Module 3Grain Feeding14. Digestion of GrainJohn NolanLearning objectivesOn completion of this topic you should be able to: Discuss the animal factors influencing starch digestion in ruminants.Discuss the dietary factors influencing starch digestion in ruminants.Describe the biochemical processes of starch digestion and the importance of volatile fattyacid production to this process.Key terms and conceptsStarch, fermentation acetate, propionate, butyrate, volatile fatty acid, site of digestionIntroduction to the topicRuminant animals have evolved with the unique capacity to digest cellulose. The digestive tract ofruminants has developed in a way that allows them to survive and produce on roughage-baseddiets. In recent years, there has been an increasing incentive to feed concentrate diets based oncereal grains. The feeding of grain-based diets can improve animal production and produce a meatproduct that is more desirable to the consumer.As ruminants have evolved to consume roughage-based diets, the provision of grain-based dietscan cause digestive problems that may lead to morbidity and mortality. Therefore, carefulmanagement of grain-based feeding to ruminants is required. With careful management, significantimprovements in the quality and quantity of production (milk, wool, meat and reproduction) can beachieved.This module will provide you with an understanding of the metabolic implications associated withintroducing grain-based diets to ruminant production systems and therefore an understanding ofthe management strategies available for commercial production.14.1 Principles of starch digestionIn all animals, digestion occurs via a combination of microbial and enzymic digestion. Microbialdigestion relies on enzymes produced by microbes whereas the host’s digestion system relies onendogenous enzymes secreted into the digestive tract. Microbes possess a far wider range ofenzymes than the animal’s digestive system and are able to break down and utilise most feedcomponents.Digestion of organic materials in feeds in the absence of oxygen is referred to as fermentation.The fermentationprovidesgutmicrobes with the energy they need to survive and to grow. The end–products of the fermentationare principally the volatile fatty acids (VFA) as well as gases such as hydrogen, methane andcarbon dioxide. VFA are rapidly absorbed from the gut and provide an important source of energyfor theApplied Animal Nutrition ANUT300/500 - 14 - 1 2009 The Australian Wool Education Trust licensee for educational activities University of New England
animal. As the microbes grow, microbial protein is synthesised and, in situations wherefermentation occurs prior to gastric digestion, microbes provide a valuable source of amino acidsfor the host animal. When microbial biomass is produced by hindgut fermentation, the animal is notable to digest the protein or absorb amino acids and they pass from the animal in the faeces.Enzymic digestion occurs in the gastric stomach and in the small intestine. Protein breakdownoccurs in two main steps. The acid conditions in the stomach denature the protein and facilitate theactivity of pepsin that mainly produces large peptide fragments and some free amino acids. Thepeptide fractions and amino acids are important in stimulating cholecystokinin (CCK) release in theduodenum which then plays a major role in gastric digestion by stimulating pancreatic enzymeproduction and intestinal enteropeptidase secretion. The supply of pancreatic enzymes into thesmall intestine is very important in providing both trypsinogen and α–amylase. Trypsinogen entersthe intestine in pancreatic juice and is converted to trypsin by enteropeptidase produced by theintestinal mucosa. In addition to trypsin, chymotrypsin and elastase are also active in the smallintestine and break down a wide range of peptides to amino acids.The digestion of starch can be considered in three stages. Starch is converted by pancreatic α–amylase to a mixture of maltose, maltotriose, dextrin and glucose. The di– and poly–saccharidesare then converted to glucose by glucosidases which are surface enzymes of the small intestinalepithelial cells. The monosaccharides (glucose, galactose and fructose) that result from thedigestion of di– and polysaccharides are then absorbed either via the sodium– dependentmonosaccharide co–transporter pathway or via sodium–independent facilitated diffusion.14.2 Digestion of carbohydrates in different species ofanimalsThere are some major differences between animal species in the efficiency of