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Human Physiology in an Aquatic Environment

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ABSTRACT

Water covers over 70% of the earth, has varying depths and temperatures and contains much of the earth's resources. Head‐out water immersion (HOWI) or submersion at various depths (diving) in water of thermoneutral (TN) temperature elicits profound cardiorespiratory, endocrine, and renal responses. The translocation of blood into the thorax and elevation of plasma volume by autotransfusion of fluid from cells to the vascular compartment lead to increased cardiac stroke volume and output and there is a hyperperfusion of some tissues. Pulmonary artery and capillary hydrostatic pressures increase causing a decline in vital capacity with the potential for pulmonary edema. Atrial stretch and increased arterial pressure cause reflex autonomic responses which result in endocrine changes that return plasma volume and arterial pressure to preimmersion levels. Plasma volume is regulated via a reflex diuresis and natriuresis. Hydrostatic pressure also leads to elastic loading of the chest, increasing work of breathing, energy cost, and thus blood flow to respiratory muscles. Decreases in water temperature in HOWI do not affect the cardiac output compared to TN; however, they influence heart rate and the distribution of muscle and fat blood flow. The reduced muscle blood flow results in a reduced maximal oxygen consumption. The properties of water determine the mechanical load and the physiological responses during exercise in water (e.g. swimming and water based activities). Increased hydrostatic pressure caused by submersion does not affect stroke volume; however, progressive bradycardia decreases cardiac output. During submersion, compressed gas must be breathed which introduces the potential for oxygen toxicity, narcosis due to nitrogen, and tissue and vascular gas bubbles during decompression and after may cause pain in joints and the nervous system. © 2015 American Physiological Society. Compr Physiol 5:1705‐1750, 2015.

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Figure 1. Figure 1. Summary of the circulatory, renal hormonal and neuro responses to thermoneutral HOWI. Central volume expansion and elevated CO are caused by compression of the dependent veins and autotransfusion. The rise in CO exceeds the metabolic demand at rest and during exercise. Activation of cardiovascular mechanoreceptors and increased atrial natriuretic peptide secretion sets into motion neural and hormonal mechanism that promote sodium and fluid loss, thus minimizing hypervolemia and theoretically readjusting blood flow to the prevailing oxygen uptake. The most powerful efferent mechanism affecting the kidney appears to be sympathetic nerves. The associated hormonal responses appear to modulate the primary renal response, with many additional factors modulating the basic renal neurohumoral responses to immersion. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().
Figure 2. Figure 2. While standing in air (A), right atrial pressure is low and gravity‐dependent veins are distended resulting in increased net plasma filtration in the limb capillaries. During HOWI (B), central volume expansion occurs with increased right atrial pressure and cardiac output. These result from hydrostatic compression of the dependent tissues and capillary reabsorption, or an autotransfusion in the dependent limbs. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().
Figure 3. Figure 3. There are three major fluid shifts that occur during HOWI (upper flow diagram): across capillary, across the cell wall and across the kidney. The lymphatic fluid shift is minor. These compartmental fluid changes are then illustrated (graph) for the 120‐min upright, thermoneutral HOWI. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().
Figure 4. Figure 4. Human plasma volume (PV) expressed as a percentage change during HOWI in TN water (35°C) (squares) and sitting in thermoneutral air (22°C) (circles) from their respective control values. The * indicate that HOWI values are significantly higher than the air counterparts.
Figure 5. Figure 5. Resting cardiac output (Q, L/m) is plotted as a function of time for humans resting HOWI in TN water (35°C) (squares) and sitting in TN air (22°C)(circles). The * indicate that HOWI values are significantly higher than the air counterparts.
Figure 6. Figure 6. Resting lower limb blood flow measured by plethysmography is plotted as function to time of HOWI. The * indicates a significant increase above air control.
Figure 7. Figure 7. Muscle blood flow measured by 133Xe washout from quadriceps muscle at rest and during cycling exercise is plotted as a function of VO2 for air and HOWI in 20°C, 25°C, and 30°C water. The dashed lines represent a constant (a‐v)O2 as a function of VO2.
Figure 8. Figure 8. Cardiac output (Q, L/min) is plotted as a function of oxygen consumption (L/min) for air and HOWI in 20°C to 35°C water at rest (VO2 < 1.0 L/min) and exercise (VO2 > 1.0 L/min).
Figure 9. Figure 9. An early version of the Gauer‐Henry hypothesis. Arrows indicate an increase or stimulation of the variable, darkened arrows indicate a decrease or inhibition of the variable. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().
Figure 10. Figure 10. Regional lung volume determined by radioactive labeling of red blood cells is plotted as a function of the depth of water immersion (progressive increasing pulmonary blood flow). The lung was divided into nine equal layers from the apex to the top of the lung. The slopes of the increase in lung volume as a function of increasing cardiac output for the nine layers were not significantly different from each other. In parallel measurements diffusion capacity (DLCO) was measured and confirmed the increase in pulmonary blood volume was a result of capillary enlargement, and not capillary recruitment (data not shown).
Figure 11. Figure 11. Net energy expenditure (E′, kW) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.
Figure 12. Figure 12. Net energy cost (C, kJ · m−1) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). The continuous lines represent iso‐metabolic power hyperbolae of 0.5, 1.0, 1.5, and 2 kW (from bottom to top). L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.
Figure 13. Figure 13. Cycle frequency (f, cycles · min−1) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). The average distance the body/boat travels per cycle (dc, m · cycle−1) is indicated by the continuous lines irradiating from the origin. L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.
Figure 14. Figure 14. Active/passive drag (D, N) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.
Figure 15. Figure 15. Alveolar‐arterial PO2 difference versus gas density in four experiments. From Moon et al. with permission ().
Figure 16. Figure 16. Dead space/tidal volume ratio versus gas density. From Moon et al. with permission ().
Figure 17. Figure 17. Mechanism of pulmonary barotrauma in a diver breathing compressed gas and ascending while holding his breath. From Vann et al. with permission ().


