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Baroreflex Control of Systemic Arterial Pressure and Vascular Bed

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Methods of Analysis
1.1 Evaluation of Baroreflex and Experimental Procedures
1.2 Elements of Control Theory for Physiological Barostat
1.3 Estimation of Open‐Loop Characteristics From Closed‐Loop Perturbations
1.4 Summary
2 Overall Performance of Baroreflex
2.1 Static Open‐Loop Performance
2.2 Dynamic Performance
2.3 Summation of Baroreflex
2.4 Modulation of Baroreceptor Reflex
2.5 Summary
3 Effector Mechanisms for Arterial Pressure Control
3.1 Reflex Control of Cardiac Output Versus Vascular Resistance
3.2 Nonuniform Efferent Signals
4 Epilogue
4.1 Efferent Innervation
4.2 Nonlinear Characteristics
4.3 Reflex in Intact Subject
4.4 Reflex Control of Cardiac Output and Vascular Resistance and Capacity
Figure 1. Figure 1.

Triphasic response of blood pressure in cat to aortic nerve stimulation with increasingly stronger stimuli. Pressor response at middle intensity probably caused by excitation of chemoreceptor fibers in cat aortic nerve. Concomitant increase in ventilation supports this explanation. •, Chemoreceptor fibers; ○, depressor fibers recruited by changing stimulus voltage (size indicates fiber diameter).

From Douglas and Schaumann
Figure 2. Figure 2.

Arterial pressure response in dog to depulsation of carotid sinus pressure (CSP) by carotid artery occlusion. During occlusion, pressure in carotid sinuses became nonpulsatile (3rd panel), which caused such a strong systemic pressor effect that CSP increased above mean level before occlusion (4th panel).

Figure 3. Figure 3.

Surgical preparation for reversible vascular isolation of carotid sinuses, which changes CSP by infusion through catheter and inflation of occluding cuffs in conscious dogs.

From Stephenson and Donald
Figure 4. Figure 4.

Surgical preparation for isolating aortic arch baroreceptor region while left ventricle ejects blood into compliance chamber with constant mean pressure and constant‐flow pump perfuses head and lower body.

From Allison, Sagawa, and Kumada
Figure 5. Figure 5.

Positive‐ vs. negative‐feedback control in open‐ or closed‐loop configuration. G, gain. Note difference in input port between conditions of closed (left) and open (right) loop.

Figure 6. Figure 6.

A: open‐loop input‐output relationship curve of a negative‐feedback system. When input is at threshold (Ith), output is maximum (Omax); when input is at saturation (Lsat), output is minimum (Omin). M, optimum input at which slope of curve (gain) is maximum; C, closed‐loop equilibrium point (input = output). M and C may or may not coincide. B: open‐loop gain (i.e., slope of curve in A) as function of input. Gmax, maximum gain when input is optimum (Io); σ, magnitude of deviation of I from Io at which gain decreases to 0.6 Gmax if gain is normal density (Gaussian) distribution curve.

Figure 7. Figure 7.

A: depressor responses of arterial pressure to series of increases in isolated right CSP in rabbit. Left carotid sinus nerve and vagi cut. Time signal, 1 min. B: Koch's Blutdruckcharakteristik (——) and its derivative (gain) curve (—–) in dog. Vertical lines (from left): threshold, optimum, and saturation pressure in isolated carotid sinus.

From Koch
Figure 8. Figure 8.

Secondary counteraction of aortic arch baroreceptor reflex with a gain (Ga) to primary effect of carotid sinus reflex with an open‐loop gain (Ge). When input ΔCSP is given to isolated carotid sinuses, output of carotid sinus reflex should be a change in systemic arterial pressure (ΔSAP = Ge · ΔCSP). However, this is weakened by aortic reflex buffering ΔSAPe with a counteraction ().

From Eq. : = (Gcs · ΔCSP)/(1 + Ga)
Figure 9. Figure 9.

Open‐loop relationship between CSP (ISP) and SAP in 16 rabbits before (○——○) and after (○‐ ‐ ‐ ‐ ‐○) transection of aortic nerves. Vertical bars, SEM. Denervation elevated relation curve only over low‐CSP range. Effect of blood volume expansion (VE) by fluid infusion until right atrial pressure increased to 10 cmH2O, which elevated relation curve only in high‐CSP range before aortic denervation

From Chen et al.
Figure 10. Figure 10.