intestinalcarbohydrate digestion and these are summarised in Table 14–1.Maize is poorly digested by the horse—even when finely ground.Maize is one of the grains with the highest apparent digestibility in poultry but is very poorlydigested in the small intestine of the horse even when it is finely ground. Similarly, sorghum grain iswell digested by poultry but is poorly digested in either the rumen or intestines of cattle when dry–rolled or ground. The differences in digestive efficiency between animal species are almostcertainly related to differences in animal enzyme systems, and/or absorption capacity of the smallintestine. An understanding of these differences may create exciting opportunities for newtechniques for preparing and feeding cereal grains.In ruminants, the digestion of starch in the small intestine may be limited by the availability ofamylase (Ørskov 1986). Thus, oligosaccharidase activity and monosaccharide transport across theenteroctye are not thought to be the limiting factors. To date the nutritional manipulation of amylasesecretion is not readily understood although it appears that protein/peptides entering the smallintestine can stimulate amylase production and increase glucose absorption (Taniguchi et al.1995).14 - 2 – Applied Animal Nutrition ANUT300/500 2009 The Australian Wool Education Trust licensee for educational activities University of New England
Table 14.1 Differences between livestock species in their ability to digest cereal grains.Source: Rowe (2006).14.3 Starch digestion and fermentation in sheep andcattleSheep are often used in nutritional studies of ruminant digestion and the data are then extrapolatedto enable interpretation of results of feeding trials in cattle. With respect to whole tract digestibilityof starch, the relationship between the proportion fermented in the rumen and that digested postruminally, differs in measurements made in sheep and cattle. The reason for the different digestiveprocesses is not entirely clear but it is likely to be related to the different sizes of sheep and cattleintestinal tracts and the dynamics of particle flow through the tract. The findings have highlightedthe risk of extrapolating the data from sheep to cattle, particularly when cracked or rolled grain isused.The patterns and efficiency of starch digestion are different in sheep and cattle.The benefits and disadvantages of fermentative and enzymic digestion in different parts of the tractare summarised in Table 14–2. From the animal’s point of view, it is beneficial, in most situations,to maximise the digestion of starch and absorption of glucose from the small intestine. This isbased on the energetic efficiency of intestinal digestion being approximately 30% higher thanfermentative digestion. The digestion of starch in the intestine carries no risk of acidosis and cansupply glucose as an important nutrient for marbling in beef production (Pethick et al. 1997).The benefits of microbial protein production associated with fermentation are not likely to be asimportant in grain feeding systems for high levels of production in ruminants as are the potentialrisks associated with acidosis and reduced fibre digestibility resulting from an accumulation of acidduring fermentation. In hindgut fermenters, such as the pig and horse, no microbial protein fromcaecal and colonic fermentation is available to the animal.On the other hand, there are considerable risks, such as laminitis, associated with fermentativeacidosis from high levels of starch reaching the hindgut (Godfrey et al. 1993; Rowe et al. 1995).There appear to be no benefits to any species associated with incomplete and inefficient digestionof starch from the small intestine. The role and manipulation of site of starch digestion is discussedin more detail elsewhere.Applied Animal Nutrition ANUT300/500 - 14 - 3 2009 The Australian Wool Education Trust licensee for educational activities University of New England
