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Avian Gastrointestinal Anatomy and Physiology Kirk C Avian Gastrointestinal Anatomy and Physiology Kirk C. Klasing, BS, MS, PhD A bird's gastrointestinal (GI) tract morphology, diges- their GI tracts appear to be morphologically tive strategy, and metabolic capability have been inti- more similar to chickens and turkeys than to mately intertwined during evolution to match the nutri- ent content and physical attributes of foods available in birds with very different dietary patterns, such as its natural habitat. The most commonly kept compan- carnivorous or herbivorous species. This permits ion species are granivorous with a tendency toward us to use the well-studied chicken as a point of omnivory. The beak, oral cavity, and tongue of granivo- reference when examining the GI tracts of com- rous birds have anatomic adaptations for shelling seeds, panion avian species. including ridges in the tomia of their beak for slicing the hull and a dexterous tongue for manipulating the seed The GI tract of a bird provides an environ- and disposing the hull. Granivorous species possess a ment for the physical and chemical reduction in sizeable crop or an expandable esophageal pouch for the size and molecular complexity of food and storing food so that large meals can be consumed. Their then absorbs the end products of digestion, proventriculus is somewhat small, and their gizzard is which are needed in widely differing quantities. large, muscular, and possesses a thick cuticle relative to carnivores or frugivores. A vestigial ceca and short A young chick requires a diet with more than rectum provide little area for use of symbiotic bacteria 15% protein, but only 0.0000003% of vitamin to aid in the digestion of the fibrous components of B12. The anatomy and digestive functions of the food. GI tract are designed to accommodate this wide Copyright 1999 by W. B. Saunders Company. range of quantitative needs. The GI tract has Key words: Digestion, companion birds, gastrointesti- sufficient morphological plasticity to accommo- nal tract. date changes in nutritional needs during the life cycle and to adapt to changing physical and nutritional characteristics of the diet. ~,4 re than 9,000 species of birds (more than ce as many as mammals) fill many of the possible niches within the earth's food web. A bird's gastrointestinal (GI) tract morphology, Anatomy and Physiology digestive strategy, and metabolic capability have of the Digestive System been intimately intertwined during evolution to The avian digestive tract is a continuous tube match the nutrient content and physical at- that opens at either end (beak and vent) to the tributes of foods available in its natural habitat. outside world and consists of a mouth, esopha- When compared across species, the GI tract is gus, crop, proventriculus, ventriculus or gizzard, the most anatomically diverse organ system. intestine, ceca, rectum, and cloaca (Fig 1). As However, distantly related species consuming food progresses through these organs, a specific similar food items often display morphological sequence of digestive events occurs, including convergence because of similar nutritional and grinding, acidifying, hydrolyzing, emulsifying, ecological selection pressures. 1,2 The most com- and transporting of the end products. monly kept companion species are granivorous The distinctive anatomy and physiology of the with a tendency toward omnivory (Table 1), and avian GI tract reflects the constraints of flight, in that most of the tract's weight is centralized within the body cavity to optimize aerial maneu- From the Department of Animal Science, University of Califor- verability. The avian GI tract has a larger number nia, Davis, Davis, CA. of organs, which have greater interorgan coopera- Address reprint requests to Kirk C. Klasing; BS, MS, PhD, tion than their mammalian counterparts. The Department of Animal Science, University of California, Davis, One Shields Ave, 2131 B. Meyo'Hall, Davis, CA 95616. precise anatomic plan of the digestive tract of Copyright 1999by W. B. Saunders Company. companion birds varies somewhat, depending 1055-937X/99/0802-0002510. 00/0 on their typical diet. 42 Seminars in Avian and Exotic Pet Medicine, Vol 8, No 2 (April), 1999:pp 42-50 Avian Digestion 43 Table 1. Eating Habits of Some Species of Captively naso-frontal hinge, permitting an increased gape Propagated Birds of the beak and providing flexion that absorbs Mostly granivorous species some of the shock associated with pecking and Gouldian Finch seed cracking. 5,6 Zebra Finch The hyoid apparatus, consisting of multiple Canary articulating bones and their associated muscula- Budgerigar Cockatiel ture, mobilizes the tongue. Psittacidae are unique Domestic Pigeons among birds in that they have additional muscles Omnivorous, but tending to be granivorous in the anterior regions of their tongues that are Conures independent of the hyoid apparatus and permit Rosella added flexibility. 