Chapter Carbohydrates 7 Function: • Source of energy for life process. • In Plant: 1.Simple sugars (glucose,ribose) involved in energy transformation and tissue synthesis. 2.Starch as energy reserve. 3.Insoluble fraction (cellulose, hemicellulose) provided structural support. • Glycan: vary in shape and size, rangeing from linear chains (polysaccharides)to highly branched molecular • Soluble components of the plant cell:starch, hemicellulose and betaglucans, which are available to fermentation. Cellulose: glucose(1-4)-β-glucoside linkage. • Microflora of the rumen of ruminants and cecum and colon of nonruminants produce cellulase to hydrolyze cellulose. • Other nonrumianats utilize cellulose by anaerobic fermentation in large intestine. • Starch is soluble carbohydrate in plant • Two important molecular of glucose: Linked via a glucose-(!-4 )-α-glucoside linkage. • Amylose : glucose(1-4)-α-glucoside linkage linear polymer of glucose, MW 15000 ~ 100000 D. Amylopectin: linear polymer with branches 10 - 100 million D. • Maize contains 24% amylose and 76% amylopectin, Waxy maize contains 1%:99% • Rice and potatoes contain 20%:80% • The hydrolysis of starch in amylose and amylopectin to glucose is similar. Carbohydrate Digestion (一) preparation for absorption • Only monosaccharides can be adsorbed from GI tract, except in newborn animals. • Poly-, tri-, disaccharides hydrolyzed (by enzymes from host or microbe) to monosaccharide Plant cell wall digestion • cellulose, hemicellulose, pectin, lignin為植 物細胞壁之主要成分, 即structure polysaccharides, insoluble polysaccharide 之生物可利用率( bioavailability ), 即影響其 營養價值( nutritive value ). • Anaerobic fermentation of carbohydrate results in the production of large quantities of volatile fatty acids (VFA, = acetic acid, propionic acid and butyric acid )and provides a large proportion of the total energy supply. Absorption and transport of monosaccharides 1.Absorption: • proximal section of the small intestine (duodenum and jejunum) has greatest capacity to absorb monosaccharides. • glucose and galactose are absorbed very efficiency. Mannose absorbed about 20% of the efficiency of glucose, arabinose at the lowest rate. (Table 7.3 ) 2.Mechanism • Sodium dependent active transport mechanism transported against its concentration gradient which is maintained by membrane bound sodium-potassium ATPase (Na- K-ATPase) • Facilitative glucose transporters transport glucose actively down a concentration gradient. • These two families of glucose transporters are protein (500-600 amino-acids). 3. Factors that affect the absorption of glucose – Reduced by short-term (24 or 48h) fasting, increased by restricted feed intake. – diabetic absorb more rapidly than normal – adrenalectomy reduce in glucose absorption but no effect on xylose absorption. – thyroidectomy and ovariectomy reduce glucose absorption. 4.Digestive enzyme deficiency and utilization of CHO • Young mammals fed large amounts of sucrose develop diarrhea and death results from an insufficiency of sucrase. • Ruminant species produce no sucrase. • Some adults have lactase deficiency, like pig and human. • Xylose feeding of young pigs results in depressed appetite and growth and causes eye cataracts. The absorption of soluble CHO exceeds 90% Factors that affect the rate of starch digestion : • particle size. • nature of starch (amylose, amylopectin content) • interaction of starch with protein, fat. • presence of antinutrient such as phytate, tannin, saponins and enzyme inhibitors. (三) Metabolic conversions • Homeostasis : maintaining the blood glucose concentration glycogenesis blood glucose glycogenolysis insuline↑ glycogen glucagon↑ Gluconeogenesis • glucose can be formed by body tissues from noncarbohydrate metabolites, including lipids, amino acids. • Glucogenic amino acids such as nonessential amino acids and several of essential (arginine, methionine, cystine, histidine, threonine, tryptophane, valine.) • The amino acids used for gluconeogensis or for energy enter TCA cycle (Fig. 7.2). • Glucose as an energy source • Glucose is metabolized to energy has two stages. (1) glycolysis : occure under anaerobic condition and