AIIMS, New Delhi. Dr. K. K. Deepak, Prof. & HOD, Physiology AIIMS, New Delhi Dr. Geetanjali Bade, Asst. Professor AIIMS, New Delhi

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1 Course : PG Pathshala-Biophysics Paper 13 : Physiological Biophysics Module 17 : Gas transport and pulmonary circulation Principal Investigator: Co-Principal Investigator: Paper Coordinator: Content Writer: Content Reviewer: Dr. Moganty R. Rajeswari, Professor AIIMS, New Delhi Dr. T. P. Singh, Prof., AIIMS, New Delhi Dr. K. K. Deepak, Prof. & HOD, Physiology AIIMS, New Delhi Dr. Geetanjali Bade, Asst. Professor AIIMS, New Delhi Dr. Renuka Sharma, Professor VMMC & SJH, New Delhi Objectives: Describe physiologic anatomy of pulmonary circulatory system Describe characteristics of pulmonary circulation Describe regional variation of circulation and ventilation-perfusion ratio Compare the composition of alveolar, inspired and expired air List the methods of transport of oxygen in blood Draw and explain O 2-Hb dissociation curve List the ways in which CO 2 is transported in blood Classify types of hypoxia with examples 1. Physiologic anatomy of pulmonary circulatory system The lung has two types of circulations A high-pressure, low-flow circulation supplies systemic arterial blood to the trachea, the bronchial tree including the terminal bronchioles, the supporting tissues of the lung, including vasculature. A low-pressure, high-flow circulation that supplies venous blood from all parts of the body to the alveolar capillaries where oxygen is added and carbon dioxide is removed. The pulmonary artery, which receives blood from the right ventricle, and its arterial branches carry blood to the alveolar capillaries for gas exchange and the pulmonary veins then return the blood to the left atrium to be pumped by left ventricle into systemic circulation. 1.2 Characteristics of pulmonary circulatory system The characteristics of pulmonary artery and its branches are 1. It is thin with a wall thickness one-third that of the aorta. 2. Its branches are very short, with larger diameters

2 3. Their walls are thin and distensible Because of all these characteristics, pulmonary arterial circulation has a large compliance and can accommodate large volume of blood pressure without much rise in pressure. Characteristics of Pulmonary veins They are very short and empty into right atrium. Pressures in the Pulmonary System Pressures in the right ventricle The systolic pressure in the right ventricle is about 25 mm Hg and the diastolic pressure averages about 0 to 1 mm Hg, values that are only one-fifth those for the left ventricle. Pressures in the pulmonary circulation The systolic pressure is 25 mm Hg and diastolic pressure is 8 mm Hg while mean arterial pressure is 15 mm Hg. The mean pulmonary capillary pressure is about 10 mm Hg whereas oncotic pressure is 25 mm Hg. Hence, the inward directed pressure gradient of 15 mm Hg will keep the alveoli free of fluid. Pressures in left Atrium and Pulmonary Veins The mean pressure in the left atrium and the major pulmonary veins averages about 2 mm Hg in the recumbent human being, varying from as low as 1 mm Hg to as high as 5 mm Hg 1.2 Effect of gravity on a. Pulmonary circulation In upright position, the upper portions of lungs are well above the level of heart and bases are below it so upper portions receive less blood supply as compared to basal portions of lungs. b. Pulmonary ventilation In upright position, the intrapleural pressure at apex is less negative as compared to base of the lungs. It causes more distension of alveoli at the apex and less ventilation per minute as compared to base of the lungs. In the upper portions of lungs blood flow is less and alveoli are large and less ventilated. Pressure in the capillaries is close to atmospheric pressure in the alveoli which is just sufficient to maintain perfusion. If capillary pressure reduces or alveolar pressure rises, some of the capillaries collapse and they do not take part in diffusion of gases thus becoming part of the physiological dead space. In the middle and lower portions of lungs, capillary pressure is greater than alveolar pressure which is in turn greater than pulmonary venous pressure, so the veins are collapsed. As the arterial pressure increases towards the base of lungs, pulmonary blood flow also increases. 1.3 Ventilation / perfusion ratio (V/P ratio) It is the ratio of pulmonary ventilation to pulmonary blood flow. V/P ratio for the whole lung at rest is about 0.8. Marked differences are observed in various parts of normal lung due to the effect of gravity. As noted above both ventilation and perfusion decline from base to apex linearly in upright position, but V/P ratio is high at apex of lungs. 2. Gas exchange in the lungs 2.1 Partial pressure of gases The pressure exerted by any one gas in a mixture of gases is called as partial pressure of that gas. It is directly proportional to concentration of gas molecules. The composition of dry air is 20.98% O 2, 0.04% CO 2, 78.06% N 2, and 0.92% other inert constituents such as argon and helium. The barometric pressure (PB) at sea level is 760 mm Hg (1 atmosphere). The partial pressure (indicated by the symbol P) of O 2 in dry air is therefore 0.21 x 760, or 160 mm Hg at sea level. The PN 2 and the other inert gases is 0.79 x 760, or 600 mm Hg; and the PCO 2 is x 760, or 0.3 mm

