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Vertebrate Zoology SCBI 305 • หน้าที่ Circulatory System – ลําเลียง อาหาร O2, CO2, ฮอร์โมน เม็ดเลือด ไปสู่เซลล์ต่างๆใน ร่างกาย – รักษาสภาพอุณหภูมิร่างกาย pH • Closed system Circulatory and Digestive Systems ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. Oxygenated blood ที่มา: Helfman et al. 1997. The Diversity of Fishes. Blackwell Science. Blood passes once through heart Blood passes through heart twice ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. Why are fish hearts small? 1 Vessels of the circulatory system Bidirectional circulation • Aorta • Vein – Blood into heart • Major veins ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. – Common cardinal (sinus) in primitive vertebrates – Portal system (hepatic portal veins, renal portal veins) – Post-, pre- cava (vena cava) – Blood out of heart • Dorsal, ventral aortae • Arteries • Aortic arches Phylogenetic modification • Correlated with functional changes Water Land Gill respiration Lung respiration Pulmonary circulation ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 2 Aortic arches • Primitive pattern of aortic arches: debatable • 6 = # taken as the basic embryonic pattern • Fishes Gills Ventral aorta Aortic arches Afferent artery Capillaries Gills Capillaries 1st arch absent in adult of most fishes Efferent artery Aortic arches Dorsal aorta Body ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. • Sharks, chondricthyans + primitive ray finned fishes Modified into small spiracle (sensory organ) Lobe finned fish (Sarcopterygii) Fishes with supplementary air-breathing organ Lung highly vascularized Little mixing between oxygenated & deoxygenated blood (undivided heart) 3 • Amphibians • Reptiles Larva: arch I, II absent III, IV, V external gills VI lung adult salamander: lost of external gills • Modification of arch IV—systemic arches (double arches) unite behind the heart to form common dorsal aorta • Subdivision of ventral aorta • Arch IV (pulmonary arch) branching out to lung Adult frog: arch III, IV, VI persist • Birds Wing vessels (subclavians) • Mammals Only 3 arches (III, IV, VI) present in adult mammals Pre-dominate systemic arch (left systemic arch not fully developed) Present in turtles & birds serving as major anterior vessels in many birds Pulmonary arch Carotid artery Systemic arch (left predominates) 4 Double systemic arches in amphibians & reptiles Single systemic arch (right) in birds (left) in mammals Aortic arches Systemic arches Pulmonary arch Arterial circuits Venous vessels • 2 systems in double circulation (Not in fishes) – Pulmonary systems lung (found in fishes with lungs, air-breathing vertebrates) – Systemic systems hepatic portal veins, renal portal veins, other veins (few differences among vertebrates, tending to retain basic characters) • Abdominal veins present in fishes, merged/ absent in tetrapods Most similar to basic pattern Hearts • Primitive heart (hagfish) • Original form contractile vessel – In amphioxus • Fishes: 4 chamber heart – Atrium – Ventricle – Conus arteriosus found in many fishes, absent in adult tetrapod – Bulbus arteriosus (lacking cardiac muscle & conal valves) found in most teleosts ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 5 • Lamprey—4 chamber heart Body Collect deoxygenated blood containing pacemaker Sinus venosus Thick wall ที่มา: Helfman et al. 1997. The Diversity of Fishes. Blackwell Science. Chondrichthyes: 4 chambers Atrium Ventricle Conus arteriosus Ventral aorta Gills ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. Body Sinus venosus Small—not chamber-like Atrium ที่มา: Helfman et al. 1997. The Diversity of Fishes. Blackwell Science. Teleosts: 2 chambers Formation of spiral valve in the conus arteriosus for regulating blood flow into left/ right pulmonary arches Ventricle Conus arteriosus—muscular extension of ventricle Bulbus arteriosus Small—not chamber-like Ventral aorta Gills ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 6 • Frog heart: 3 chambers Body – 2 atria, 1 ventricle Right atrium Ventricle Lungs/ skin Close similarity between lungfish & amphibians Left atrium Ventricle Body ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. • Mixed blood in amphibian heart • Why do their hearts evolve in this direction? • Reptiles (lizards) – 5 chambered heart (2 atria, 3 compartments of ventricle) Reduced sinus venosus but same functions Completely divided atrium • Amniote heart: 4 major chambers – 2 atria – 2 ventricles Ventricle with 3 compartments ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 7 • Reptile (alligator) Completely divided ventricle • Cardiac shunt permitting heart to produce different pressures within systemic & pulmonary circuits • Useful ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. • Bird ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 4 chambered heart Reduced sinus venosus Conus arteriosus pulmonary & aortic trunks • Mammal 4 chambered heart Reduced sinus venosus ( Purkinje fiber = pacemaker) in wall of right atrium Conus arteriosus pulmonary & aortic trunks ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 8 • Birds & mammals 4 chambered heart BUT independent evolution – Seen in embryonic development • Adaptation to environment ≠ improvement of design • Incompletely divided heart ≠ imperfect – Perfectly matching with environment • Evolution ≠ progressive improvement – Alternative way of adaptation to demands of different lifestyles Pulmonary circuit Systemic circuit Heat transfer • Another function of the circulatory system • Reptiles basking in the sun absorb heat in their peripheral blood vessels to warm tissues • Body heat dissipates to surface of body via capillaries • Endothermic vs. ectothermic • Homeothermic vs. poikilothermic Animals that maintain constant body temperature Animals that show a change in body temperature • Birds & mammals body temperature regulations using own metabolism • Other vertebrates body temperature depending on environments – Adaptation behaviors (movements) • Effects of climate change • Tolerant ranges, limitations geographic distribution • Adaptations to climate change? 