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ANIMAL PHYSIOLOGY BIOL 3151: Principles of Animal Physiology Dr. Tyler Evans Email: [email protected] Phone: 510-885-3475 Office Hours: F 8:30-11:30 or appointment Website: http://evanslabcsueb.weebly.com/ PREVIOUS LECTURE CARDIOVASCULAR PHYSIOLOGY CHARACTERISTICS OF CIRCULATORY SYSTEMS • circulatory systems can either be OPENED or CLOSED • in a CLOSED circulatory system, fluid remains within blood vessels at all points and substances must diffuse across the walls of blood vessels to enter tissues • in an OPEN circulatory system, fluid enters a SINUS (space) at some point and there comes into direct contact with tissues allowing exchange • there is often uncertainty as to which type of system an animal possesses • decapod crustaceans have both sinuses and fine branching blood vessels. Their circulatory systems are usually classified as open, but like closed systems, diffusion can across the membrane of some fine blood vessels textbook Fig 8.8 pg 356 PREVIOUS LECTURE CARDIOVASCULAR PHYSIOLOGY ADVANTAGES OF A CLOSED CIRCULATORY SYSTEM • closed circulatory systems provide two main advantages over open systems: 1. ability to generate high pressure and flow rates 2. ability to better control and direct blood flow to specific tissues • these features are important for oxygen delivery to metabolically active tissue and closed systems tend to be found in highly active organisms or those living in low-oxygen environments PREVIOUS LECTURE CARDIOVASCULAR PHYSIOLOGY VERTEBRATE CIRCULATORY SYSTEMS • although left and right sides of the heart are joined together in a single organ, in birds and mammals these sides are functionally separated • functionally, more like a single circuit with two pumps • having separated pulmonary and systemic circuits has an important advantage: allows pressure in each circuit to be different • in lungs, capillaries must be thin to allow for gas exchange and cannot withstand high pressure • in contrast high pressure is needed in the systemic circuit to force blood throughout the body PREVIOUS LECTURE VERTEBRATE CIRCULATORY SYSTEMS • trade-off to completely separated circuit is that the circulatory system becomes relatively inflexible • for example, if a mammal holds it’s breath, blood must still flow to the lungs despite that the fact that blood is not becoming oxygenated when it arrives • because breathe continuously, has been selection pressure to divert blood flow from pulmonary circuit • unlike birds and mammals, amphibians and reptiles have incompletely divided hearts and because ventricles of the heart are interconnected, blood can be diverted from pulmonary to systemic circuit or vice-versa. textbook Fig 8.13 pg 362 TODAY’S LECTURE WHAT IS A HEART? • chambered hearts, like those in humans, evolved from simple pulsatile blood vessels or tubular peristaltic hearts independently many times in different animals • for this reason we find substantial differences in the structure and function of hearts among animals and the difference between a heart and a contractile blood vessel can be cloudy textbook Fig 8.6 pg 355 HEARTS ARTHROPOD HEARTS • Hemolymph is the circulating fluid of open circulatory systems • When hemolymph enters sinuses, it mixes with other fluids like EXTRACELLULAR FLUID and LYMPH (fluid with only small molecules and few proteins) • because these fluids are constantly mixing it is difficult to distinguish each fluid and it is instead referred to as HEMOLYMPH HEARTS ARTHROPOD HEARTS • arthropod hearts generally pump HEMOLYMPH out into the circulation via arteries and blood returns to the heart via a series of holes or OSTIA. • valves within the ostia open and close to regulate the flow of hemolymph • neurons of the CARDIAC GANGLION send a signal to close the ostia and initiate contraction, squeezing blood out of the heart and around the body • arthropods hearts are attached to SUSPENSORY LIGAMENTS, so that after the heart contracts these ligaments pull the heart to increase volume • as this occurs, the ostia open and hemolymph is drawn into the heart, ready for the next contraction (i.e. contraction) (i.e. relaxation) textbook Fig 8.16 pg 368 HEARTS STRUCTURE OF VERTEBRATE HEARTS: FISH • Bony Fish hearts contain four-chambers arranged in a series: blood enter the SINUS VENOSUS, flows into the