Digestive Physiology of Primates
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Digestive physiology of primates Marcus Clauss Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse Faculty, University of Zurich, Switzerland Institute of Anthropology 2011 based on a true story Why are primates fascinating for digestive physiologists? •!A broad variety of anatomical and physiological digestive adaptations within one taxonomic group •!Experimental work is particularly challenging Major ‘limitation’: ethical responsibility 1000.0000 Caecum fermenter Colon fermenter 100.0000 Nonruminant foregut Ruminant foregut Papio anubis 10.0000 1.0000 0.1000 GIT contents (kg) 0.0100 0.0010 0.0001 0.01 0.10 1.00 10.00 100.00 1000.00 10000.00 BM (kg) from Clauss et al. (2007) Major ‘limitation’: ethical responsibility 1000.0000 Caecum fermenter Colon fermenter 100.0000 Nonruminant foregut Ruminant foregut Papio anubis 10.0000 1.0000 0.1000 GIT contents (kg) 0.0100 0.0010 0.0001 0.01 0.10 1.00 10.00 100.00 1000.00 10000.00 BM (kg) from Clauss et al. (2007) Major limitation: diets used in digestion studies Major limitation: diets used in digestion studies 90 80 70 60 50 40 aD DM (%) 30 20 10 0 0 20 40 60 80 Dietary NDF (%DM) Data collection on langurs from Nijboer & Clauss (2006) Major limitation: diets used in digestion studies 90 80 70 60 50 40 aD DM (%) 30 20 10 0 0 20 40 60 80 Dietary NDF (%DM) Data collection on langurs from Nijboer & Clauss (2006) Major limitation: diets used in digestion studies 90 80 70 ? 60 50 40 aD DM (%) 30 20 10 0 0 20 40 60 80 Dietary NDF (%DM) Data collection on langurs from Nijboer & Clauss (2006) Comparing fibre fractions: Foods in “increasing quality”-order from Gaulin (1979) Comparing fibre fractions: Foods in “increasing quality”-order from Gaulin (1979) Comparing fibre fractions: Foods in “increasing quality”-order from Gaulin (1979) Against many evidence, “fruits” are considered higher quality! Comparing fibre fractions: Foods in “increasing quality”-order from Gaulin (1979) Evident discrepancies in the data given! Outline •!Digestive anatomy •!Digestive physiology I •!The Jarman-Bell principle •!Digestive physiology II •!Feed your monkey Digestive anatomy Hindgut Fermentation - Caecum from Stevens & Hume (1995) Hindgut Fermentation - Colon from Stevens & Hume (1995) Hindgut Fermentation - Colon from Stevens & Hume (1995) Primate caecum/colon fermenters from Stevens & Hume (1995) Caecum volume 1000 ) 100 3 10 olume (cm olume Prosimians Caecum v 1 New World Monkeys Cercopithecine monkeys Colobine monkeys Apes 0.1 10 100 Body length (cm) Data from Chivers & Hladik (1980) Foregut Fermentation from Stevens & Hume (1995) Foregut Fermentation Photos A. Schwarm/ M. Clauss Foregut Fermentation - Ruminant aus Stevens & Hume (1995) Photo Llama: A. Riek Stomach volume 10000 Prosimians New World Monkeys Cercopithecine monkeys ) 3 Colobine monkeys 1000 Apes 100 Stomach volume (cm Stomach volume 10 10 100 Body length (cm) Data from Chivers & Hladik (1980) Stomach volume 100000 Prosimians ) ! 10000 New World Monkeys Cercopithecine monkeys Colobine monkeys Apes 1000 Domestic pig Domestic horse Domestic ruminant Stomach volume (cm Stomach volume 100 Sloth 10 10 100 1000 Body length (cm) Data from Chivers & Hladik (1980) Colobines: fermentation capacity 100 Ruminants Langurs Sloth 10 1 Fermentation contents (kg) Fermentation 0.1 1 10 100 1000 Body mass (kg) Data collection from Parra (1978) Colobines: fermentation capacity Data collection from Schwarm et al. (2010) Digestive constraint I: The ‘foregut fermentation trap‘ Foregut fermentation = Ruminant digestion? Foregut fermentation = Ruminant digestion? Foregut vs. Hindgut Fermentation from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation after enzymatic digestion and absorption: from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation after enzymatic digestion and absorption: ‘Loss’ of bacterial protein, bacterial products (B- Vitamins?)? from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation after enzymatic digestion and absorption: ‘Loss’ of bacterial protein, bacterial products (B- Vitamins?)? (coprophagy) from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation after enzymatic digestion and absorption: ‘Loss’ of bacterial protein, bacterial products (B- Vitamins?)? (coprophagy) Use of easily digestible substrates from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation Fermentation prior to after enzymatic enzymatic digestion and digestion and absorption: absorption: ‘Loss’ of bacterial protein, bacterial products (B- Vitamins?)? (coprophagy) Use of easily digestible substrates from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation Fermentation prior to after enzymatic enzymatic digestion and digestion and absorption: absorption: Use of ‘Loss’ of bacterial bacterial protein, protein, bacterial bacterial products (B- products (B- Vitamins) Vitamins?)