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Circulation and gas exchange • Overview • Circulation • Gas exchange Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Overview • Overview: Trading with the Environment • Every organism must exchange materials with its environment – And this exchange ultimately occurs at the cellular level Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In unicellular organisms – These exchanges occur directly with the environment • For most of the cells making up multicellular organisms – Direct exchange with the environment is not possible Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The feathery gills projecting from a salmon – Are an example of a specialized exchange system found in animals Figure 42.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Circulation • Overview • Circulatory systems – Open and closed – Circulation in invertebrates – Circulation in vertebrates • Mammalian circulation system – The Pathway – The Pumping – The Movement – The Exchange Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Circulation • Diffusion alone is not adequate for transporting substances over long distances in animals – Time for a substance to diffuse is proportional to the square of the distance • Transport systems – Functionally connect the organs of exchange with the body cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Most complex animals have internal transport systems – That circulate fluid, providing a lifeline between the aqueous environment of living cells and the exchange organs, such as lungs, that exchange chemicals with the outside environment Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gastrovascular Cavities • Simple animals, such as cnidarians – Have a body wall only two cells thick that encloses a gastrovascular cavity • The gastrovascular cavity – Functions in both digestion and distribution of substances throughout the body Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Some cnidarians, such as jellies – Have elaborate gastrovascular cavities Circular canal Mouth Radial canal 5 cm Figure 42.2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Open and Closed Circulatory Systems • More complex animals – Have one of two types of circulatory systems: open or closed • Both of these types of systems have three basic components – A circulatory fluid (blood) – A set of tubes (blood vessels) – A muscular pump (the heart) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In insects, other arthropods, and most molluscs – Blood bathes the organs directly in an open circulatory system Heart Hemolymph in sinuses surrounding ograns Anterior vessel Figure 42.3a Lateral vessels Ostia Tubular heart (a) An open circulatory system Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In a closed circulatory system – Blood is confined to vessels and is distinct from the interstitial fluid Heart Interstitial fluid Small branch vessels in each organ Dorsal vessel (main heart) Auxiliary hearts Figure 42.3b Ventral vessels (b) A closed circulatory system Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Open systems – Less energetically costly – Can function as a hydrostatic skeleton • Closed systems – Are more efficient at transporting circulatory fluids to tissues and cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Survey of Vertebrate Circulation • Humans and other vertebrates have a closed circulatory system – Often called the cardiovascular system • Blood flows in a closed cardiovascular system – Consisting of blood vessels and a two- to fourchambered heart Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Arteries carry blood to capillaries – The sites of chemical exchange between the blood and interstitial fluid • Veins – Return blood from capillaries to the heart Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Vertebrate circulatory systems AMPHIBIANS REPTILES (EXCEPT BIRDS) MAMMALS AND BIRDS Lung and skin capillaries Lung capillaries Lung capillaries FISHES Gill capillaries Artery Pulmocutaneous circuit Gill circulation Heart: ventricle (V) A Atrium (A) Systemic Vein circulation Systemic capillaries Right systemic aorta Pulmonary circuit A A V Right V Left Right Systemic circuit Systemic capillaries Figure 42.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Pulmonary circuit Left Systemic V aorta Left A Systemic capillaries A V Right A V Left Systemic circuit Systemic capillaries Fishes • A fish heart has two main chambers – One ventricle and one atrium • Blood pumped from the ventricle – Travels to the gills, where it picks up O2 and disposes of CO2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Amphibians • Frogs and other amphibians – Have a three-chambered heart, with two atria and one ventricle • The ventricle pumps blood into a forked artery – That splits the ventricle’s output into the pulmocutaneous circuit and the systemic circuit Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Reptiles (Except Birds) • Reptiles have double circulation – With a pulmonary circuit (lungs) and a systemic circuit • Turtles, snakes, and lizards – Have a three-chambered heart Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mammals and Birds • In all mammals and birds – The ventricle is completely divided