ATRIUM, then into the muscular VENTRICLE and finally through the BULBOUS ARTERIOSIS • all components except the BULBOUS ARTERIOSIS are contractile in bony fish textbook Fig 8.18 pg 370 HEARTS STRUCTURE OF VERTEBRATE HEARTS: AMPHIBIANS • amphibians have a three-chambered heart: two atria and one ventricle • the ventricle pumps blood through the CONUS ARTERIOSIS to both the PULMONARY (lung) and SYSTEMIC (whole-body) circuit • oxygenated blood returns to the left atrium, while deoxygenated blood flows into the right atrium • the two atria supply blood to the single ventricle, which is either pumped ot the lungs or the rest of the body • mechanisms separating oxygenated and deoxygenated are not fully understood, but involves re-direction using a SPIRAL FOLD textbook Fig 8.18 pg 370 HEARTS STRUCTURE OF VERTEBRATE HEARTS: MOST REPTILES • non-crocodilian reptiles have five-chambered hearts • two atria as in amphibians, but the ventricle is divided into three interconnected compartments: CAVUM VENOSUM, CAVUM PULMONALE, CAVUM ARTERIOSUM • despite incompletely separated ventricle, oxygenated and de-oxygenated blood are typically separated. • de-oxygenated blood enters right atrium and flow to cavum venosum, then across the muscular ridge to the cavum pulmonale • oxygenated blood enters right atrium, then into cavum areteriosum and out aortas to rest of body textbook Fig 8.19 pg 371 HEARTS STRUCTURE OF VERTEBRATE HEARTS • reptiles can also distribute blood selectively between pulmonary and systemic circuits-called a SHUNT. • in a shunt, some fraction of oxygenated poor blood bypasses the pulmonary circuit and re-enters the systemic circuit (and vice-versa too) • reptiles are intermittent breathers often holding their breath for long periods of time and allows for blood to be directed to the body diving rather than to the non-functional lungs HEARTS STRUCTURE OF VERTEBRATE HEARTS: MOST REPTILES • in crocodiles, the ventricles are separated and shunting occurs with the help of a special valve a the entrance of the pulmonary artery (going to lungs) • rather than opening or closing in response to changes in pressure like most cardiac valves, this valve in controlled by the hormone EPINEPHRINE • when crocodiles are at rest underwater, epinephrine is low and the valve is closed diverted blood away from the pulmonary circuit • when active, epinephrine is higher and the valve is open • crocodiles can remain submerged for several hours HEARTS STRUCTURE OF VERTEBRATE HEARTS: BIRDS & MAMMALS • birds and mammals have four chambered hearts • the left side of the heart consists of a thin-walled atrium and a thick walled ventricle, on the right side the ventricle is much thinner textbook Fig 8.20 pg 373 • left ventricle pumps blood through the high resistance systemic circuit and must therefore contract more forcefully than the right ventricle • ATRIOVENTRICULAR VALVES allow blood to flow from artium to ventricle, but not in reverse. • SEMILUNAR VALVES on blood vessels prevent blood from flowing back into ventricles HEARTS THE CARDIAC CYCLE • the vertebrate heart acts as a integrated organ, with each chamber contracting at appropriate times to properly move blood FISH • In fish, each chamber contracts in series • Starts in SINUS VENOSUS-too thin walled to be involved in moving blood long distances, but instead important in initiating contraction • pressure in sinus venosus opens valve and blood flows to atrium • building pressure opens valve from atrium and blood flows into the ventricle • contraction of the ventricle propels blood through the body • the bulbus arteriosis is stretchy and helps dampen blood pressure changes HEARTS THE CARDIAC CYCLE • the vertebrate heart acts as a integrated organ, with each chamber contracting at appropriate times to properly move blood MAMMALS • in mammals, blood entering atria first flows passively into the ventricles as the AV valves are open. • the atria then contract pumping additional blood into the ventricles, reaching END DIASTOLIC VOLUME (max amount of blood in ventricle) • as pressure builds in the ventricles, the semilunar valves are forced open and blood flows out into the arteries in the VENTRICULAR EJECTION PHASE • at this point ventricles have reached minimum or END-SYSTOLIC VOLUME • throughout ventricular contraction, the atria are relaxed so blood is once again