? (coprophagy) Use of easily digestible substrates from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation Fermentation prior to after enzymatic enzymatic digestion and digestion and absorption: absorption: Use of ‘Loss’ of bacterial bacterial protein, protein, bacterial bacterial products (B- products (B- Vitamins) Vitamins?)? Bacterial (coprophagy) detoxification? Use of easily digestible substrates from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation Fermentation prior to after enzymatic enzymatic digestion and digestion and absorption: absorption: Use of ‘Loss’ of bacterial bacterial protein, protein, bacterial bacterial products (B- products (B- Vitamins) Vitamins?)? Bacterial (coprophagy) detoxification? Use of easily ‘Loss’ of easily digestible digestible substrates substrates and bacterial modification from Stevens & Hume (1995) Foregut vs. Hindgut Fermentation Fermentation Fermentation prior to after enzymatic enzymatic digestion and digestion and absorption: particularly absorption: Use of suited for ‘Loss’ of bacterial fibre fermen- bacterial protein, protein, bacterial tation bacterial products (B- products (B- Vitamins) Vitamins?) Bacterial (coprophagy) detoxification? Use of easily ‘Loss’ of easily digestible digestible substrates substrates from Stevens und Hume (1995) Intake and Passage in Primates 60 50 40 30 MRT (h) MRT 20 10 0 0 20 40 60 80 100 120 140 DMI (g/kg0.75/d) simple-stomached with forestomach Eulemur/Varecia from Clauss et al. (2008) Intake and Passage in Primates 60 50 40 30 MRT (h) MRT 20 10 0 0 20 40 60 80 100 120 140 DMI (g/kg0.75/d) simple-stomached with forestomach Eulemur/Varecia from Clauss et al. (2008) Intake and Passage in Primates Hindgut fermenters can have either - 60 high or low intake and (hence) short or long ingesta retention 50 40 30 MRT (h) MRT 20 10 0 0 20 40 60 80 100 120 140 DMI (g/kg0.75/d) simple-stomached with forestomach Eulemur/Varecia from Clauss et al. (2008) Intake and Passage in Primates Nonrum. ff appear limited to a low food intake and 60 (hence) long ingesta retention 50 40 30 MRT (h) MRT 20 10 0 0 20 40 60 80 100 120 140 DMI (g/kg0.75/d) simple-stomached with forestomach Eulemur/Varecia from Clauss et al. (2008) Two Preconditions 1.! It is energetically favourable to digest ‘autoenzymatically digestible’ components autoenzymatically, not by fermentative digestion. 2.! Autoenzymatically digestible components are fermented at a drastically higher rate than plant fiber. 80 70 60 50 40 30 20 10 Gas production (ml/h/200mgTS) 0 0 6 12 18 24 Time (h) from Hummel et al. (2006ab) Digestive Strategies Low intake !! long passage High intake !! short passage Digestive Strategies Low intake Autoenzymatic digestion followed ! long passage ! by thorough fermentative ! digestion High intake !! short passage Digestive Strategies Low intake Autoenzymatic digestion followed ! long passage ! by thorough fermentative ! digestion Autoenzymatic digestion followed by cursory ! High intake fermentative !! short passage digestion Digestive Strategies Low intake Autoenzymatic Fermentative digestion digestion followed followed by !! long passage by thorough autoenzymatic ! fermentative ! digestion of products digestion (and remains) Autoenzymatic digestion followed by cursory ! High intake fermentative !! short passage digestion Digestive Strategies Low intake Autoenzymatic Fermentative digestion digestion followed followed by !! long passage by thorough autoenzymatic ! fermentative ! digestion of products digestion (and remains) Autoenzymatic Cursory fermentative digestion mainly of digestion followed autoenzymatically by cursory High intake ! digestible components fermentative followed by ineffective !! short passage digestion autoenzymatic digestion of undigested fiber? Digestive Strategies Low intake Autoenzymatic Fermentative digestion digestion followed followed by !! long passage by thorough autoenzymatic ! fermentative ! digestion of products digestion (and remains) Autoenzymatic Cursory fermentative digestion mainly of digestion followed autoenzymatically by cursory High intake ! digestible components fermentative followed by ineffective !! short passage digestion autoenzymatic digestion of undigested fiber? From Digestive to Metabolic Strategies Low intake !! long passage ! ! !! low BMR High intake ! !! short passage !! high BMR How can you increase fermentative digestive efficiency? •! Digestion of plant fibre by bacteria is the more efficient ... –! the more time is available for it = the longer the mean gastrointestinal retention time. –! the finer the plant fibre particles are = the finer the ingesta is chewed.