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Head‐out Water Immersion: Animal Studies

John A. Krasney

10.1002/cphy.cp040238

Source: Supplement 14: Handbook of Physiology, Environmental Physiology

Originally published: 1996

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 The Gauer‐Henry Hypothesis
2 Ratio of Blood Flow to Metabolism in Water Immersion
3 Transcapillary Fluid Shift in Water Immersion
4 Water Immersion as a Model for Hypervolemia
4.1 Methods for Eliciting Hypervolemia
4.2 Methodological Considerations for Animal Studies
5 Cardiovascular Receptors: Circulatory, Renal, and Hormonal Influences
5.1 Cardiac Receptors
5.2 Arterial Baroreceptors
6 Cardiovascular Responses to Water Immersion
6.1 Central Hemodynamics
6.2 Transcapillary Fluid Shift
7 Regional Vascular Responses to Water Immersion
8 The Renal Response to Water Immersion
9 Hormonal Responses to Water Immersion
9.1 Methodological Considerations
9.2 Vasopressin
9.3 Renin‐Angiotensin II‐Aldosterone System
9.4 Atrial Natriuretic Peptide (ANP)
9.5 Renal Prostaglandins
9.6 Adrenergic System in Water Immersion
10 Conclusions
Figure 1. Figure 1.

Early version of the Gauer‐Henry hypothesis. Open arrows indicate an increase or stimulation of the variable, darkened arrows indicate a decrease or inhibition of the variable.
Figure 2. Figure 2.

Water immersion increases the ratio of blood flow (Q) to metabolism () at rest (left panel) where is constant. Autoregulation of blood flow occurs, but the level at which systemic flow is held constant over a range of perfusion pressures is shifted upward. As exercise raises the level (right panel), the level of systemic flow is elevated for any level during water immersion. [From Christie et al. 18 with permission.]
Figure 3. Figure 3.

While standing in air (left panel), right atrial pressure (Pra) is low, dependent veins are distended, and net filtration occurs in limb capillaries. During water immersion (right panel), central volume expansion occurs with increased Pra and cardiac output because (I) there is hydrostatic compression of dependent tissues and (II) capillary reabsorption or an autotransfusion occurs in dependent limbs.
Figure 4. Figure 4.

Sling frame assembly for the study of conscious animals during water immersion. PAO = aortic pressure; PLA, Pra = left and right atrial pressures, respectively; PPL = pleural pressure; QAO = aortic blood flow (electromagnetic flow transducer); PLV = left ventricular pressure (solid state pressure transducer).

Reprinted with permission of the American Physiological Society; from Hajduczok et al. 62
Figure 5. Figure 5.

Expanded version of the Gauer‐Henry hypothesis. Arrow symbols as in Figure 1.
Figure 6. Figure 6.

Resetting and change of sensitivity of heart rate limb of arterial baroreflex in an awake dog during water immersion. TSAP = transmural systolic arterial pressure; Saturation TSAP = point at which heart rate stopped changing as TSAP was elevated. Threshold TSAP could not be determined in the standing dog.

Reprinted with permission of the American Physiological Society, from Yoshino et al. 152
Figure 7. Figure 7.

Percent change of plasma volume (PV), pressure in an implanted Guyton capsule in subcutaneous tissue of dog forelimb (Pcapsule), calculated capillary hydrostatic pressure (Pcapillary) and plasma oncotic pressure (πP) in an awake animal undergoing water immersion at 37°C.

Reprinted with permission from the Best Publishing Company; from Krasney 84
Figure 8. Figure 8.

Changes in cephalic vein pressure (Pcephv); implanted Guyton capsule pressure (Pcps); and wick catheter pressure (Pwick) relative to the external hydrostatic reference pressure (Pref) during graded immersion in awake dogs.

[Reprinted with permission from the American Physiological Society; from Miki et al. 101
Figure 9. Figure 9.

Increase in extracellular fluid volume (Δ ECF) during water immersion commencing at time 0 in an anesthetized, splenectomized, bilaterally nephrectomized dog. ECF volume was estimated by the 125I‐iothalamate space.

Reprinted by permission of the American Physiological Society, from Miki et al. 102
Figure 10. Figure 10.

Plasma volume responses to immersion in anesthetized, nephrectomized dogs and in dogs with intact kidneys. The role of the kidney is to minimize hypervolemia during immersion.
Figure 11. Figure 11.