3 Hg. The water vapor in the air reduces these percentages, and therefore the partial pressures, to a slight degree. Air equilibrated with water is saturated with water vapor, and inspired air is saturated by the time it reaches the lungs. The partial pressure of water at body temperature (37 C) is 47 mm Hg. Therefore, the partial pressures at sea level of the other gases in the air reaching the lungs are PO 2, 149 mm Hg; PCO 2, 0.3 mm Hg; and PN 2 (including the other inert gases), 564 mm Hg. Gas diffuses from areas of high pressure to areas of low pressure, with the rate of diffusion depending on the concentration gradient and the nature of the barrier between the two areas. When a mixture of gases comes in contact with a liquid and equilibrates with it, each gas in the mixture dissolves in the liquid to an extent determined by its partial pressure and its solubility in the fluid. The solubility coefficient of the gas determines the partial pressure exerted by the gas for a given concentration of dissolved gas. It can be expressed as (Henry s law) Partial pressure= concentration of dissolved gas/solubility coefficient At body temperature, solubility coefficient for O 2 is and for CO 2 it is As CO 2 is more than 20 times as soluble as O 2, the partial pressure of CO 2 is less than one twentieth that exerted by O Partial pressures of gases (mm Hg) in various parts of respiratory system and circulatory system Oxygen concentration and partial pressure in alveoli Oxygen is continuously absorbed from alveoli into the blood and new oxygen is continuously breathed into alveoli from atmosphere. So concentration and partial pressure of O 2 at alveoli is determined by 1) rate of absorption of O 2 in to the blood and 2) entry of new O 2 in to the lungs by ventilatory process. Carbon dioxide concentration and partial pressure in alveoli CO 2 is continually produced by the body, carried by the blood to alveoli and then removed from alveoli by ventilation. So alveolar PCO 2 increases in proportion to rate of CO 2 excretion and decreases in inverse proportion to alveolar ventilation. Composition of alveolar air (Figure is taken from textbook. Kindly redraw it) In the steady state, inspired air mixes with the alveolar gas, replacing the O 2 that has entered the blood and diluting the CO 2 that has entered the alveoli. A part of this mixture is expired. The O 2 content of the alveolar gas then falls and its CO 2 content rises until the next inspiration. Because the volume of gas in the alveoli is about 2 L at the end of expiration (functional residual capacity), each 350 ml increment of inspired and expired air has relatively little effect on PO 2 and PCO 2. Indeed, the composition of alveolar gas