Top: effect of sequentially opening multiple negative‐feedback reflex loops (with open‐loop gains G2, G3, G4 all having a value of unity) on pressure output (ΔPo) of an opened negative‐feedback loop with a gain G1 = 1, in response to pressure input (ΔPi). Bottom: apparent gain of initially opened reflex (ΔPo/ΔPi) increasing nonlinearly as counteracting loops (with unity gain) are eliminated sequentially. , , and , elimination of loops represented by these gains. True gain of unity cannot be observed until all counteracting loops are removed.

Figure 11. Figure 11.

Top: how sequential opening of 4 parallel negative‐feedback loops, all with unity gain, affects weakening of a pressure disturbance (ΔPd, e.g., posthemorrhage hypotension) by these loops. Bottom: •———•, disturbance effect reduced to 20% (ΔPo = ΔPd)/5) when all loops are intact but gradually and nonlinearly increased as loops were sequentially removed (G1–G4). ○‐ ‐ ‐ ‐○, Overall system gain [(ΔPd/ΔPo) − 1; Eq. ].

Figure 12. Figure 12.

Variations in relationship of left aortic nerve (A) to vagus nerve, superior laryngeal nerve, nodose ganglion, and superior cervical ganglion in dog.

From Hashimoto and Hirohata
Figure 13. Figure 13.

Mean CSP‐mean SAP relationship with 4 pulse pressures. Average from 14–17 dogs anesthetized with chloralose‐urethane. Note leftward shift of portion with steepest slope during increase in pulse pressure.

Adapted from Schmidt, Kumada, and Sagawa
Figure 14. Figure 14.

Two ways the cardiovascular system is perturbed to estimate arterial baroreflex gain from closed‐loop responses. Perturbation I: cases like head‐level change, carotid artery occlusion, or application of a negative pressure on the neck. Perturbation II: circumstances like hemorrhage and transfusion. SAPd1 and SAPd2, 2 kinds of disturbing pressures for perturbations I and II, respectively.

Figure 15. Figure 15.

A: plaster cylinder around neck to apply positive or negative CSP to change transmural pressure in anesthetized and intubated dogs. B: average response (———) of arterial pressure to changes in estimated transmural pressure of carotid sinuses from 12 dogs. Data fit to growth (or logistic function) curve; ‐ ‐ ‐ ‐ ‐, derivative (i.e., gain curve). Maximum gain (1.2) occurred at unperturbed CSP (155 mmHg), and magnitude of deviation (σ) in Eq. was about ±25 mmHg.

From Shubrooks
Figure 16. Figure 16.

A: baroreceptor reflex where SAPd1 is introduced. Gbaro gain of baroreflex. B: ΔSAP in response to SAPd1. Eq. estimates Gbaro from ΔSAP and ΔCSP.

Figure 17. Figure 17.

A: baroreceptor reflex into which SAPd2 is introduced. B: response of arterial pressure to SAPd2. Unlike the case in Fig. , ΔSAP = ΔCSP and a different formula (Eq. ) is needed to estimate Gbaro.

Figure 18. Figure 18.

A: hysteresis in mean sinus nerve (multifiber) activity as mean aortic pressure decreased and increased. Average from 7 nerves as percent of maximum. B: hysteresis in arterial pressure response to stepwise increases and decreases in pulsatile CSP. Average from 11 cats.

A from Pelletier et al. , by permission of the American Heart Association, Inc.; B adapted from Kendrick et al.
Figure 19. Figure 19.

Isolated CSP (ISP) in relation to arterial pressure (A and C) and gain (ΔSAP/ΔISP; B and D). Data from 15 dogs anesthetized with chloralose. o, Data during 25‐s stimulation of hypothalamic defense area; • with bars, means ± SE in control condition. Curves in C and D are best‐fit normal cumulative distribution curve and normal density distribution curve, respectively.

From Kumada, Sagawa, et al.
Figure 20. Figure 20.

Effect of aortic baroreceptor reflex on systemic pressure perfusion and cardiac performance with isolated aortic arch preparation of Fig. . Percent changes from control collected and means ± SE plotted from 20 dogs.

From Allison, Sagawa, and Kumada
Figure 21. Figure 21.