There are three major fluid shifts (Δ J) which occur during immersion; across the capillary (Jcap), across the cell wall (Jcell), and across the kidney (Ju). The fourth fluid shift in the lymphatics (JL is minor. ISF = interstitial fluid; ICF = intracellular fluid.
Figure 12. Figure 12.

a: If protein fails to move into the plasma compartment during immersion, the rise of interstitial protein concentration could cause cell water and perhaps ions to leave the cell osmotically. b: Graded hydrostatic compression could activate cell membrane mechanoreceptors and membrane pumps which would extrude ions and water from the cells (133A). c: Compliance differences between cell, interstitial fluid, and capillary could cause fluid to shift from cell to plasma in immersion.

Reprinted with permission from the Best Publishing Company; from Krasney 84
Figure 13. Figure 13.

As predicted by Guyton's analysis 61, the equilibrium condition for cardiac output (Q) and venous pressure (Pv) would move from the control state (A) to point B due to decreased venous capacity resulting from hydrostatic compression, and further to point C because of hypervolemia from autotransfusion. The net result is a higher cardiac output at a higher central venous pressure as predicted by the Frank‐Starling relationship.
Figure 14. Figure 14.

Percent changes (%Δ) in the distribution of regional blood flows () measured by radiolabeled microspheres during immersion in conscious dogs. Flow increases to certain tissues (skin, fat, respiratory muscles) can be accounted for while flow increases to abdominal viscera and skeletal muscle are not readily explained.

Reprinted with permission from the Best Publishing Company; from Krasney 84
Figure 15. Figure 15.

Renal responses to water immersion in volume replete (R) and nonreplete (NR) conscious dogs. Shaded and dashed areas represent responses during air‐timed control studies.
Figure 16. Figure 16.

Renal responses to water immersion in intact vs. cardiac‐denervated (CD) dogs. = urine flow, = sodium excretion, Cosm = osmotic clearance, = free water clearance. An identical increase in occurred in the CD dogs, but it was caused by a rise in instead of by the rise in Cosm and observed in intact dogs.

Reprinted with permission from the Best Publishing Company; from Krasney 84
Figure 17. Figure 17.

The role of the kidney in water immersion is to minimize the hypervolemia due to the shift of fluid out of the cell compartment.
Figure 18. Figure 18.

Summary of the circulatory, renal, hormonal, and neural responses to water immersion. Central volume expansion and the rise of cardiac output is caused by compression of dependent veins and by autotransfusion. The rise in cardiac output exceeds metabolic demand at rest and during exercise. Activation of cardiovascular mechanoreceptors and increased ANP secretion sets into motion neural and hormonal mechanisms which promote salt and fluid loss to minimize the hypervolemia and theoretically readjust systemic to the prevailing . The most powerful efferent mechanism affecting the kidney appears to be the sympathetic nerves. The associated hormonal responses appear to modulate the primary renal response. A large number of factors can modulate the basic renal neurohumoral response to immersion.


Figure 1.

Early version of the Gauer‐Henry hypothesis. Open arrows indicate an increase or stimulation of the variable, darkened arrows indicate a decrease or inhibition of the variable.



Figure 2.

Water immersion increases the ratio of blood flow (Q) to metabolism () at rest (left panel) where is constant. Autoregulation of blood flow occurs, but the level at which systemic flow is held constant over a range of perfusion pressures is shifted upward. As exercise raises the level (right panel), the level of systemic flow is elevated for any level during water immersion. [From Christie et al. 18 with permission.]



Figure 3.

While standing in air (left panel), right atrial pressure (Pra) is low, dependent veins are distended, and net filtration occurs in limb capillaries. During water immersion (right panel), central volume expansion occurs with increased Pra and cardiac output because (I) there is hydrostatic compression of dependent tissues and (II) capillary reabsorption or an autotransfusion occurs in dependent limbs.



Figure 4.

Sling frame assembly for the study of conscious animals during water immersion. PAO = aortic pressure; PLA, Pra = left and right atrial pressures, respectively; PPL = pleural pressure; QAO = aortic blood flow (electromagnetic flow transducer); PLV = left ventricular pressure (solid state pressure transducer).

Reprinted with permission of the American Physiological Society; from Hajduczok et al. 62


Figure 5.

Expanded version of the Gauer‐Henry hypothesis. Arrow symbols as in Figure 1.



Figure 6.

Resetting and change of sensitivity of heart rate limb of arterial baroreflex in an awake dog during water immersion. TSAP = transmural systolic arterial pressure; Saturation TSAP = point at which heart rate stopped changing as TSAP was elevated. Threshold TSAP could not be determined in the standing dog.

