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==What are Carbohydrates?==

==What are Carbohydrates?==

Carbohydrates (saccharides) are a group of '''carbon-based molecules''', that are naturally found as '''monosaccharides, disaccharides, or polysaccharides'''.<ref name="NRC">National Research Council (NRC). (2006) '''Carbohydrates and Fiber. In Nutrient Requirements for Dogs and Cats.''' 2006 ''Washington, DC: National Academies Press ''p.49-80.</ref> Carbohydrates are a common source of energy, however they are '''not considered an essential nutrient in dogs or cats'''. Functionally carbohydrates are classified as '''absorbable, digestible, fermentable''' or '''non-fermentable'''. When consumed in food, absorbable and digestible carbohydrates typically provide approximately 4 kcal of energy per gram, whereas fermentable and non-fermentable carbohydrates provide little to no usable energy.   

Carbohydrates (saccharides) are a group of '''carbon-based molecules''', that are naturally found as '''monosaccharides, disaccharides, or polysaccharides'''.<ref name="NRC">National Research Council (NRC). (2006) '''Carbohydrates and Fiber. In Nutrient Requirements for Dogs and Cats.''' 2006 ''Washington, DC: National Academies Press ''p.49-80.</ref> Carbohydrates are a common source of [[Energy - Nutrition|energy]], however they are '''not considered an essential nutrient in dogs or cats'''. Functionally carbohydrates are classified as '''absorbable, digestible, fermentable''' or '''non-fermentable'''. When consumed in food, absorbable and digestible carbohydrates typically provide approximately 4 kcal of energy per gram, whereas fermentable and non-fermentable carbohydrates provide little to no usable energy.   

[[Sugars - Nutrition|'''Absorbable''' carbohydrates]] include monosaccharides, such as the '''glucose, fructose,''' and '''galactose'''; the sugar alcohols '''mannitol, sorbitol''' and '''xylitol'''; disaccharides, such as '''lactose''' (glucose + galactose), '''sucrose '''(glucose + fructose) and '''maltose''' (glucose + glucose); and the polysaccharides such as '''[[Starch - Nutrition|starch]], glycogen''' (from animal muscle and liver) and '''amylose''' and '''amylopectin''' from plants.

[[Sugars - Nutrition|'''Absorbable''' carbohydrates]] include monosaccharides, such as the '''glucose, fructose,''' and '''galactose'''; the sugar alcohols '''mannitol, sorbitol''' and '''xylitol'''; disaccharides, such as '''lactose''' (glucose + galactose), '''sucrose '''(glucose + fructose) and '''maltose''' (glucose + glucose); and the polysaccharides such as '''[[Starch - Nutrition|starch]], glycogen''' (from animal muscle and liver) and '''amylose''' and '''amylopectin''' from plants.

==Roles in the Body==

==Roles in the Body==

#'''Energy Production''': All cells in the body have a requirement for '''glucose''' and it must be obtained regularly either from the diet or synthesised through hepatic gluconeogenesis. Glucose absorbed from the diet can be used directly in intermediate metabolism (ATP production) or to synthesise glycogen and fatty acid.<ref name="Ebiner">Ebiner JR, ''et al.'' (1979)''' Comparison of carbohydrate utilization in man using indirect calorimetry and mass spectrometry after oral load of 100 g naturally-labelled (13C) glucose.''''' Br J Nutr'' 1979;41:419-429.</ref><ref name="Flatt">Flatt JP, ''et al''. (1985)'''Effects of dietary fat on postprandial substrate oxidation and on carbohydrate and fat balances.''''' J Clin Invest ''1985;76:1019-1024.</ref> In the absence of dietary starches or sugars, '''hepatic gluconeogenesis''' can support maintenance of normal blood glucose levels, gluconeogenic amino acids and glycerol in dogs<ref>Romsos DR,'' et al.'' (1976)''' Effects of dietary carbohydrate, fat and protein on growth, body composition, and blood metabolite levels in the dog. '''''J Nutr ''1976;106:1452-1456.</ref> and cats.<ref name="Morris">Morris JG, ''et al.'' (1977)''' Carbohydrate digestion in the domestic cat ''(Felis catus)'''. Br J Nutr'' 1977;37:365-373.</ref>