Marked effect of pulsating isolated CSP on constant‐flow perfusion pressure in total systemic vascular bed of dog (left) as opposed to no effect of pulsating isolated aortic arch pressure in same dog (right). • and ○, Data when receptor region pressure was pulsated with peak‐to‐peak amplitude of 62 mmHg; ▴ and ▵, data with 0 pulse pressure. Proximity of solid and open symbols simply indicates reproducibility of data.

From Angell‐James and Daly
Figure 22. Figure 22.

Effect of stimulating cervical sympathetic nerve on carotid sinus reflex control of mean arterial pressure (A) and heart rate (B). •, Control; ○, during nerve stimulation. Top: mean difference ± SE caused by sympathetic stimulation. Effect depends on CSP level.

From Bolter and Ledsome
Figure 23. Figure 23.

Dynamic performance of canine aortic baroreceptor reflex control of systemic vascular resistance (1st channel), cardiac output (CO; 3rd channel), and heart rate (4th channel) as studied by sinusoidal change of isolated aortic arch pressure (2nd channel) with a peak‐to‐peak amplitude of 36 mmHg and at frequencies of 1/30 Hz (left), 1/8 Hz (middle), and 1/4 Hz (right). In preparation of Fig. .

From Allison
Figure 24. Figure 24.

Bode plots of frequency response from 3 dogs (different symbols) and response of nonlinear model (curves). Upper part of plot: amplitude ratio of output arterial pressure sinusoids to input CSP sinusoids (gain) as function of frequency of input sinusoid. Lower part: phase lag of output sinusoids to input sinusoids. ‐ ‐ ‐→, Phase margin and gain margin for instability.

From Levison et al. , by permission of the American Heart Association, Inc.
Figure 25. Figure 25.

Top: effects of pulsation frequency in isolated carotid sinus with fixed amplitude (50 mmHg peak to peak) on arterial pressure, CO, and total peripheral resistance. Bottom: effects of pulsation amplitude with fixed frequency (2 Hz) in 14–17 dogs.

Adapted from Schmidt, Kumada, and Sagawa
Figure 26. Figure 26.

Definition of linear (additive), facilitatory, and inhibitory (occlusive) summation. Latter 2 from interaction within a system. S, black‐box system; I, input; O, output; subscripts a, b, and c, variables.

Figure 27. Figure 27.

Summation of bilateral carotid sinus reflex control of left ventricular contractile state measured by change in isovolumic peak systolic pressure (ΔLVSP). Effects of 9 combinations of left and right CSP studied in 5 dogs and average fit to a nonlinear (interactive) model (Eq. ). Dots are not plots of data.

From Martin et al. , by permission of the American Heart Association, Inc.
Figure 28. Figure 28.

Linear (additive) summation between carotid sinus reflex control and aortic arch reflex control of arterial pressure (P) in anesthetized dogs. Abscissa, sum of separate reflex effects on P; ordinate, magnitude of simultaneous effect of both reflexes; line of identity, additive summation between 2 reflexes.

From Donald and Edis
Figure 29. Figure 29.

Left: effect of vagotomy on CSP‐arterial pressure relationship. •, Prevagotomy data; ○, postvagotomy data. Average from 5 dogs. Right: effect of anesthetizing same dogs with chloralose (60–80 mg/kg). Arrows, increase in arterial pressure on bilateral carotid occlusion. ‐ ‐ ‐ ‐, Control arterial pressure when carotid sinuses were not isolated.

From Stephenson and Donald
Figure 30. Figure 30.

Absence of marked effect of denervating carotid sinus and aortic baroreflex afferents on hemodynamic responses to moderate exercise in dog.

From McRitchie et al.
Figure 31. Figure 31.

Marked attenuation of carotid sinus baroreceptor reflex in conscious dog with heart failure caused by pulmonary stenosis (right) as opposed to strong pressor and other hemodynamic responses to bilateral common carotid artery occlusion (left). Heart rate response is absent despite similar preocclusion rate and marked attenuation of increase in mesenteric vascular resistance (bottom rows).

From Higgins et al.
Figure 32. Figure 32.