Reprinted with permission of the American Physiological Society, from Yoshino et al. 152


Figure 7.

Percent change of plasma volume (PV), pressure in an implanted Guyton capsule in subcutaneous tissue of dog forelimb (Pcapsule), calculated capillary hydrostatic pressure (Pcapillary) and plasma oncotic pressure (πP) in an awake animal undergoing water immersion at 37°C.

Reprinted with permission from the Best Publishing Company; from Krasney 84


Figure 8.

Changes in cephalic vein pressure (Pcephv); implanted Guyton capsule pressure (Pcps); and wick catheter pressure (Pwick) relative to the external hydrostatic reference pressure (Pref) during graded immersion in awake dogs.

[Reprinted with permission from the American Physiological Society; from Miki et al. 101


Figure 9.

Increase in extracellular fluid volume (Δ ECF) during water immersion commencing at time 0 in an anesthetized, splenectomized, bilaterally nephrectomized dog. ECF volume was estimated by the 125I‐iothalamate space.

Reprinted by permission of the American Physiological Society, from Miki et al. 102


Figure 10.

Plasma volume responses to immersion in anesthetized, nephrectomized dogs and in dogs with intact kidneys. The role of the kidney is to minimize hypervolemia during immersion.



Figure 11.

There are three major fluid shifts (Δ J) which occur during immersion; across the capillary (Jcap), across the cell wall (Jcell), and across the kidney (Ju). The fourth fluid shift in the lymphatics (JL is minor. ISF = interstitial fluid; ICF = intracellular fluid.



Figure 12.

a: If protein fails to move into the plasma compartment during immersion, the rise of interstitial protein concentration could cause cell water and perhaps ions to leave the cell osmotically. b: Graded hydrostatic compression could activate cell membrane mechanoreceptors and membrane pumps which would extrude ions and water from the cells (133A). c: Compliance differences between cell, interstitial fluid, and capillary could cause fluid to shift from cell to plasma in immersion.

Reprinted with permission from the Best Publishing Company; from Krasney 84


Figure 13.

As predicted by Guyton's analysis 61, the equilibrium condition for cardiac output (Q) and venous pressure (Pv) would move from the control state (A) to point B due to decreased venous capacity resulting from hydrostatic compression, and further to point C because of hypervolemia from autotransfusion. The net result is a higher cardiac output at a higher central venous pressure as predicted by the Frank‐Starling relationship.



Figure 14.

Percent changes (%Δ) in the distribution of regional blood flows () measured by radiolabeled microspheres during immersion in conscious dogs. Flow increases to certain tissues (skin, fat, respiratory muscles) can be accounted for while flow increases to abdominal viscera and skeletal muscle are not readily explained.

Reprinted with permission from the Best Publishing Company; from Krasney 84


Figure 15.

Renal responses to water immersion in volume replete (R) and nonreplete (NR) conscious dogs. Shaded and dashed areas represent responses during air‐timed control studies.



Figure 16.

Renal responses to water immersion in intact vs. cardiac‐denervated (CD) dogs. = urine flow, = sodium excretion, Cosm = osmotic clearance, = free water clearance. An identical increase in occurred in the CD dogs, but it was caused by a rise in instead of by the rise in Cosm and observed in intact dogs.

Reprinted with permission from the Best Publishing Company; from Krasney 84


Figure 17.

The role of the kidney in water immersion is to minimize the hypervolemia due to the shift of fluid out of the cell compartment.



Figure 18.

Summary of the circulatory, renal, hormonal, and neural responses to water immersion. Central volume expansion and the rise of cardiac output is caused by compression of dependent veins and by autotransfusion. The rise in cardiac output exceeds metabolic demand at rest and during exercise. Activation of cardiovascular mechanoreceptors and increased ANP secretion sets into motion neural and hormonal mechanisms which promote salt and fluid loss to minimize the hypervolemia and theoretically readjust systemic to the prevailing . The most powerful efferent mechanism affecting the kidney appears to be the sympathetic nerves. The associated hormonal responses appear to modulate the primary renal response. A large number of factors can modulate the basic renal neurohumoral response to immersion.

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Article Title: Head‐out Water Immersion: Animal Studies
 

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John A. Krasney. Head‐out Water Immersion: Animal Studies. Compr Physiol 2011, Supplement 14: Handbook of Physiology, Environmental Physiology: 855-887. First published in print 1996. doi: 10.1002/cphy.cp040238