4 remains remarkably constant, not only at rest but also under a variety of other conditions. Diffusion of gases across alveolar- capillary membrane (Figure is taken from textbook. Kindly redraw it) 2.3 Gases diffuse across alveolar- capillary membrane. Blood takes 0.75 s to traverse the pulmonary capillaries at rest. The time required for each gas to reach equilibrium with blood depends up on their reaction with substances in blood. For example, the anesthetic gas nitrous oxide (N 2O) does not react and reaches equilibrium in about 0.1 s. In this situation, the amount of N 2O taken up is not limited by diffusion but by the amount of blood flowing through the pulmonary capillaries; that is, it is flow-limited. On the other hand, carbon monoxide (CO) is taken up by hemoglobin in the red blood cells at such a high rate that the partial pressure of CO in the capillaries stays very low and equilibrium is not reached in the 0.75 s the blood is in the pulmonary capillaries. Therefore, the transfer of CO is not limited by perfusion at rest and instead is diffusionlimited. O 2 is intermediate between N 2O and CO; it is taken up by hemoglobin, but much less avidly than CO, and it reaches equilibrium with capillary blood in about 0.3 s. Thus, its uptake is perfusion-limited. Diffusing capacity-volume of gas that will diffuse through the respiratory membrane each minute for a partial pressure difference of 1 mm Hg. Diffusing capacity for CO (DL CO) is taken as the index of diffusing capacity as its uptake is diffusion limited. Normal value-25 ml/min/mm Hg. It increases during exercise, upto three folds the normal. The PO 2 of alveolar air is 100 mm Hg and that of blood entering capillaries is 40 mm Hg. Oxygen diffuses across the respiratory membrane and PO 2 of blood rises to 97 mm Hg. It falls to 95 mm Hg in the aorta. Similarly, PCO 2 of venous blood is 46 mm Hg and that of alveolar air is 40 mm Hg. CO 2 moves from blood to alveoli and PCO 2 of blood leaving the lungs is 40 mm Hg. 3. Transport of oxygen Methods of oxygen transport Oxygen diffuses from alveoli into pulmonary blood and then gets transported to tissue capillaries in two forms 1. Combined with hemoglobin 2. Dissolved in plasma 3.1 Role of hemoglobin in oxygen transport

5 97 % of the O 2 transported from lungs to the tissue is in combination with hemoglobin present in RBCs. Hemoglobin is made up of four subunits. Each subunit contains a heme moiety which is a porphyrin ring complex having one atom of ferrous iron. Thus each of the four iron atoms in Hb can reversibly bind with one O 2 molecule. When PO 2 is high as in pulmonary capillaries, O 2 binds with Hb but when PO 2 is low as in tissue capillaries, O 2 is released from Hb. Oxygen- hemoglobin dissociation curve (Figure is taken from textbook. Kindly redraw it) It relates percentage saturation of the O 2 carrying capacity of hemoglobin to the PO 2. This curve has a characteristic sigmoid shape because combination of the first heme in the Hb molecule with O 2 increases the affinity of the second heme for O 2, and oxygenation of the second increases the affinity of the third, and so on, so that the affinity of Hb for the fourth O 2 molecule is many times that for the first. Maximum amount of oxygen that can combine with hemoglobin When blood is equilibrated with 100% O 2, the normal hemoglobin becomes 100% saturated. When fully saturated, each gram of normal hemoglobin contains 1.39 ml of O 2. However, blood normally contains small amounts of inactive hemoglobin derivatives, and the measured value in vivo is lower, 1.34 ml of O 2. Maximum amount of oxygen released from hemoglobin to tissues. In arterial blood (Hb 97% saturated), O 2 content is 19.4 ml per 100 ml In venous blood (Hb 75% saturated), O 2 content is 14.4 ml per 100 ml Thus 5 ml of O 2 is transported from the lungs to the tissues by 100 ml of blood. Bohr effect The decrease in O2 affinity of Hb when the ph of blood falls is called as Bohr effect Factors that shift the oxygen-hemoglobin curve to right are 1. Increased carbon dioxide concentration 2. Increase temperature 3. Decreased ph of blood 4. Increased 2,3-biphosphoglycerate in RBCs All these conditions lead to release of more oxygen to the tissues. 3.2 Transport of oxygen in dissolved state Only 3% O 2 is transported to tissues in dissolved state. At PO 2 95 mm Hg, 0.29 ml O 2 is dissolved in plasma. At PO 2 40 mm Hg, 0.12 ml O 2 remains dissolved in plasma ml of O 2 is transported in the dissolved state to the tissues by each 100 ml of arterial blood.