Figure 1. Summary of the circulatory, renal hormonal and neuro responses to thermoneutral HOWI. Central volume expansion and elevated CO are caused by compression of the dependent veins and autotransfusion. The rise in CO exceeds the metabolic demand at rest and during exercise. Activation of cardiovascular mechanoreceptors and increased atrial natriuretic peptide secretion sets into motion neural and hormonal mechanism that promote sodium and fluid loss, thus minimizing hypervolemia and theoretically readjusting blood flow to the prevailing oxygen uptake. The most powerful efferent mechanism affecting the kidney appears to be sympathetic nerves. The associated hormonal responses appear to modulate the primary renal response, with many additional factors modulating the basic renal neurohumoral responses to immersion. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().


Figure 2. While standing in air (A), right atrial pressure is low and gravity‐dependent veins are distended resulting in increased net plasma filtration in the limb capillaries. During HOWI (B), central volume expansion occurs with increased right atrial pressure and cardiac output. These result from hydrostatic compression of the dependent tissues and capillary reabsorption, or an autotransfusion in the dependent limbs. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().


Figure 3. There are three major fluid shifts that occur during HOWI (upper flow diagram): across capillary, across the cell wall and across the kidney. The lymphatic fluid shift is minor. These compartmental fluid changes are then illustrated (graph) for the 120‐min upright, thermoneutral HOWI. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().


Figure 4. Human plasma volume (PV) expressed as a percentage change during HOWI in TN water (35°C) (squares) and sitting in thermoneutral air (22°C) (circles) from their respective control values. The * indicate that HOWI values are significantly higher than the air counterparts.


Figure 5. Resting cardiac output (Q, L/m) is plotted as a function of time for humans resting HOWI in TN water (35°C) (squares) and sitting in TN air (22°C)(circles). The * indicate that HOWI values are significantly higher than the air counterparts.


Figure 6. Resting lower limb blood flow measured by plethysmography is plotted as function to time of HOWI. The * indicates a significant increase above air control.


Figure 7. Muscle blood flow measured by 133Xe washout from quadriceps muscle at rest and during cycling exercise is plotted as a function of VO2 for air and HOWI in 20°C, 25°C, and 30°C water. The dashed lines represent a constant (a‐v)O2 as a function of VO2.


Figure 8. Cardiac output (Q, L/min) is plotted as a function of oxygen consumption (L/min) for air and HOWI in 20°C to 35°C water at rest (VO2 < 1.0 L/min) and exercise (VO2 > 1.0 L/min).


Figure 9. An early version of the Gauer‐Henry hypothesis. Arrows indicate an increase or stimulation of the variable, darkened arrows indicate a decrease or inhibition of the variable. Redrawn from Krasney () and reproduced with permission from Elsevier Limited, Oxford, United Kingdom ().


Figure 10. Regional lung volume determined by radioactive labeling of red blood cells is plotted as a function of the depth of water immersion (progressive increasing pulmonary blood flow). The lung was divided into nine equal layers from the apex to the top of the lung. The slopes of the increase in lung volume as a function of increasing cardiac output for the nine layers were not significantly different from each other. In parallel measurements diffusion capacity (DLCO) was measured and confirmed the increase in pulmonary blood volume was a result of capillary enlargement, and not capillary recruitment (data not shown).


Figure 11. Net energy expenditure (E′, kW) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.


Figure 12. Net energy cost (C, kJ · m−1) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). The continuous lines represent iso‐metabolic power hyperbolae of 0.5, 1.0, 1.5, and 2 kW (from bottom to top). L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.


Figure 13. Cycle frequency (f, cycles · min−1) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). The average distance the body/boat travels per cycle (dc, m · cycle−1) is indicated by the continuous lines irradiating from the origin. L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.


Figure 14. Active/passive drag (D, N) is plotted as a function of the speed (v, m · s−1) for different forms of aquatic locomotion (original data were interpolated over the aerobic speed range, see text for details). L: swimming by using the leg kick at the surface; UWF: underwater swimming with SCUBA diving equipment; AL: swimming the front crawl; K: kayaking; R: rowing. Taken from Pendergast et al. () with permission.


Figure 15. Alveolar‐arterial PO2 difference versus gas density in four experiments. From Moon et al. with permission ().


Figure 16. Dead space/tidal volume ratio versus gas density. From Moon et al. with permission ().


Figure 17. Mechanism of pulmonary barotrauma in a diver breathing compressed gas and ascending while holding his breath. From Vann et al. with permission ().
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David R. Pendergast, Richard E. Moon, John J. Krasney, Heather E. Held, Paola Zamparo. Human Physiology in an Aquatic Environment. Compr Physiol 2015, 5: 1705-1750. doi: 10.1002/cphy.c140018