Effects of volume expansion (VE) and vagally mediated reflexes (VAGI) on ISP‐SAP relationship in 16 rabbits anesthetized with pentobarbital sodium (35 mg/kg). Aortic nerve transected and vagal efferent signal blocked with methylatropine (0.5 mg/kg). Dextran solution in saline was infused until atrial pressure reached 10 cmH2O for VE. VE effects are confined to lower ISP region. In presence of vagal afferents, VE reduces SAPmax and reflex gain; in absence of vagal afferents, VE mildly elevates SAPmax (cf. Fig. ).

From Chen et al.
Figure 33. Figure 33.

Influence of various parameters of carotid sinus reflex on CSP‐SAP relationship fitted to normal cumulative distribution curve function (Eq. ). Only those events that affect SAPmax and no other parameters of reflex system (Gmax, CSPopt, σ) cause a parallel vertical shift of relation curve. For examples of analysis see Fig. and Table .

Figure 34. Figure 34.

Cardiovascular processes that determine mean SAP (blocks with solid arrows) and the arterial baroreceptor reflex system efferents (open arrows), which control major components of cardiovascular processes. Mathematical functions (f1 and f2) relate input to output.

Figure 35. Figure 35.

A: effects of ISP on Starling CO curve represented by aortic flow (AF)‐mean right atrial pressure (MRAP) relation curve. ISP set at 4 levels, and at each the Starling curve was determined by bleeding and transfusing a solution of blood and dextran in closed‐chest dogs; top curve from pooled data with CSP at 75 and 100 mmHg. Number of dogs in parentheses below ISP value. Mean aortic pressure held at 100 mmHg throughout measurements by servo‐pump system that infused or withdrew blood from or into systemic artery .‐ ‐ ‐ ‐, 3rd‐Order polynomial function of MRAP best fit to experimental data. With this function, carotid sinus reflex simply modifies Starling relation parameter. B: effect of afterload mean aortic pressure (MAP) on ISP‐AF relationship. When MAP was fixed at 100 mmHg, increase in ISP significantly decreased AF. When MAP could change with ISP, variation of AF was greatly attenuated because of afterload effect.

From Kostiuk, Sagawa, and Shoukas , by permission of the American Heart Association, Inc.
Figure 36. Figure 36.

Changes in reservoir blood volume (top panel) caused by ΔCSP (2nd panel). Reservoir and constant‐flow pump inserted between caval veins and right atrium of dog. Constant inferior vena caval pressure (4th panel) and systemic perfusion flow (5th panel), but carotid sinus reflex was allowed to change SAP (3rd panel). ‐ ‐ ‐ ‐ ‐ On top panel reconstructs original trace (———), which had to be biased for maximum recording sensitivity. Increase in reservoir blood volume signifies decrease in blood volume in lumped venous capacity vessel in excess of passively increased blood volume in lumped arterial bed due to reflex increase in arterial pressure.

From Shoukas and Sagawa , by permission of the American Heart Association, Inc.
Figure 37. Figure 37.

Apparent arterial pressure‐flow relation (o———o) when CO is changed under closed‐loop control by arterial baroreflex. Variety of rectilinear resistances under open‐loop baroreflex (parametric) control (‐ ‐ ‐ ‐). In closed‐loop condition, pressure‐flow relation shifts from 1 line to another as reflex control varies with arterial pressure. P0, pressure axis intercept of extrapolated linear resistance; P0,a, apparent arterial pressure at 0 flow from extrapolation of quasi‐linear range of pressure‐flow relation curves under closed‐loop reflex control.

Based on data from Sagawa and Eisner
Figure 38. Figure 38.

Regional resistances as function of CSP under different anesthetics. □, α‐Chloralose (80 mg/kg, n = 14); Δ, pentobarbital sodium (25 mg/kg, n = 14); •, halothane (1%, n = 12). Bars, SEM. All data from vagotomized dogs. Different scales for ordinate.

From Cox and Bagshaw


Figure 1.

Triphasic response of blood pressure in cat to aortic nerve stimulation with increasingly stronger stimuli. Pressor response at middle intensity probably caused by excitation of chemoreceptor fibers in cat aortic nerve. Concomitant increase in ventilation supports this explanation. •, Chemoreceptor fibers; ○, depressor fibers recruited by changing stimulus voltage (size indicates fiber diameter).

From Douglas and Schaumann


Figure 2.

