1. Appendix 2 - Diets for pigs based on hulles barley varieties supplemented with a non-starch polysaccharides degrading enzymes did not improve starch digestion potential in vitro 148 3.3. Inclusion of pasta by-products into diet improve starch degradation, fermentation kinetics and lower production of volatile fatty acids in an in vitro model of the porcine gastrointestinal tract 165 3.4. Kinetics of starch digestion and in vitro gas production affected by maize particle size 186 5 4. Research projects – Part 2 – Porcine taste receptors and nutritional homeostasis 211 4.1. Sugar, amino and fatty acids receptor gene expression in pigs fed iso- energetic diets enriched with β-glucans or arabinoxylans 211 4.2. Identification and expression profile of the porcine bitter taste receptor repertoire along the gastrointestinal tract 241 5. Conference papers 259 6. Conclusions 281 Acknowledgments 285 Marcin 6 Words index AA Amino Acid AACC American Association for Clinical Chemistry ADFI Average daily feed intake ADG Average daily gain; AUC Area under the curve AX Arabinoxylan BCFA Branched-chain fatty acid (sum of iso-butyric and iso-valeric acid) BG Beta-glucan BW Body weight C∞ Final in vitro digestion coefficient CCNFSDU The Codex Committee on Nutrition and Foods for Special cDNA Complementary DNA Co In vitro digestion coefficient at 0 min, CP Crude protein Ct In vitro digestion coefficient of starch at time t DF Dietary fibre DM Dry matter DNA Deoxyribonucleic acid EDTA Ethylenediaminetetraacetic acid EXP Exponential model GC Gas chromatography GI Glycaemic index HI Hydrolysis index k Process rate LAG discrete lag phase; 7 LCFA Long-chain fatty acid MCFA Medium-chain fatty acid MM Mentel model NS Not significant NSP Non-starch polysaccharide OM Organic matter pGI Predicted glycaemic index RNA Ribonucleic acid RS Resistant starch SCFA Short-chain fatty acid SEM Standard error of mean T/2 Half-time to asymptote TS Total starch Vf Final asymptotic gas volume; VFA Volatile-fatty acid 8 1. General introduction Plant carbohydrates are primary source of energy for both humans and livestock. The need to improve and optimize the efficiency of starch digestion is an important research focus in animal nutrition. In monogastric animals, such as swine, starch from cereals grains and theirs co-products is mainly digested in the small intestine. However, some fractions of carbohydrates known as resistant starch (RS) or fibrous non-starch polysaccharide can escape enzymatic digestion, reach the large intestine to be a source of fermentable material for microbiota. In vivo studies on digestibility and performance represent the natural methods with appropriate environment to obtain necessary data and information. However, they are time-consuming and expensive and requires a large number of animal subjects, sometimes with specific features. A good alternative for the in vivo trials are in situ and in vitro techniques used to estimate starch digestion potential of feed grains. In swine nutrition, in vitro methods, proposed in the last decade, are based on multi-enzymatic reactions mimicking the digestion processes occurring in the gastro-intestinal tract. Besides some limitations, in vitro procedures have been shown to accurately and precisely estimate the metabolic response of starch digestion in swine. Efforts have been put to study the complexities of starch digestion kinetics using several mathematical models. Cereals and their co-products included into diets are choose on the basis of their nutrient composition, usually without taking into account possible differences in palatability. Recent trials proved that feed preference in pigs is affected by cereal type, inclusion rate and diet form. Such approach is strategic, particularly during critical stages like weaning, and could help to improve feed intake. The past several years, since the discovery of taste receptors in the early 2000s, have been an stimulating time for the field of taste biology and the coming years will most likely remain still very rich in research activities. Mammals distinguish the five basic taste qualities, sweet, sour, umami, salty, and bitter, to assess the quality of consumed food. But recent findings challenge the canon of five basic sensations, strongly suggesting significant role of taste in dietary fatty acids perception. The sensory cells that 9 allow taste perception, the taste receptor cells, are distributed in the oral cavity, with the tongue being the prime taste organ. When the main role of tongue is to taste flavours of food before ingestion, a spread network of receptors are present also along the gastrointestinal system, within enteroendocrine cells population, to sense and transmit the information to regulatory systems, which adjust activities according to the luminal content. Consequently, the composition of the diet can affect the way in which gut hormones, secreted by entero-endocrine, could affect the gastrointestinal physiological mechanisms influencing nutrient uptake, animal growth and development. The present work has been divided into two parts. At first, research projects attempt to evaluate technologies that increase digestibility of energy and other nutrients in cereal grains and their co- products. Differences in starch digestibility have been attributed to various factors, including the the type of corn endosperm, the presence of proteins and prolamins, the presence of lipids, the amylose- amylopectin ratio, the starch granule structure, the particle size, the conservation and processing methods. Use of various processing techniques and exogenous enzymes could improve the nutritional value of feed. Studies were carried out to select barley verities, which could partially or totally replace corn in swine nutrition. Even if, barley contains anti-nutritional factors such as arabinoxylans and β- glucans (non-starch polysaccharides, NSP), supplementation of exogenous NSP enzymes capable of degrading polysaccharide cage structures around nutrients and reduce the viscosity of the intestinal contents improving the availability and digestibility of nutrients, and at the end the animal performance. Additionally, these trials are in the line with The European Environment Agency indications for sustainability, as Europe's water resources are seriously threatened. Barley belongs to cereals which in contrary to corn do not require long-term irrigation for efficient production. Therefore, agriculture sector could possibly achieve some benefits in terms of water usage performance. Second part of my work was dedicated to investigate the sensors of diet compounds which could be considered as potential markers to monitor nutrition status of organism. Activation of these receptors is believed to influence the hunger-satiety cycle, and their expression levels may be altered by dietary composition. Thus, the aim was to investigate for the first time the effect of arabinoxylans (AX) and 10 β-glucans (BG) on the relative level of expression of carbohydrates, amino and fatty acid nutrient sensors genes in porcine oral and non-oral tissues. Moreover a special chapter describe pattern of bitter receptors family (Tas2R) along gastrointestinal tract of pig. 11 2. Literature review 2.1. Carbohydrates Carbohydrates are the most important energy source in human nutrition, with the general formula (CH2O)n. They comprise a group of substances with different structures and varying physical, chemical, and physiological properties. The type and composition of dietary carbohydrates vary greatly among different food products. In this way they have wide range of influence on the food texture and flavor or variability and palatability of diet (Giuberti et al., 2014; Zijlstra and Jha, 2012). Population studies have shown that general carbohydrates supply in developed countries is approximately 45% of energy requirements and up to 85% in developing countries. They are considered as a fundamental source
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