into separate right and left chambers • The left side of the heart pumps and receives only oxygen-rich blood – While the right side receives and pumps only oxygen-poor blood Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • A powerful four-chambered heart – Was an essential adaptation of the endothermic way of life characteristic of mammals and birds Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mammalian Circulation: The Pathway • The mammalian cardiovascular system 7 Capillaries of head and forelimbs Anterior vena cava Pulmonary artery Aorta Pulmonary artery 9 6 Capillaries of right lung Capillaries of left lung 2 4 3 Pulmonary vein 5 1 Right atrium 3 11 Left atrium Pulmonary vein 10 Left ventricle Right ventricle Aorta Posterior vena cava 8 Figure 42.5 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Capillaries of abdominal organs and hind limbs • Heart valves – Dictate a one-way flow of blood through the heart • Blood begins its flow – With the right ventricle pumping blood to the lungs • In the lungs – The blood loads O2 and unloads CO2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Oxygen-rich blood from the lungs – Enters the heart at the left atrium and is pumped to the body tissues by the left ventricle • Blood returns to the heart – Through the right atrium Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mammalian Circulation: The Pumping • The heart contracts and relaxes – In a rhythmic cycle called the cardiac cycle • The contraction, or pumping, phase of the cycle – Is called systole • The relaxation, or filling, phase of the cycle – Is called diastole Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The heart rate, also called the pulse – Is the number of beats per minute • The cardiac output – Is the volume of blood pumped into the systemic circulation per minute • It depends on the heart rate and stroke volume Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Some cardiac muscle cells are self-excitable – Meaning they contract without any signal from the nervous system • A region of the heart called the sinoatrial (SA) node, or pacemaker – Sets the rate and timing at which all cardiac muscle cells contract Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The SA node generates electrical impulses that spread rapidly through the wall of the atria • Purkinje fibers conduct the signals to the apex of the heart and throughout ventricular walls • This stimulates the ventricles to contract from the apex toward the atria, driving blood to the arteries Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The pacemaker is influenced by – Nerves, hormones, body temperature, and exercise Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mammalian Circulation: The Movement • Physical principles govern blood circulation • The same physical principles that govern the movement of water in plumbing systems – Also influence the functioning of animal circulatory systems Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The “infrastructure” of the circulatory system – Is its network of blood vessels Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • All blood vessels – Are built of similar tissues – Have three similar layers Artery Vein Basement membrane Endothelium 100 µm Valve Endothelium Smooth muscle Connective tissue Endothelium Capillary Smooth muscle Connective tissue Artery Vein Venule Figure 42.9 Arteriole Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Structural differences in arteries, veins, and capillaries – Correlate with their different functions • Capillaries have very thin walls – To enhance exchange • Arteries have thicker walls – To accommodate the high pressure of blood pumped from the heart Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In the thinner-walled veins – Blood flows back to the heart mainly as a result of muscle action Direction of blood flow in vein (toward heart) Valve (open) Skeletal muscle Valve (closed) Figure 42.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Blood Flow Velocity • Physical laws governing the movement of fluids through pipes – Influence blood flow and blood pressure • Blood travels over a thousand times faster in the aorta than in capillaries • Law of continuity – Fluid flows through narrower segments of the pipe faster than it flows through wider segments Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The velocity of blood flow varies in the circulatory system Systolic pressure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Veins Venules Venae cavae Figure 42.11 Arterioles Diastolic pressure Capillaries 120 100 80 60 40 20 0 Arteries Velocity (cm/sec) 50 40 30 20 10 0 Aorta Area (cm2) 5,000 4,000 3,000 2,000 1,000 0 Pressure (mm Hg) – And is slowest in the capillary beds as a result of the large total cross-sectional area Blood Pressure • Fluids exert a force called hydrostatic pressure – Fluids flow from areas of high to low pressure • Blood pressure – Is the hydrostatic pressure that blood exerts against the wall of a vessel; it is much greater in the arteries when the heart contracts during systole Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Surge of pressure felt during pulse is due to narrow openings of arterioles impeding the exit of blood from arteries, the peripheral