Arterial pressure response in dog to depulsation of carotid sinus pressure (CSP) by carotid artery occlusion. During occlusion, pressure in carotid sinuses became nonpulsatile (3rd panel), which caused such a strong systemic pressor effect that CSP increased above mean level before occlusion (4th panel).



Figure 3.

Surgical preparation for reversible vascular isolation of carotid sinuses, which changes CSP by infusion through catheter and inflation of occluding cuffs in conscious dogs.

From Stephenson and Donald


Figure 4.

Surgical preparation for isolating aortic arch baroreceptor region while left ventricle ejects blood into compliance chamber with constant mean pressure and constant‐flow pump perfuses head and lower body.

From Allison, Sagawa, and Kumada


Figure 5.

Positive‐ vs. negative‐feedback control in open‐ or closed‐loop configuration. G, gain. Note difference in input port between conditions of closed (left) and open (right) loop.



Figure 6.

A: open‐loop input‐output relationship curve of a negative‐feedback system. When input is at threshold (Ith), output is maximum (Omax); when input is at saturation (Lsat), output is minimum (Omin). M, optimum input at which slope of curve (gain) is maximum; C, closed‐loop equilibrium point (input = output). M and C may or may not coincide. B: open‐loop gain (i.e., slope of curve in A) as function of input. Gmax, maximum gain when input is optimum (Io); σ, magnitude of deviation of I from Io at which gain decreases to 0.6 Gmax if gain is normal density (Gaussian) distribution curve.



Figure 7.

A: depressor responses of arterial pressure to series of increases in isolated right CSP in rabbit. Left carotid sinus nerve and vagi cut. Time signal, 1 min. B: Koch's Blutdruckcharakteristik (——) and its derivative (gain) curve (—–) in dog. Vertical lines (from left): threshold, optimum, and saturation pressure in isolated carotid sinus.

From Koch


Figure 8.

Secondary counteraction of aortic arch baroreceptor reflex with a gain (Ga) to primary effect of carotid sinus reflex with an open‐loop gain (Ge). When input ΔCSP is given to isolated carotid sinuses, output of carotid sinus reflex should be a change in systemic arterial pressure (ΔSAP = Ge · ΔCSP). However, this is weakened by aortic reflex buffering ΔSAPe with a counteraction ().

From Eq. : = (Gcs · ΔCSP)/(1 + Ga)


Figure 9.

Open‐loop relationship between CSP (ISP) and SAP in 16 rabbits before (○——○) and after (○‐ ‐ ‐ ‐ ‐○) transection of aortic nerves. Vertical bars, SEM. Denervation elevated relation curve only over low‐CSP range. Effect of blood volume expansion (VE) by fluid infusion until right atrial pressure increased to 10 cmH2O, which elevated relation curve only in high‐CSP range before aortic denervation

From Chen et al.


Figure 10.

Top: effect of sequentially opening multiple negative‐feedback reflex loops (with open‐loop gains G2, G3, G4 all having a value of unity) on pressure output (ΔPo) of an opened negative‐feedback loop with a gain G1 = 1, in response to pressure input (ΔPi). Bottom: apparent gain of initially opened reflex (ΔPo/ΔPi) increasing nonlinearly as counteracting loops (with unity gain) are eliminated sequentially. , , and , elimination of loops represented by these gains. True gain of unity cannot be observed until all counteracting loops are removed.



Figure 11.

Top: how sequential opening of 4 parallel negative‐feedback loops, all with unity gain, affects weakening of a pressure disturbance (ΔPd, e.g., posthemorrhage hypotension) by these loops. Bottom: •———•, disturbance effect reduced to 20% (ΔPo = ΔPd)/5) when all loops are intact but gradually and nonlinearly increased as loops were sequentially removed (G1–G4). ○‐ ‐ ‐ ‐○, Overall system gain [(ΔPd/ΔPo) − 1; Eq. ].



Figure 12.

Variations in relationship of left aortic nerve (A) to vagus nerve, superior laryngeal nerve, nodose ganglion, and superior cervical ganglion in dog.

From Hashimoto and Hirohata


Figure 13.

Mean CSP‐mean SAP relationship with 4 pulse pressures. Average from 14–17 dogs anesthetized with chloralose‐urethane. Note leftward shift of portion with steepest slope during increase in pulse pressure.