7 The tongues of lories and Sulfur-crested Cockatoo African Grey Parrot lorikeets are relatively long and end in fine, Amazon Parrot hairlike processes for the collection of sap or Macaw nectar through capillary action, s Tongues of Chicken many other psittacine species are thick and Omnivorous, but tending to be frugivorous muscular, acting as fingers for the manipulation Toucan Mynah and extraction of seeds from their husks or from Barbets cones. The tongues of passerines, such as finches Omnivores, but tending to be nectarivorous and fru- and canaries, are narrow, short, and not very givorous muscular. Lofikeets The anterior part of the oral cavity is roofed Lories with a hard palate, which in finches, canaries, budgerigars, and cockatiels contains two ridges Beak, Tongue, and Oral Cavity that facilitate removal of the shell from seeds The beak, tongue, and oral cavity function in before consumption. Typically, granivorous spe- grasping, testing, mechanical processing, and cies have well-developed compound tubular sali- lubricating food and propelling the food to the vary glands scattered in groups around the lower esophagus. Their specific morphology reflects oral cavity, tongue, and pharynx. the physical requirements of obtaining and pro- The rapid transit of food through the mouth, cessing food items. The granivorous tendency of lack of mastication, relatively low saliva addition most companion birds is evident in their short to the food, and low numbers of taste receptors stout beaks with pronounced ridges designed for result in poor taste acuity relative to humans. cracking seeds and dexterous tongue for manipu- Parrots have about 350 taste receptors, com- lating the seed when shelling it. pared with 9,000 in hmnans. 9 Most taste buds are A keratin sheath (rhamphotheca) growing located on the palate and on the posterior from the upper and lower mandible forms the tongue. The palatal taste buds are usually located beak. The keratin is continually lost by wear and in regions where the epithelium is soft and replaced by new growth. The location and rate of glandular, typically near the salivary glands. The growth and wear influence the exact shape of the tongue, oral cavity, and beak have a rich supply beak, and subtle changes may occur as food types of touch receptors that augment the bird's rela- change. The tomia, or outer edges, are some- tively poor gustatory capacity with a strong tactile what sharp to facilitate cutting seed coats. Sev- sense. eral structural attributes of the beak, jaws, and Esophagus and Crop skull provide unique functional characteristics that are important in obtaining food. The attach- The esophagus extends down the neck into ment of the lower beak to the skull is somewhat the thoracic cavity and terminates in the proven- loose, permitting a large gape. A bird's gape triculus. In the budgerigar, the esophagus is places the upper limit on the size of food item dorsal to the trachea in the anterior regions of that can be consumed and is particularly large in the neck and then runs along the right side. 1~ frugivorous species (eg, toucans). A second struc- Peristaltic contraction of inner circular and outer tural adaptation found in parrots is the articula- longitudinal muscles in the tunica muscularis tion of the upper mandible in the cranium at the propels food posteriorly through the esophagus. 44 Kirk C. Klasing Cro :ulus izzard Pancreas / "Cloacal bursa Cloaca lal loop loop Figure 1. The digestive tract of a Budgerigar. (Reprinted with permission. 10 ) To aid in swallowing large food items, the esopha- side of the neck, and this organ extends trans- gus is expandable as a result of a series of versely to the left side. The presence of large longitudinal folds. This accordion-like arrange- folds of mucosa allows considerable expansion ment is enriched with mucous glands to provide and shrinkage, depending on the amount of lubrication. The epithelial lining is thick and contents. Immediately cranial to the thoracic cornified for protection against mechanical dam- inlet, the crop narrows to reform an esophageal age as a result of swallowing food items whole. In tube and passes between the coracoid bones to many species, the esophagus widens into a crop the right of the syrinx and dorsal to the heart, i~ just before entering the thoracic cavity. In the Some granivorous species (eg, Cardueline budgerigar, the inlet to the crop is on the right Finches) do not have a true crop, but they Avian Digestion 45 usually have a very expandable esophageal pouch The thickness and physical properties of the that can store food items. cuticle are highly correlated with food con- sumed, being especially thick in granivores, but Stomach thin and soft in frugivores and nectarivores. It is The stomach consists of the proventriculus thickest and toughest directly under the thick (glandular stomach) and the gizzard (muscular muscles, which provide much of the grinding stomach), which is sometimes called the ventricu- within the gizzard. The cuticle lining wears down lus. The proventriculus is the site at which steadily and undergoes continual renewal.
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