Topic 13: Energetics & Performance. How are gas exchange, circulation & metabolism inter-related?

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Topic 3: Energetics & Performance How are gas exchange, circulation & metabolism interrelated? How is it done in air and water? What organs are involved in each case? How does ventilation differ among taxa? What are the main components of circulation? How does circulation work? How does the anatomy of the heart differ among taxa? What are the different types of metabolic rates? What are the two types of metabolism? How is performance influenced by temperature? How are gas exchange, circulation & metabolism inter-related? What are the components of the gas exchange system? What are the components of the circulatory system? How are gas exchange, circulation & metabolism interrelated? Mammalian circulation ignore the details McGraw-Hill

How are gas exchange, circulation & metabolism inter-related? Mammalian circulation ignore the details McGraw-Hill

Two main purposes from environment to tissues _ from tissues Gas exchange occurs in both air and water Reptiles generally exchange gases in air Amphibians do this in both air and water Ambystoma tigrinum Two types of structures for respiration _ Lungs _ Gills, skin, pharynx, cloaca Amphibians have many nonpulmonary gas exchange structures (also often lungs) Reptile have predominantly pulmonary gas exchange R Tramontano, PJB Non-pulmonary: Amphibians Gas exchange and water balance are related through same structures Gas exchange structures: plethodontids (cutaneous respiration) All amphibians Especially hellbender, hairy frog All larval amphibians Paedomorphic adults Non-pulmonary: Reptiles Not prevalent because of keratinized skin Hydrophiinae (sea snakes) Some Testudines Rheodytes leukops, an Australian Pleurodire Has lined with villi Pump water in and out of bursae ~80 times/minute http://www.divegallery.com/sea_krait.htm Stebbins & Cohen 995, Fig. 3.5; EJ Routman Pough et al 2004, Fig 4-35 Pulmonary using lungs Widespread (obviously) Requires of the lungs Exhalation of poor, C rich air Inhalation of rich, C poor air Most amphibians use Pulmonary ventilation Lepidosaurs use Expansion of thoracic cavity creates negative pressure Inhalation In lizards, air moves between lungs during locomotion In snakes, pressure changes In lung where gas exchange occurs Not much in part, where air is stored Liem et al. Fig. 8-4 Pough et al 2004, Fig 7-5, 7-6

Pulmonary ventilation Testudines have a rigid shell mm. transversus abdominis & pectoralis control size of visceral cavity, facilitating ventilation Crocodylia use the liver m. diaphragmaticus pulls liver anteriorly This compresses lungs, leading to exhalation Components of the circulatory system Heart Vessels (veins, arteries, capillaries) Blood The heart pumps the blood through the vessels to: Lungs/gills Organs Muscles Transports & Mammalian circulation ignore the details Pough et al 2004, Fig 7-7 McGraw-Hill pattern depends on gas exchange structure Gills (larval amphibians) _ Heart Heart -poor blood goes to gills -rich blood goes to body Same as fishes Lungs (most others) Heart Heart Heart Pulmonary circuit has lower blood pressure than systemic circuit Variation in how heart is divided among taxa www.universe-review.ca www.universe-review.ca arteries Paired vessels in adult amphibians Divide into: Pulmonary arteries Cutaneous arteries Skin is part of pulmonary circuit, provides: 20-90% uptake 30-00% C release What amphibians would have almost all gas exchange through skin? Variation in heart morphology: Anura 3 chambers Oxygenated and deoxygenated blood mix in heart! and channels formed by trabeculae help keep rich and poor blood separate Good enough for slower metabolism Photo EJ Routman Pough et al 2004, Fig 7-8 2

Variation in heart morphology: Testudines & Squamata 3 chambers (2 atria, ventricle) Ventricle subdivided into 3 chambers: Variation in heart morphology: Crocodylia 4 chambered heart: poor rich Atrioventricular valve blocks interventricular canal between CA & CV Muscular ridge divides CV & CP Separation of pulmonary and systemic circuits acts as a shunt Allows the movement of blood between circuits poor rich Pough et al 2004, Fig 7-9 Pough et al 2004, Fig 7-0 What are shunts for? Controlling [ ] affects ph Can supply more oxygen to tissues when animal is not breathing Controlling amount of blood in the systemic circuit affects heating and cooling rates R L shunt Blood bypasses the lungs during diving, etc. The use of sugars to produce energy This energy is used for: Movement Reproduction Anything else the animal does Sugars are transported to cells by Sugars are metabolized in Proceedings of the Royal Society Photos PJB, B Rourke Metabolic rate Varies with activity metabolic rate Minimum energy consumption to stay alive Postabsorptive, inactive, inactive part of day metabolic rate Postabsorptive, inactive, but alert metabolic rate Maximal aerobic rate At cusp of switching to anaerobic metabolism Maximal intake Maximum metabolic rate occurs at maximum aerobic speed (MAS) During sustainable activity Glucose is metabolized Water and C are produced During unsustainable activity Glycogen is metabolized Lactate is produced, leading to fatigue Pough et al. 2004, Fig 7- Pough et al. 2004, Fig 7-

All metabolism produces ATP, which is used as energy is aerobic sugar 35 ATP + CO2 +H2O Efficient, but slow Good for sustained activity is anaerobic sugar 3 ATP + Lactic Acid Inefficient but fast Good for short term, high output activity Sceloporus jarrovii use glycolytic metabolism during territorial defense Routine activity Little build up Territorial defense Intense Includes running and biting other males [Lactate] proportional to# bites/minute www.palaeos.com PJB; Pough et al. 2004 Fig 7-3 Temperature affects performance in ectotherms Affects locomotion in Phelsuma dubia Log max velocity (m/s) At high temps, they take Stride duration is affected by how fast muscles can Stride length is Velocity 0.5 0.0-0.5 -.0 0 5 20 25 30 35 40 Temperature (C) a ab b Stride Duration (s) 0.3 0.2 0. Stride Duration a ab bc cd d Stride Length (mm) 0 0.0 0 5 20 25 30 35 40 0 5 20 25 30 35 40 Temperature (C) Temperature (C) Bergmann & Irschick 2006; Photo PJB 80 70 60 50 40 30 20 0 Stride Length a ab b b Post-meal metabolism in snakes Digestion uses a lot of energy, especially when meal is big Some pythons consume ~0X more oxygen at peak digestion Also, intestine of a fed snake: More blood More mucous Thicker intestinal lining Ott & Secor 2007 Time

Calling in frogs is also energetically expensive The higher the calling rate, the higher the metabolic rate Some call 500X/hour V0 2 increases 0X??; Pough et al. 2004, Fig 7-8 Calling in frogs is energetically expensive but is it worth it? Pough et al. 2004, Fig 7-9, 7-20