resistance Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Blood pressure is determined partly by cardiac output – And partly by peripheral resistance due to variable constriction of the arterioles • Nerve impulses, hormones and other signals control the arteriole wall muscles – Stress, both physical and emotional can raise blood pressure by triggering nervous and hormonal responses that will constrict blood vessels Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Cardiac output is adjusted in concert with changes in peripheral resistance – During heavy exercise, arterioles in working muscles dilate admitting greater flow of oxygen and decreasing peripheral resistance – At the same time cardiac output increases Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Capillary Function • Capillaries in major organs are usually filled to capacity – But in many other sites, the blood supply varies Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Two mechanisms – Regulate the distribution of blood in capillary beds • In one mechanism – Contraction of the smooth muscle layer in the wall of an arteriole constricts the vessel Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In a second mechanism – Precapillary sphincters control the flow of blood between arterioles and venules Precapillary sphincters Thoroughfare channel (a) Sphincters relaxed Arteriole Venule Capillaries Arteriole Venule (b) Sphincters contracted (c) Capillaries and larger vessels (SEM) Figure 42.13 a–c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 20 m Mammalian Circulation: The Exchange • The critical exchange of substances between the blood and interstitial fluid – Takes place across the thin endothelial walls of the capillaries Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The difference between blood pressure and osmotic pressure – Drives fluids out of capillaries at the arteriole end and into capillaries at the venule end Tissue cell Capillary Capillary Red blood cell INTERSTITIAL FLUID Net fluid movement out Net fluid movement in 15 m Direction of blood flow Blood pressure Osmotic pressure Inward flow Pressure At the arterial end of a capillary, blood pressure is greater than osmotic pressure, and fluid flows out of the capillary into the interstitial fluid. At the venule end of a capillary, blood pressure is less than osmotic pressure, and fluid flows from the interstitial fluid into the capillary. Outward flow Figure 42.14 Arterial end of capillary Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Venule end Fluid Return by the Lymphatic System • The lymphatic system – Returns fluid to the body from the capillary beds – Aids in body defense Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Fluid reenters the circulation – Directly at the venous end of the capillary bed and indirectly through the lymphatic system Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gas exchange • Respiratory surfaces • Respiratory adaptations for aquatic animals • Respiratory adaptations for terrestrial animals • Breathing across vertebrates • Diffusion of gases Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Gas exchange occurs across specialized respiratory surfaces • Gas exchange – Supplies oxygen for cellular respiration and disposes of carbon dioxide Respiratory medium (air of water) O2 CO2 Respiratory surface Organismal level Circulatory system Cellular level Energy-rich molecules from food Cellular respiration Figure 42.19 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ATP • Animals require large, moist respiratory surfaces for the adequate diffusion of respiratory gases – Between their cells and the respiratory medium, either air or water Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • For simple animals, plasma membrane of every cell is close enough to outside environment for gases to diffuse in and out • Most animals, the bulk of the body lacks direct access to the respiratory medium – Some animals like earthworms and amphibians use entire outer skin as a respiratory organ, how? Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gills in Aquatic Animals • Gills are outfoldings of the body surface – Specialized for gas exchange Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In some invertebrates – The gills have a simple shape and are distributed over much of the body (a) Sea star. The gills of a sea star are simple tubular projections of the skin. The hollow core of each gill is an extension of the coelom (body cavity). Gas exchange occurs by diffusion across the gill surfaces, and fluid in the coelom circulates in and out of the gills, aiding gas transport. The surfaces of a sea star’s tube feet also function in gas exchange. Gills Coelom Figure 42.20a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Tube foot • Many segmented worms have flaplike gills – That extend from each segment of their body (b) Marine worm. Many polychaetes (marine worms of the phylum Annelida) have a pair of flattened appendages called parapodia on each body segment. The parapodia serve as gills and also function in crawling and swimming. Parapodia Figure 42.20b Gill Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The gills of clams, crayfish, and many other animals – Are restricted to a local body