#'''Energy Production''': All cells in the body have a requirement for '''glucose''' and it must be obtained regularly either from the diet or synthesised through hepatic gluconeogenesis. Glucose absorbed from the diet can be used directly in intermediate metabolism (ATP production) or to synthesise glycogen and [[Fatty Acids Overview - Nutrition|fatty acids]].<ref name="Ebiner">Ebiner JR, ''et al.'' (1979)''' Comparison of carbohydrate utilization in man using indirect calorimetry and mass spectrometry after oral load of 100 g naturally-labelled (13C) glucose.''''' Br J Nutr'' 1979;41:419-429.</ref><ref name="Flatt">Flatt JP, ''et al''. (1985)'''Effects of dietary fat on postprandial substrate oxidation and on carbohydrate and fat balances.''''' J Clin Invest ''1985;76:1019-1024.</ref> In the absence of dietary starches or sugars, '''hepatic gluconeogenesis''' can support maintenance of normal blood glucose levels, gluconeogenic amino acids and glycerol in dogs<ref>Romsos DR,'' et al.'' (1976)''' Effects of dietary carbohydrate, fat and protein on growth, body composition, and blood metabolite levels in the dog. '''''J Nutr ''1976;106:1452-1456.</ref> and cats.<ref name="Morris">Morris JG, ''et al.'' (1977)''' Carbohydrate digestion in the domestic cat ''(Felis catus)'''. Br J Nutr'' 1977;37:365-373.</ref>

#'''Intestinal Health''': Non-absorbable carbohydrates (oligosaccharide and polysaccharide dietary fibres) are resistant to degradation by mammalian enzymes. These carbohydrates are more commonly referred to as '''dietary fibres''' and can be divided into two broader categories depending on whether they can be further metabolized (fermented) by intestinal bacteria or not.<ref name="NRC" />  

#'''Intestinal Health''': Non-absorbable carbohydrates (oligosaccharide and polysaccharide [[Fibre - Nutrition|dietary fibres]]) are resistant to degradation by mammalian enzymes. These carbohydrates are more commonly referred to as '''dietary fibres''' and can be divided into two broader categories depending on whether they can be further metabolized (fermented) by intestinal bacteria or not.<ref name="NRC" />  

#'''Fermentable, non-absorbable carbohydrates '''can be utilized as an energy substrate by bacteria in the [[Large Intestine Overview - Anatomy & Physiology|large intestine]]. By-products of bacterial fermentation include CO, H₂, methane, and the short-chain fatty acids acetate, propionate and butyrate, which can support optimal colonocyte function and intestinal health. Diffusion of acetate and proprionate across the colonic mucosa facilitates water reabsorption. Butyrate is the preferred energy substrate of colonocytes.<ref>Herschel DA, ''et al.'' (1981)''' Absorption of volatile fatty acids and H₂O by the colon of the dog. '''''AJVR ''1981;42:1118-1124.</ref><ref>Reinhart GA,'' et al.'' (1994)''' Source of dietary fiber and its effects on colonic microstructure, function and histopathology of the beagle dogs. '''''J Nutr ''1994;124:2701S-2703S.</ref><ref>Howard MD,'' et al.'' (1999)''' Blood flow and epithelial cell proliferation of the canine colon are altered by source of dietary fiber. '''''Vet Clin Nutr'' 1999;6:8-15.</ref>

#'''Fermentable, non-absorbable carbohydrates '''can be utilized as an energy substrate by bacteria in the [[Large Intestine Overview - Anatomy & Physiology|large intestine]]. By-products of bacterial fermentation include CO, H₂, methane, and the short-chain fatty acids acetate, propionate and butyrate, which can support optimal colonocyte function and intestinal health. Diffusion of acetate and proprionate across the colonic mucosa facilitates water reabsorption. Butyrate is the preferred energy substrate of colonocytes.<ref>Herschel DA, ''et al.'' (1981)''' Absorption of volatile fatty acids and H₂O by the colon of the dog. '''''AJVR ''1981;42:1118-1124.</ref><ref>Reinhart GA,'' et al.'' (1994)''' Source of dietary fiber and its effects on colonic microstructure, function and histopathology of the beagle dogs. '''''J Nutr ''1994;124:2701S-2703S.</ref><ref>Howard MD,'' et al.'' (1999)''' Blood flow and epithelial cell proliferation of the canine colon are altered by source of dietary fiber. '''''Vet Clin Nutr'' 1999;6:8-15.</ref>

#'''Non-fermentable, non-absorbable carbohydrates''' include structural components of plant cell walls such as cellulose, lignin and bran. These forms of carbohydrates resist degradation by animal or bacterial enzymes and pass through the intestinal tract intact.

#'''Non-fermentable, non-absorbable carbohydrates''' include structural components of plant cell walls such as cellulose, lignin and bran. These forms of carbohydrates resist degradation by animal or bacterial enzymes and pass through the intestinal tract intact.