production of pyruvate. Referred to as the Embden-meyerhof pathway ( in the cytosol). (2)oxidation of pyruvate: pyruvate produced by glycolysis is transported into the mitosol and is oxidized to CO2 and H2O with production of energy • Energetics of glucose catabolism – 1 mole of glucose to 2 mole of pyruvate (produce 8 mole ATP(10-2)) – 2 mole of pyruvate to 2 mole of acetyl-CoA (produce 6 mole) – 2 mole acetyl-CoA to CO2 and H2O (produce 24 mole) Energetics of glucose catabolism 異化1mole glucose 產生38mole ATP glycolytic pathway 8 mole of ATP pyruvate to acetyl CoA 6 mole of ATP acetate to CO2 and H2O 24 mole of ATP Totle 38 mole of ATP • One mole of ATP has a value of 8Kcal/mole • 1 mole glucose 氧化產生之能量為 8Kcal x 38 = 304Kcal (45% of efficiency). • C6H12O6 → 6CO2 + 6H2O + 673 Kcal • The conversion of the free energy of the oxidation of glucose has an efficiency of 40-65%. Acetic acid as an energy source • Acetic acid is the major product of carbohydrate digestion in the rumen and only present in the peripheral blood. It is used as a source of energy by various tissues Propionic acid as an energy • In ruminant propionate are produced from carbohydrate breakdown in the rumen. → • Pass across the rumen wall to the liver→ • Change into glucose. : The first stage is conversion to succinyl-COA.(fig.9.7) Butyric acid as an energy source • Butyric acid is converted toβhydroybutyrate in its passage across the rumen walls and used as a source of energy by number of tissues Abnormal carbohydrate metabolism (一) Ketosis: • 因碳水化合物及脂肪代謝異常, 使過量之酮 體(acetone, acetoacetate, Bhydroxybutyrate)累積於體組織。 • 血中濃度增加稱Ketonemia(acetomia), 出現 在尿中則稱Ketoneuria. • 特別容易出現於反芻動物開始泌乳(牛)或懷 孕期(羊),對葡萄糖需要量大時,Vit B12缺 乏時。 hypoglycemia, depleted liver glycogen ↓ elevated mobilization of adipose tissue lipids ↓ increased production of ketones and Lipemia ↓ Increase breakdown of tissue protein for energy ↓ • body weight loss (breakdown of tissue protein) milk production decrease abortion in pregnant animals water consumption increase (loss body fluid ) loss of electrolytes (Na, K) • Treatment : injection of intravenous glucose, adrenal corticoid hormones, or adenocorticotropic hormone(ACTH) Diabetes mellitus • hyperglycemia ( excessive blood glucose ) • three types : – insulin–dependent diabetes (typeⅠ) – Non-insulin–dependent diabetes (typeⅡ) – gestational diabetes insulin • • • • promote glucose utilization increase protein synthesis decrease protein catabolism increase the entry of amino acids into the cell • stimulates formation of triglycerides (TG) • inhibit TG breakdown Diabetes mellitus • TypeⅠ: – destruction of the beta cell of Islets of Langerhans in the pancrease (缺insulin) – appear any age (genetic) – require insulin administration Diabetes mellitus • Type Ⅱ: – defect in utilization of insulin by liver and other tissue (insulin作用被破壞), defective cellular uptake of glucose and impaired release of glucose from liver, 造成高血壓。 Diabetes mellitus • Type 3: – increased tissue resistance to the action of insulin and increase the blood glucose. Insulin deficiency • carbohydrate, lipid, amino acid metabolism abnormal ↓ » negative N balance » glucose loss in urine » polyuria » electrolyte loss Insulin – like growth factors (IGF) • IGF I and IGFⅡ are small polypeptides that control the growth of several types of cells, have a structure very similar to that of insulin. • IGFⅠand bone growth 有關 IGFⅡand fetal growth 有關 • IGFⅠand IGFⅡand their binding proteins are major links between nutrient intake and anabolic response of cells. Plant fiber as energy source • Plant fiber :cellulose, hemicellulose, liglin, gum, and β-glucan are present as cellular membrane and storage form of energy in the plant. • Tannin present in the seed coat of dark seed (sorghum), it is unpalatable to animal and birds. • Van Soest partition plant tissues into two fractions. – Cell wall constituents • Lignin, cellulose, hemicellulose • Gum: pectin, glucan • -Cell contents • Simple sugars, starches Summary • • • • • • • • Monosaccharide Disaccharide Oligosaccharide : 3-10 sugar unit Polysaccharide : >10 sugar unit Crude fiber Dietary fiber Neutral detergent fiber Acid detergent fiber