region (c) Scallop. The gills of a scallop are long, flattened plates that project from the main body mass inside the hard shell. Cilia on the gills circulate water around the gill surfaces. (d) Crayfish. Crayfish and other crustaceans have long, feathery gills covered by the exoskeleton. Specialized body appendages drive water over the gill surfaces. Gills Gills Figure 42.20c, d Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The effectiveness of gas exchange in some gills, including those of fishes – Is increased by ventilation and countercurrent flow of blood and water Oxygen-poor blood Gill arch Gill arch Water flow Blood vessel Oxygen-rich blood Lamella Operculum O2 Figure 42.21 Water flow over lamellae showing % O2 Gill filaments Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Blood flow through capillaries in lamellae showing % O2 Countercurrent exchange Tracheal Systems in Insects • The tracheal system of insects – Consists of tiny branching tubes that penetrate the body Air sacs Tracheae Spiracle (a) The respiratory system of an insect consists of branched internal tubes that deliver air directly to body cells. Rings of chitin reinforce the largest tubes, called tracheae, keeping them from collapsing. Enlarged portions of tracheae form air sacs near organs that require a large supply of oxygen. Air enters the tracheae through openings called spiracles on the insect’s body surface and passes into smaller tubes called tracheoles. The tracheoles are closed and contain fluid (blue-gray). When the animal is active and is using more O2, most of the fluid is withdrawn into the body. This increases the surface area of air in contact with cells. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 42.22a • The tracheal tubes – Supply O2 directly to body cells Body cell Air sac Tracheole Trachea Air Tracheoles Mitochondria Body wall Myofibrils (b) This micrograph shows cross sections of tracheoles in a tiny piece of insect flight muscle (TEM). Each of the numerous mitochondria in the muscle cells lies within about 5 µm of a tracheole. Figure 42.22b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2.5 µm Lungs • Spiders, land snails, and most terrestrial vertebrates – Have internal lungs Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mammalian Respiratory Systems: A Closer Look • A system of branching ducts – Conveys air to the lungs Branch from the pulmonary artery (oxygen-poor blood) Branch from the pulmonary vein (oxygen-rich blood) Terminal bronchiole Nasal cavity Pharynx Left lung Alveoli 50 µm 50 µm Larynx Esophagus Trachea Right lung Bronchus Bronchiole Diaphragm SEM Heart Figure 42.23 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Colorized SEM • Breathing ventilates the lungs • The process that ventilates the lungs is breathing – The alternate inhalation and exhalation of air Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings How an Amphibian Breathes • An amphibian such as a frog – Ventilates its lungs by positive pressure breathing, which forces air down the trachea Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings How a Mammal Breathes • Mammals ventilate their lungs – By negative pressure breathing, which pulls air into the lungs Rib cage expands as rib muscles contract Air inhaled Rib cage gets smaller as rib muscles relax Air exhaled Lung Diaphragm INHALATION Diaphragm contracts (moves down) Figure 42.24 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings EXHALATION Diaphragm relaxes (moves up) • Lung volume increases – As the rib muscles and diaphragm contract Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings How a Bird Breathes • Besides lungs, bird have eight or nine air sacs – That function as bellows that keep air flowing through the lungs Air Air Anterior air sacs Trachea Posterior air sacs Lungs Lungs Air tubes (parabronchi) in lung EXHALATION Air sacs empty; lungs fill INHALATION Air sacs fill Figure 42.25 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 1 mm • Air passes through the lungs – In one direction only • Every exhalation – Completely renews the air in the lungs Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Role of Partial Pressure Gradients • Gases diffuse down pressure gradients – In the lungs and other organs • Diffusion of a gas – Depends on differences in a quantity called partial pressure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • A gas always diffuses from a region of higher partial pressure – To a region of lower partial pressure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In the lungs and in the tissues – O2 and CO2 diffuse from where their partial pressures are higher to where they are lower Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Inhaled air Exhaled air 160 0.2 O2 CO2 120 27 Alveolar spaces O2 CO2 104 Alveolar epithelial cells 40 O2 CO2 Blood entering alveolar capillaries 40 O2 CO2 2 1 O2 Alveolar capillaries of lung 45 O2 CO2 104 Pulmonary veins Systemic arteries Systemic veins CO2 40 40 O2 CO2 Pulmonary arteries Blood leaving tissue capillaries Blood leaving alveolar capillaries Heart Tissue capillaries O2 3 4 45 O2 CO2 Tissue cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 100 40 O2 CO2 O2 CO2 Figure 42.27 Blood entering tissue capillaries <40 >45 O2 CO2