====Cats and Carbohydrates:====  

====Cats and Carbohydrates:====  

Cats are able to digest and absorb dietary sugars and starches well<ref name="Morris" /> but have low glucokinase activity in the liver<ref>Tanaka A, ''et al.'' (2005)''' Comparison of expression of glucokinase gene and activities of enzymes related to glucose metabolism in livers between dog and cat. '''''Vet Res Commun ''2005;29:477-485. </ref> and do not adapt carbohydrate metabolism to dietary intake.<ref>Buddington RK, ''et al.'' (1991) '''Dietary regulation of intestinal brush-border sugar and amino acid transport in carnivores.''''' Am J Physiol ''1991;261:R793–801.</ref> There has been controversy over the role of dietary carbohydrate in development of obesity and [[DM|diabetes mellitus]] in cats but carbohydrate intake has not been shown to be a risk factor in development of obesity,<ref> Backus RC, ''et al.'' (2007)''' Gonadectomy and high dietary fat but not high dietary carbohydrate induce gains in body weight and fat of domestic cats.''''' Br J Nutr ''2007;98:641-650.</ref> hyperglycaemia<ref>Hoenig M, ''et al.'' (2012)''' Evaluation of long-term glucose homeostasis in lean and obese cats using continuous glucose monitoring.''''' AJVR'' 2012:73:1100-1106.</ref> or diabetes mellitus<ref>Verbrugghe A,'' et al.'' (2012)''' Nutritional modulation of insulin resistance in the true carnivorous cat: a review. '''''Crit Rev Food Sci Nutr'' 2012;52:172–182.</ref> in otherwise healthy adult cats.

Cats are able to digest and absorb dietary sugars and starches well<ref name="Morris" /> but have low glucokinase activity in the liver<ref>Tanaka A, ''et al.'' (2005)''' Comparison of expression of glucokinase gene and activities of enzymes related to glucose metabolism in livers between dog and cat. '''''Vet Res Commun ''2005;29:477-485. </ref> and do not adapt carbohydrate metabolism to dietary intake.<ref>Buddington RK, ''et al.'' (1991) '''Dietary regulation of intestinal brush-border sugar and amino acid transport in carnivores.''''' Am J Physiol ''1991;261:R793–801.</ref> There has been controversy over the role of dietary carbohydrate in development of obesity and [[DM|diabetes mellitus]] in cats but carbohydrate intake has not been shown to be a risk factor in development of obesity,<ref> Backus RC, ''et al.'' (2007)''' Gonadectomy and high dietary fat but not high dietary carbohydrate induce gains in body weight and fat of domestic cats.''''' Br J Nutr ''2007;98:641-650.</ref> hyperglycaemia<ref>Hoenig M, ''et al.'' (2012)''' Evaluation of long-term glucose homeostasis in lean and obese cats using continuous glucose monitoring.''''' AJVR'' 2012:73:1100-1106.</ref> or [[Diabetes Mellitus|diabetes mellitus]]<ref>Verbrugghe A,'' et al.'' (2012)''' Nutritional modulation of insulin resistance in the true carnivorous cat: a review. '''''Crit Rev Food Sci Nutr'' 2012;52:172–182.</ref> in otherwise healthy adult cats.

==Consequences of Deficiency==

==Consequences of Deficiency==

The sugar-alcohol, '''xylitol''' (used as a low glycaemic index sweetener in many human foods) '''is toxic to dogs and cats''' and ingestion can lead to severe hypoglycaemia, [[Liver Failure|liver failure]] and death.<ref>Xia Z, ''et al.'' (2009)''' Experimental acute toxicity of xylitol in dogs.''''' J Vet Pharmacol Ther ''2009;32:465-469.</ref> No toxicity has been associated with high intake of other carbohydrates in otherwise healthy dogs and cats, though in animals with pre-existing diabetes mellitus increased intake of sugars and starches can contribute to post-prandial hyperglycaemia and increase [[insulin]] requirements.<ref>Bennett N, ''et al.'' (2006)''' Comparison of a low carbohydrate-low fiber diet and a moderate carbohydrate-high fiber diet in the management of feline diabetes mellitus. '''''J Feline Med Surg ''2006;8:73-84.</ref><ref>Elliot KF, ''et al.'' (2012)''' A diet lower in digestible carbohydrate results in lower postprandial glucose concentrations compared with a traditional canine diabetes diet and an adult maintenance diet in healthy dogs.''''' Res Vet Sci'' 2012;93:288-295.</ref>

The sugar-alcohol, '''xylitol''' (used as a low glycaemic index sweetener in many human foods) '''is toxic to dogs and cats''' and ingestion can lead to severe hypoglycaemia, [[Liver Failure|liver failure]] and death.<ref>Xia Z, ''et al.'' (2009)''' Experimental acute toxicity of xylitol in dogs.''''' J Vet Pharmacol Ther ''2009;32:465-469.</ref> No toxicity has been associated with high intake of other carbohydrates in otherwise healthy dogs and cats, though in animals with pre-existing diabetes mellitus increased intake of sugars and starches can contribute to post-prandial hyperglycaemia and increase [[insulin]] requirements.<ref>Bennett N, ''et al.'' (2006)''' Comparison of a low carbohydrate-low fiber diet and a moderate carbohydrate-high fiber diet in the management of feline diabetes mellitus. '''''J Feline Med Surg ''2006;8:73-84.</ref><ref>Elliot KF, ''et al.'' (2012)''' A diet lower in digestible carbohydrate results in lower postprandial glucose concentrations compared with a traditional canine diabetes diet and an adult maintenance diet in healthy dogs.''''' Res Vet Sci'' 2012;93:288-295.</ref>

