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CHAPTER 49 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Gas Exchange • One of the major physiological challenges facing all multicellular animals is obtaining sufficient oxygen and disposing of excess carbon dioxide • In vertebrates, the gases diffuse into the aqueous layer covering the epithelial cells that line the respiratory organs • Diffusion is passive, driven only by the difference in O2 and CO2 concentrations on the two sides of the membranes and their relative solubilities in the plasma membrane 2 Gas Exchange • Gases diffuse directly into unicellular organisms • However, most multicellular animals require system adaptations to enhance gas exchange • Amphibians respire across their skin • Insects have an extensive tracheal system • Fish use gills • Mammals have a large network of alveoli 3 4 5 6 Gills • Specialized extensions of tissue that project into water • Increase surface area for diffusion • External gills are not enclosed within body structures – Found in fish and amphibians – Two main disadvantages • Must be constantly moved to ensure contact with oxygen-rich fresh water • Are easily damaged 7 Gills • Gills of bony fishes are located between the oral (buccal or mouth) cavity and the opercular cavities • These two sets of cavities function as pumps that alternately expand • Move water into the mouth, through the gills, and out of the fish through the open operculum or gill cover 8 Gills 9 Gills • Some bony fish have immobile opercula – Swim constantly to force water over gills – Ram ventilation • Most bony fish have flexible gill covers • Remora switch between ram ventilation and pumping action 10 Gills • 3–7 gill arches on each side of a fish’s head • Each is composed of two rows of gill filaments • Each gill filament consist of lamellae – Thin membranous plates that project into water flow – Water flows past lamellae in 1 direction only 11 12 Gills • Within each lamella, blood flows opposite to direction of water movement – Countercurrent flow – Maximizes oxygenation of blood – Increases Dp • Fish gills are the most efficient of all respiratory organs 13 14 Gills • Many amphibians use cutaneous respiration for gas exchange • In terrestrial arthropods, the respiratory system consists of air ducts called trachea, which branch into very small tracheoles – Tracheoles are in direct contact with individual cells – Spiracles (openings in the exoskeleton) can 15 be opened or closed by valves Lungs • Gills were replaced in terrestrial animals because – Air is less supportive than water – Water evaporates • The lung minimizes evaporation by moving air through a branched tubular passage • A two-way flow system – Except birds 16 Lungs • Air exerts a pressure downward, due to gravity • A pressure of 760 mm Hg is defined as one atmosphere (1.0 atm) of pressure • Partial pressure is the pressure contributed by a gas to the total atmospheric pressure 17 Lungs • Lungs of amphibians are formed as saclike outpouchings of the gut • Frogs have positive pressure breathing – Force air into their lungs by creating a positive pressure in the buccal cavity • Reptiles have negative pressure breathing – Expand rib cages by muscular contractions, creating lower pressure inside the lungs 18 19 Lungs • Lungs of mammals are packed with millions of alveoli (sites of gas exchange) • Inhaled air passes through the larynx, glottis, and trachea • Bifurcates into the right and left bronchi, which enter each lung and further subdivide into bronchioles • Alveoli are surrounded by an extensive capillary network 20 Lungs 21 Lungs • Respiration in birds occurs in two cycles – Cycle 1 = Inhaled air is drawn from the trachea into posterior air sacs, and exhaled into the lungs – Cycle 2 = Air is drawn from the lungs into anterior air sacs, and exhaled through the trachea • Blood flow runs 90o to the air flow – Crosscurrent flow – Not as efficient as countercurrent flow 22 Lungs 23 Gas Exchange • Gas exchange is driven by differences in partial pressures • Blood returning from the systemic circulation, depleted in oxygen, has a partial oxygen pressure (PO2) of about 40 mm Hg • By contrast, the PO2 in the alveoli is about 105 mm Hg • The blood leaving the lungs, as a result of this gas exchange, normally contains a PO2 of about 