Adapted from Schmidt, Kumada, and Sagawa


Figure 14.

Two ways the cardiovascular system is perturbed to estimate arterial baroreflex gain from closed‐loop responses. Perturbation I: cases like head‐level change, carotid artery occlusion, or application of a negative pressure on the neck. Perturbation II: circumstances like hemorrhage and transfusion. SAPd1 and SAPd2, 2 kinds of disturbing pressures for perturbations I and II, respectively.



Figure 15.

A: plaster cylinder around neck to apply positive or negative CSP to change transmural pressure in anesthetized and intubated dogs. B: average response (———) of arterial pressure to changes in estimated transmural pressure of carotid sinuses from 12 dogs. Data fit to growth (or logistic function) curve; ‐ ‐ ‐ ‐ ‐, derivative (i.e., gain curve). Maximum gain (1.2) occurred at unperturbed CSP (155 mmHg), and magnitude of deviation (σ) in Eq. was about ±25 mmHg.

From Shubrooks


Figure 16.

A: baroreceptor reflex where SAPd1 is introduced. Gbaro gain of baroreflex. B: ΔSAP in response to SAPd1. Eq. estimates Gbaro from ΔSAP and ΔCSP.



Figure 17.

A: baroreceptor reflex into which SAPd2 is introduced. B: response of arterial pressure to SAPd2. Unlike the case in Fig. , ΔSAP = ΔCSP and a different formula (Eq. ) is needed to estimate Gbaro.



Figure 18.

A: hysteresis in mean sinus nerve (multifiber) activity as mean aortic pressure decreased and increased. Average from 7 nerves as percent of maximum. B: hysteresis in arterial pressure response to stepwise increases and decreases in pulsatile CSP. Average from 11 cats.

A from Pelletier et al. , by permission of the American Heart Association, Inc.; B adapted from Kendrick et al.


Figure 19.

Isolated CSP (ISP) in relation to arterial pressure (A and C) and gain (ΔSAP/ΔISP; B and D). Data from 15 dogs anesthetized with chloralose. o, Data during 25‐s stimulation of hypothalamic defense area; • with bars, means ± SE in control condition. Curves in C and D are best‐fit normal cumulative distribution curve and normal density distribution curve, respectively.

From Kumada, Sagawa, et al.


Figure 20.

Effect of aortic baroreceptor reflex on systemic pressure perfusion and cardiac performance with isolated aortic arch preparation of Fig. . Percent changes from control collected and means ± SE plotted from 20 dogs.

From Allison, Sagawa, and Kumada


Figure 21.

Marked effect of pulsating isolated CSP on constant‐flow perfusion pressure in total systemic vascular bed of dog (left) as opposed to no effect of pulsating isolated aortic arch pressure in same dog (right). • and ○, Data when receptor region pressure was pulsated with peak‐to‐peak amplitude of 62 mmHg; ▴ and ▵, data with 0 pulse pressure. Proximity of solid and open symbols simply indicates reproducibility of data.

From Angell‐James and Daly


Figure 22.

Effect of stimulating cervical sympathetic nerve on carotid sinus reflex control of mean arterial pressure (A) and heart rate (B). •, Control; ○, during nerve stimulation. Top: mean difference ± SE caused by sympathetic stimulation. Effect depends on CSP level.

From Bolter and Ledsome


Figure 23.

Dynamic performance of canine aortic baroreceptor reflex control of systemic vascular resistance (1st channel), cardiac output (CO; 3rd channel), and heart rate (4th channel) as studied by sinusoidal change of isolated aortic arch pressure (2nd channel) with a peak‐to‐peak amplitude of 36 mmHg and at frequencies of 1/30 Hz (left), 1/8 Hz (middle), and 1/4 Hz (right). In preparation of Fig. .

From Allison


Figure 24.

Bode plots of frequency response from 3 dogs (different symbols) and response of nonlinear model (curves). Upper part of plot: amplitude ratio of output arterial pressure sinusoids to input CSP sinusoids (gain) as function of frequency of input sinusoid. Lower part: phase lag of output sinusoids to input sinusoids. ‐ ‐ ‐→, Phase margin and gain margin for instability.

From Levison et al. , by permission of the American Heart Association, Inc.


Figure 25.