Excessive intake of non-absorbable carbohydrates (both fermentable and non-fermentable dietary fibres) can increase stool bulk and slow gastrointestinal transit time.<ref>Eastwood MA. (1992) '''The physiologic effects of dietary fiber: An update. '''''Annu Rev Nutr ''1992;12:19-35.</ref><ref>Lewis LD, ''et al''. (1994)''' Stool characteristics, transit time, and nutrient digestibility in dogs fed different fiber sources.''''' J Nutr ''1994;124:2716S-2718S.</ref> This may potentially result in constipation in healthy dogs and cats or worsening dysmotility in animals with underlying intestinal disease.  

Excessive intake of non-absorbable carbohydrates (both fermentable and non-fermentable dietary fibres) can increase stool bulk and slow gastrointestinal transit time.<ref>Eastwood MA. (1992) '''The physiologic effects of dietary fiber: An update. '''''Annu Rev Nutr ''1992;12:19-35.</ref><ref>Lewis LD, ''et al''. (1994)''' Stool characteristics, transit time, and nutrient digestibility in dogs fed different fiber sources.''''' J Nutr ''1994;124:2716S-2718S.</ref> This may potentially result in [[Colonic Impaction - Dog and Cat|constipation]] in healthy dogs and cats or worsening dysmotility in animals with underlying intestinal disease.  

Undigested disaccharides and [[Starch - Nutrition|starches]] can act as osmotic agents drawing water into the intestinal lumen and can be fermented by intestinal bacteria.<ref>Washabau RJ,'' et al.'' (1986)''' Evaluation of intestinal carbohydrate malabsorption by pulmonary hydrogen gas excretion.''''' AJVR ''1986;47:1402-1406.</ref><ref>Muir P, ''et al. '' (1991) '''Evaluation of carbohydrate malassimilation and intestinal transit time in cats by measurement of breath hydrogen excretion. '''''AJVR'' 1991;52:1104-1109.</ref> Maldigestion and malabsorption of dietary carbohydrate (specifically starch) may be a feature of feline [[Inflammatory Bowel Disease|inflammatory bowel disease]].<ref>Ugarte C, ''et al.'' (2004)''' Carbohydrate malabsorption is a feature of feline inflammatory bowel disease but does not increase clinical gastrointestinal signs. '''''J Nutr'' 2004;134:2068S–2071S.</ref>

Undigested disaccharides and [[Starch - Nutrition|starches]] can act as osmotic agents drawing water into the intestinal lumen and can be fermented by intestinal bacteria.<ref>Washabau RJ,'' et al.'' (1986)''' Evaluation of intestinal carbohydrate malabsorption by pulmonary hydrogen gas excretion.''''' AJVR ''1986;47:1402-1406.</ref><ref>Muir P, ''et al. '' (1991) '''Evaluation of carbohydrate malassimilation and intestinal transit time in cats by measurement of breath hydrogen excretion. '''''AJVR'' 1991;52:1104-1109.</ref> Maldigestion and malabsorption of dietary carbohydrate (specifically starch) may be a feature of feline [[Inflammatory Bowel Disease|inflammatory bowel disease]].<ref>Ugarte C, ''et al.'' (2004)''' Carbohydrate malabsorption is a feature of feline inflammatory bowel disease but does not increase clinical gastrointestinal signs. '''''J Nutr'' 2004;134:2068S–2071S.</ref>

==Dietary Sources==

==Dietary Sources==

Dietary carbohydrates (both absorbable and non-absorbable) are primarily found in as '''fruits, vegetables, cereal grains, and pulses '''(i.e., legumes). Animal sourced carbohydrates (glycogen) are also found in '''muscle and liver'''.  

Dietary carbohydrates (both absorbable and non-absorbable) are primarily found in '''fruits, vegetables, cereal grains, and pulses '''(i.e., legumes). Animal sourced carbohydrates (glycogen) are also found in '''muscle and liver'''.  

==Diagnosing Carbohydrate Deficiency==

==Diagnosing Carbohydrate Deficiency==

[[Category:To Do - Nutrition]]

[[Category:To Do - Nutrition]]

[[Category:To Do - Nutrition GGP]]

[[Category:To Do - Nutrition preMars]]