9 Temperature Lower end Limiting factor for distribution in cold environments 135 species 26 species Cold temperature Species cannot survive Hot temperature กิน • Phylogenetic similarity • Higher temperature narrower cold temperature tolerance, NOT increasing hot temperature tolerance Limiting factor for distribution in warm environments Natural selection physiological responses to lower temperature ที่มา: Araujo et al. 2013. Ecology Letters 16: 1206-1219. – Similar responses to temperatures in closely related species Upper end • • • • Eat = put food in mouth, swallow & chew it Consume = Ingest = take food into body usually by swallowing Digest = be changed after eating into a form that the body can use • Absorb = take in a substance from the surface • Egest = discharge undigested food as forces – ≠excretion 10 โครงสร้างระบบทางเดินอาหาร • • • • • • Buccal cavity: palate, teeth, tongue, oral glands Pharynx Esophagus Stomach Intestines: small, large Others • Teeth in fishes – Buccal cavity, pharynx • Tetrapods – Palate, vomer, palatine, pterygoid bones • Crocodiles, birds, mammals Buccal cavity • Palate = roof of the buccal cavity – Fusion of ventral skull bones • Teeth = derivations of dermal armor – Toothless vertebrates agnathans, sturgeons, some toads, turtles, birds, baleen whales – Toothed vertebrates fishes (highly variables), most amphibians, some reptiles, some birds, mammals • Reduction in numbers and distribution (evolution) • Succession of teeth • Tooth replacement in waves (back to front, every other tooth) • Mammals 2 sets of teeth – Jaws 11 • Shape of teeth similar in vertebrates except mammals • Mammalian teeth incisors, canines, premolars, molars (heterodont dentition) • Tongue • Primary tongue (crescent-shaped elevation in the floor of the buccal cavity caused by the underlying hyoid skeleton) – Found in most fishes, primitive amphibians • Primary tongue + glandular field – Found in most amphibians • Primary tongue + glandular field + lateral lingual swelling – Found in reptiles & mammals • Tongue mobility – Immobilized turtles, crocs, some birds, whales – Highly mobile snakes, insectivorous lizards & amphibians & birds – Attached to floor of buccal cavity but some mobility mammals • Oral gland • Saliva – Not found in fishes • Venom (lizards, snakes, mammals) • Anticoagulant (vampire bats) • Functions: Capturing, gathering food (woodpeckers), Taste, Swallowing, Grooming, Thermoregulation (dogs), Human speech 12 Esophagus Pharynx • Fish respiratory organ • Tetrapods foregut Stomach • Muscular chamber • Storage & macerating & secreting digestive enzymes • Weakly developed in cyclostomes • Muscular tube connecting pharynx and stomach • Crop (birds) Diversity of digestive tracts in fishes Straight tube ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. 13 • Differentiated from the esophagus in amphibians, reptiles • Glandular stomach + muscular stomach (gizzard) in birds • Well-developed in mammals – Multi-chambered stomach • Amphibians coiled small intestine, short & straight large intestine • Reptiles & birds coiled small intestine, short large intestine emptying to the cloaca • Mammals coiled, differentiated, long small intestine, long large intestine Intestine • Variation in fishes Spiral value—modification of intestine in chondrichthyans, sturgeons, lungfishes, bichirs, coelacanths, bowfin, gars—increases absorptive surface areas ที่มา: Helfman et al. 1997. The Diversity of Fishes. Blackwell Science. Others • Liver • Gall bladder – Cyclostomes, most birds, some mammals have no gall bladder • Pancreas 14 • Ceca • Cloaca – Pyloric & duodenal ceca common in teleosts (for digestion & absorption) ที่มา: Kardong, KV. 2009. Vertebrates: Comparative Anatomy, Function, Evolution (5th ed.). McGraw-Hill, Inc., New York. – Herbivorous Tetrapods (cellulose digestion) Remain Cloaca well-developed, subdivided, incorporated into other structures Lost • Well-developed in sharks & lungfishes • Remain in amphibians, reptiles, birds, monotremes • Lost in chimaera, ray-finned fishes, coelacanth, eutherian mammals – End of digestive tract (collection of intestine, urinary, genital ducts) open to outside – Opening to outside through a cloacal opening/ vent – Latin: cloaca = sewer – Found in all vertebrates during embryonic development – Shallow or none in lampreys, ray-finned fishes, mammals (except monotremes) – Anus (end of digestive tract if no cloaca present) Diet change • Seasonal variation – Opportunistic feeders – Generalists – Specialists • Ontogenetic niche shift – Evolutionary adapatation – Anus 15 Ontogenetic niche shift • Irreversible, genetically determined • Morphological changes, age – Habitat change? • Driving forces (Claessen & Dieckmann 2002—Evolutionary Ecology Research vol. 4: 189-217) – Predator avoidance – Niche variation 16