6 4. Transport of carbon dioxide Carbon dioxide is transported in blood in following forms 1. In the dissolved state 2. As bicarbonate ions 3. In combination with hemoglobin and plasma proteins 4.1 In the dissolved state The amount of CO 2 dissolved in plasma at 45 mm Hg is 2.7 ml/dl The amount of CO 2 dissolved in plasma at 40 mm Hg is 2.4 ml/dl Thus only 0.3 ml (7% of total) CO 2 is transported by 100 ml of blood in dissolved form. 4.2 As bicarbonate ions The CO 2 that diffuses into red blood cells is rapidly hydrated to H 2CO 3 because of the presence of carbonic anhydrase. The H 2CO 3 dissociates to H + and HCO 3, and the H + is buffered, primarily by hemoglobin, while the HCO 3 enters the plasma. Because the rise in the HCO 3 content of red cells is much greater than that in plasma as the blood passes through the capillaries, about 70% of the HCO 3 formed in the red cells enters the plasma. The excess HCO 3 leaves the red cells in exchange for Cl which is called as chloride shift. This process is mediated by anion exchanger 1, a major membrane protein in the red blood cell. Because of this, the Cl content of the red cells in venous blood is significantly greater than that in arterial blood. This mode of transport accounts for 70% of CO 2 transported from tissues to the lung. Haldane effect Combination of oxygen with hemoglobin in the lungs causes hemoglobin to become a stronger acid and it displaces CO 2 from blood into alveoli. 4.3 In combination with hemoglobin and plasma proteins Carbon dioxide reacts directly with plasma proteins and amine radicals of Hb molecule to form carbaminohemoglobin. This is a reversible reaction and accounts for 30% of CO 2 transported by blood. 5. Hypoxia Hypoxia is deficiency of O 2 at issue level. It could be due to inadequate supply of O 2 to the tissues or inability of the tissues to utilize available O 2. Types of hypoxia are 1. Hypoxic hypoxia 2. Anemic hypoxia 3. Stagnant hypoxia 4. Histotoxic hypoxia 5.1 Hypoxic hypoxia When PO 2 of arterial blood is reduced, it is called as hypoxic hypoxia. The causes for hypoxic hypoxia are low PO 2 in inspired air, hypoventilation, and diffusion defect at lungs or ventilation-perfusion mismatch. A common example is difficulty in breathing at high altitudes. For hypoxic hypoxia, oxygen therapy is the treatment of choice.

7 5.2 Anemic hypoxia In this hypoxia, PO 2 of arterial blood is normal but Hb to carry O 2 is inadequate. Conditions that lead to anemic hypoxia are anemia, carbon monoxide poisoning and presence of altered Hb like methemoglobin. Treatment for anemic hypoxia is oxygen therapy which increases dissolved O 2 and for carbon monoxide poisoning, hyperbaric oxygen therapy is required as affinity of CO for Hb is 210 times greater than the affinity of O Stagnant hypoxia Stagnant hypoxia occurs due to decreased blood flow to the tissues. It is seen in heart failure, shock and vascular obstruction. O 2 therapy is not much useful in this type of hypoxia, underlying cause needs to be treated. 5.4 Histotoxic hypoxia In this hypoxia, tissue cannot utilize oxygen despite of normal O 2 supply. It is present in cyanide poisoning and severe diphtheria. O 2 therapy is not much useful in this type of hypoxia, however hyperbaric O 2 therapy is beneficial. Effects of hypoxia Hypoxia has its effects at cellular, tissue and organ level. It can alter cellular transcription factors and thus protein expression. It has effect on brain function and produces symptoms like dizziness, drowsiness and headache. It can also affect ventilation. Chronic or Long term hypoxia can result in cell and tissue death. Summary The pressure gradient in pulmonary circulation system is much less than that in systemic circulation. Both ventilation and perfusion are greater at base of the lung as compared to apex but ventilation-perfusion ratio is lower at base as compared to apex. Partial pressures of oxygen and carbon dioxide at alveoli and pulmonary capillary determine net flow of these gases. The amount of O 2 present in blood is determined by the amount dissolved and amount bound to Hb. One molecule of Hb binds with four molecules of O 2. CO 2 in blood is rapidly converted to H 2CO 3 due to activity of enzyme carbonic anhydrase. It also readily forms carbamino compounds with blood proteins. It is also transported in dissolved form. Hypoxia is deficiency of O 2 at issue level. There are four types of hypoxia and they are hypoxic hypoxia, anemic hypoxia, stagnant hypoxia and histotoxic hypoxia. O 2 therapy is most useful in hypoxic hypoxia.

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