passively entering these chambers and the cycle repeats. CARDIAC CYCLE OF MAMMALS Textbook Fig 8.21 pg 374 HEARTS THE CARDIAC CYCLE-MAMMALS • remember that although the two ventricles of the mammalian heart contract simultaneously, the left ventricle contracts much more forcefully than the right ventricle… WHY? textbook Fig 8.20 pg 373 HEARTS CONTROL OF CONTRACTION • clearly, cardiac contraction must be tightly regulated • unlike the muscles described previously that required neural stimulus (i.e. NEUROGENIC), vertebrate CARDIOMYOCYTES (i.e. heart muscle cells) are MYOGENIC, they produce rhythmic spontaneous depolarizations that initiate contraction • cardiomyocytes are electrically coupled by __________________, so that depolarizations can spread from once cell to another. Rate of spontaneous depolarization varies (i.e. some faster, some slow), but those with the fastest rates are called PACEMAKER CELLS In vertebrates (except fish), the pacemakers cells are located in the SINOATRIAL (SA) NODE PACEMAKER CELLS • • • • HEARTS pacemaker cells are small, have few myofibrils, and thus do not contract these cells have unstable resting membrane potentials slowly drift up from -60mV until an action potential is initiated The result of a FUNNY CURRENT (names for unusual behavior) • FUNNY CHANNELS open during textbook Fig 8.23 pg 376 hyperpolarization (i.e. after action potential) and allow Na+ to gradually enter cell. • when membrane potential hits threshold, voltage gated calcium channels open that add to depolarization and trigger an action potential • during repolarization, K+ channels open (more slowly than muscle cells though). • processes is repeated during hyperoplarization HEARTS ENDOCRINE SIGNALS REGULATE PACEMAKER CELLS • endocrine signals alter the rate of pacemaker potentials from the SA node • NOREPINEPRHINE and EPINEPHRINE bind to beta-adrenergic receptors on pacemaker cells • this binding alters the activity of the funny channels and calcium channels, in effect speeding the rate of Na+ depolarization and increases the frequency of action potentials and ultimately increases heart rate Pathway to increase heart rate textbook Fig 8.24 pg 377 HEARTS ENDOCRINE SIGNALS REGULATE PACEMAKER CELLS • ACETYLCHOLINE acts to reduce heart rate • binding of acetylcholine to MUSCARINIC RECEPTORS on pacemaker cells increases permeability of K+ and causes increased hyperpolarization • hyperpolarization means longer for Na+ and calcium to reach threshold • slows the rate of action potential generation and thus heart rate textbook Fig 8.25 pg 378 HEARTS ELECTRICAL ACTIVITY • in addition to electrical signals in the heart are spread via special conduction pathways • gap junctions spread signal within a chamber, but contraction at different chambers at different times relies on alternative pathway • after an action potential in the AV node, the depolarization spreads along an INTERNODAL PATHWAY to the ATRIOVENTRICULAR (AV) node • The AV node delays the signal, then passes on through the BUNDLE OF HIS and PURKINJE FIBERS to the ventricles • delay ensures that atria finish contracting before ventricles start contraction textbook Fig 8.27 pg 380 HEARTS ELECTRICAL ACTIVITY • cardiac muscle cell depolarization produces a strong electrical signal that can be detected by an ELECTROCARDIOGRAM • the fluctuations represent the combined effects of all action potentials • The P-WAVE is the atria depolarizing • The QRS-COMPLEX is the result of ventricle depolarization and atrial repolarization • VENTRICULAR FIBRILLATION results form uncoordinated contraction and results in effective pumping of blood to cells • A DEFRIBRILLATOR machine rapidly depolarizes all the cells, reseting the system and allowing the pacemaker cells to take over LECTURE SUMMARY • structure of vertebrate and invertebrate hearts (i.e. # of chambers) • importance of SHUNTING in reptiles and the ability to divert blood away from the pulmonary circuit when holding breath • described the CARDIAC CYCLE in fish and mammals • control of heart contraction • FUNNY CHANNELS and FUNNY CURRENTS • Role of endocrine signals like acetylcholine, epinephrine and norepinephrine • the spread of electrical signal across the heart and the role of the INTERNODAL PATHWAY • ended by describing the causes of the fluctuations on electrocardiogram NEXT LECTURE BLOOD