Table 14.2 Significance of site of digestion in determining nutritional value of grain.Source: Rowe (2006).The differences between animals in their digestive capacity with different grains highlight theimportance of enzymic digestion. The most marked differences highlighted in Table 14–1 are thosebetween the traditional grain eaters, poultry and pigs and the traditional roughage eaters, cattle andhorses, particularly with respect to digestion of grains such as sorghum and maize. It is not clearexactly what enzyme systems within the animal are responsible for these major differences.Readings!The following readings are available on CD:1. Cheng, K.J., McAllister, T.A., Popp, J.D., Hristov, A.N., Mir, A. and Shin, H.T. 1998, ‘Areview of bloat in feedlot cattle, Journal of Animal Science, vol. 76 pp. 299-308.2. Ghorbani, G.R., Morgari, D.P., Beauchemin, K.A. and Leedle, J.A.Z. 2002, ‘Effects ofbacterial direct-fed microbials on rumen fermentation, blood variables and the microbialpopulations of feedlot cattle’, Journal of Animal Science, vol. 80 pp. 1977-1986.3. Huntington, G.B. 1997, ‘Starch utilization by ruminants: from basics to the bunk’, Journal ofAnimal Science, vol. 75 pp. 852-867.ActivitiesAvailable on WebCTMulti-ChoiceQuestionsSelf AssessmentQuestionsSubmit answers via WebCT1.Rank grains in terms of their efficiency ofintestinal starch digestion.2.Why do you think that poultry are able to digeststarch more efficiently than ruminant animals?3.Do measurements of grain digestibility in sheepgive a good indication of the likely digestibility in cattle?4.What are the benefits of enzymatic starchdigestion compared to fermentative digestion?5.Comment on the value of ground maize as afeed for horses.Useful Web LinksAvailable on WebCTAssignment QuestionsChoose ONE question from ONE of the topics asyour assignment. Short answer questions appear onWebCT. Submit your answer via WebCT14 - 4 – Applied Animal Nutrition ANUT300/500 2009 The Australian Wool Education Trust licensee for educational activities University of New England
ReferencesBeever, D.E., Coelha da Silva, J.F. and Armstrong, D.G. 1970, ‘The effect of processing maize andits digestion in sheep’. Proceedings of the Nutrition Society Vol 29, pp 43A–44A.Choct, M. and Annison, G. 1990. ‘Anti–nutritive activity of wheat pentosans in broiler diets’. BritishPoultry Science Vol 31, pp 811–822.Godfrey, S. I.; Boyce, M. D.; Rowe, J. B.; Speijers, E. J. 1993. ‘Changes within the digestive tract ofsheep following engorgement with barley’. Australian Journal of Agricultural Research Vol 44, pp1093–1101.Householder, D.D., Potter, G.D. and Lichtenwalner, R.E. 1977. ‘Nutrient utilization in differentsegments of the equine digestive tract’. Proceedings of the 5th Equine Nutrition and PhysiologySymposium. p. 44–45. (Missouri, USA).Huntington, G.B. 1997. ‘Starch utilisation in ruminants, from basics to bunk’. Journal of AnimalScience Vol 75, pp 852–867.Meyer, H., Radicke, S., Kienzle, E., Wilke, S., Kleffken, D. and Illenseer, M. 1995. ‘Investigation onpreileal digestion of starch from grain, potato and manioc in horses’. Journal of VeterinaryMedicine Vol 42, pp 372–381.Mollah, Y., Bryden, W. L., Wallis, I. R., Balnave, D. and Annison, E. F. 1983. ‘Studies on lowmetabolisable energy wheats for poultry using conventional and rapid assay procedures and theeffects of processing’. British Poultry Science Vol 24, pp 81–89.Ørskov, E.R, Fraser, C. and Kay, R.N.B. 1969. ‘Dietary factors influencing the digestion of starch inthe rumen and small intestine of early weaned lambs’. British Journal of Nutrition Vol 23, pp 217–226.Ørskov, E.R. 1986. ‘Starch digestion and utilisation in ruminants’. Journal of Animal Science Vol63, pp 1624–1633.Owsley, W.F., Knabe, D.A. and Tanksley, T.D. 1981. ‘Effect of sorghum particle size on digestibilityof nutrients at the terminal ileum and over the total digestive tract of growing–finishing pigs’.Journal of Animal Science Vol 52, pp557–566.Pethick, D.W., McIntyre, B.L., Tudor, G., and Rowe, J.B. 1997. ‘The partitioning of fat inruminants—can nutrition be used as a tool to regulate marbling’. In ‘Recent Advances in AnimalNutrition in Australia.’ (Eds. J.L. Corbett, M. Choct, J.V. Nolan and J.B. Rowe). pp.151–158.(University of New England, Armidale).Reisenfeld, G., Sklan, D., Bar, A., Eisner, U. and Hurwitz, S. 1980. ‘Glucose absorption and starchdigestion in the intestine of the chicken’. Journal of Nutrition 110, pp117–121.Rowe, J.B., Pethick, D.W. and Johnson, K.G. 1995, 'Control of lactic acid and pH during invitrofermentation, of hind gut digesta in the presence of high levels of glucose’. ‘Controlling acidosisin the equine hindgut’. Recent Advances in Animal Nutrition Vol 10, pp 136-143.Taniguchi, K., Huntington, G.B. and Glenn, B.P. 1995. ‘Net nutrient flux by visceral tissues of beefsteers given abomasal and ruminal infusions of casein and starch’. Journal of Animal Science Vol73, pp 236–249.Applied Animal Nutrition ANUT300/500 - 14 - 5 2009 The Australian Wool Education Trust licensee for educational activities University of New England
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