100 mm 24 25 Lung Structure and Function • Outside of each lung is covered by the visceral pleural membrane • Inner wall of the thoracic cavity is lined by the parietal pleural membrane • Space between the two membranes is called the pleural cavity – Normally very small and filled with fluid – Causes 2 membranes to adhere – Lungs move with thoracic cavity 26 Lung Structure and Function • During inhalation, thoracic volume increases through contraction of two muscle sets – Contraction of the external intercostal muscles expands the rib cage – Contraction of the diaphragm expands the volume of thorax and lungs • Produces negative pressure which draws air into the lungs 27 28 Lung Structure and Function • Tidal volume – Volume of air moving in and out of lungs in a person at rest • Vital capacity – Maximum amount of air that can be expired after a forceful inspiration • Hypoventilation – Insufficient breathing – Blood has abnormally high PCO2 • Hyperventilation – Excessive breathing – Blood has abnormally low PCO2 29 Lung Structure and Function • Each breath is initiated by neurons in a respiratory control center in the medulla oblongata • Stimulate external intercostal muscles and diaphragm to contract, causing inhalation • When neurons stop producing impulses, respiratory muscles relax, and exhalation occurs • Muscles of breathing usually controlled automatically – Can be voluntarily overridden – hold your breath 30 Lung Structure and Function • Neurons are sensitive to blood PCO2 changes • A rise in PCO2 causes increased production of carbonic acid (H2CO3), lowering the blood pH • Stimulates chemosensitive neurons in the aortic and carotid bodies • Send impulses to respiratory control center to increase rate of breathing • Brain also contains central chemoreceptors that are sensitive to changes in the pH of cerebrospinal fluid (CSF) 31 32 Respiratory Diseases • Chronic obstructive pulmonary disease (COPD) – Refers to any disorder that obstructs airflow on a long-term basis – Asthma • Allergen triggers the release of histamine, causing intense constriction of the bronchi and sometimes suffocation 33 Respiratory Diseases • Chronic obstructive pulmonary disease (COPD) (cont.) – Emphysema • Alveolar walls break down and the lung exhibits larger but fewer alveoli • Lungs become less elastic • People with emphysema become exhausted because they expend three to four times the normal amount of energy just to breathe • Eighty to 90% of emphysema deaths are caused by cigarette smoking 34 Respiratory Diseases • Lung cancer accounts for more deaths than any other form of cancer • Caused mainly by cigarette smoking • Follows or accompanies COPD • Lung cancer metastasizes (spreads) so rapidly that it has usually invaded other organs by the time it is diagnosed • Chance of recovery from metastasized lung cancer is poor, with only 3% of patients surviving for 5 years after diagnosis 35 36 Hemoglobin • Consists of four polypeptide chains: two a and two b • Each chain is associated with a heme group • Each heme group has a central iron atom that can bind a molecule of O2 • Hemoglobin loads up with oxygen in the lungs, forming oxyhemoglobin • Some molecules lose O2 as blood passes through capillaries, forming deoxyhemoglobin 37 Hemoglobin • Hemoglobin’s affinity for O2 is affected by pH and temperature • The pH effect is known as the Bohr shift – Increased CO2 in blood increases H+ – Lower pH reduces hemoglobin’s affinity for O2 – Results in a shift of oxyhemoglobin dissociation curve to the right – Facilitates oxygen unloading • Increasing temperature has a similar effect38 Hemoglobin 39 Transportation of Carbon Dioxide • About 8% of the CO2 in blood is dissolved in plasma • 20% of the CO2 in blood is bound to hemoglobin • Remaining 72% diffuses into red blood cells – Enzyme carbonic anhydrase combines CO2 with H2O to form H2CO3 – H2CO3 dissociates into H+ and HCO3– – H+ binds to deoxyhemoglobin – HCO3– moves out of the blood and into plasma – One Cl– exchanged for one HCO3– – “chloride shift” 40 41 Transportation of Carbon Dioxide • When the blood passes through pulmonary capillaries, these reactions are reversed • The result is the production of CO2 gas, which is exhaled • Other dissolved gases are also transported by hemoglobin – Nitric oxide (NO) and carbon monoxide (CO) 42