Top: effects of pulsation frequency in isolated carotid sinus with fixed amplitude (50 mmHg peak to peak) on arterial pressure, CO, and total peripheral resistance. Bottom: effects of pulsation amplitude with fixed frequency (2 Hz) in 14–17 dogs.

Adapted from Schmidt, Kumada, and Sagawa


Figure 26.

Definition of linear (additive), facilitatory, and inhibitory (occlusive) summation. Latter 2 from interaction within a system. S, black‐box system; I, input; O, output; subscripts a, b, and c, variables.



Figure 27.

Summation of bilateral carotid sinus reflex control of left ventricular contractile state measured by change in isovolumic peak systolic pressure (ΔLVSP). Effects of 9 combinations of left and right CSP studied in 5 dogs and average fit to a nonlinear (interactive) model (Eq. ). Dots are not plots of data.

From Martin et al. , by permission of the American Heart Association, Inc.


Figure 28.

Linear (additive) summation between carotid sinus reflex control and aortic arch reflex control of arterial pressure (P) in anesthetized dogs. Abscissa, sum of separate reflex effects on P; ordinate, magnitude of simultaneous effect of both reflexes; line of identity, additive summation between 2 reflexes.

From Donald and Edis


Figure 29.

Left: effect of vagotomy on CSP‐arterial pressure relationship. •, Prevagotomy data; ○, postvagotomy data. Average from 5 dogs. Right: effect of anesthetizing same dogs with chloralose (60–80 mg/kg). Arrows, increase in arterial pressure on bilateral carotid occlusion. ‐ ‐ ‐ ‐, Control arterial pressure when carotid sinuses were not isolated.

From Stephenson and Donald


Figure 30.

Absence of marked effect of denervating carotid sinus and aortic baroreflex afferents on hemodynamic responses to moderate exercise in dog.

From McRitchie et al.


Figure 31.

Marked attenuation of carotid sinus baroreceptor reflex in conscious dog with heart failure caused by pulmonary stenosis (right) as opposed to strong pressor and other hemodynamic responses to bilateral common carotid artery occlusion (left). Heart rate response is absent despite similar preocclusion rate and marked attenuation of increase in mesenteric vascular resistance (bottom rows).

From Higgins et al.


Figure 32.

Effects of volume expansion (VE) and vagally mediated reflexes (VAGI) on ISP‐SAP relationship in 16 rabbits anesthetized with pentobarbital sodium (35 mg/kg). Aortic nerve transected and vagal efferent signal blocked with methylatropine (0.5 mg/kg). Dextran solution in saline was infused until atrial pressure reached 10 cmH2O for VE. VE effects are confined to lower ISP region. In presence of vagal afferents, VE reduces SAPmax and reflex gain; in absence of vagal afferents, VE mildly elevates SAPmax (cf. Fig. ).

From Chen et al.


Figure 33.

Influence of various parameters of carotid sinus reflex on CSP‐SAP relationship fitted to normal cumulative distribution curve function (Eq. ). Only those events that affect SAPmax and no other parameters of reflex system (Gmax, CSPopt, σ) cause a parallel vertical shift of relation curve. For examples of analysis see Fig. and Table .



Figure 34.

Cardiovascular processes that determine mean SAP (blocks with solid arrows) and the arterial baroreceptor reflex system efferents (open arrows), which control major components of cardiovascular processes. Mathematical functions (f1 and f2) relate input to output.



Figure 35.

A: effects of ISP on Starling CO curve represented by aortic flow (AF)‐mean right atrial pressure (MRAP) relation curve. ISP set at 4 levels, and at each the Starling curve was determined by bleeding and transfusing a solution of blood and dextran in closed‐chest dogs; top curve from pooled data with CSP at 75 and 100 mmHg. Number of dogs in parentheses below ISP value. Mean aortic pressure held at 100 mmHg throughout measurements by servo‐pump system that infused or withdrew blood from or into systemic artery .‐ ‐ ‐ ‐, 3rd‐Order polynomial function of MRAP best fit to experimental data. With this function, carotid sinus reflex simply modifies Starling relation parameter. B: effect of afterload mean aortic pressure (MAP) on ISP‐AF relationship. When MAP was fixed at 100 mmHg, increase in ISP significantly decreased AF. When MAP could change with ISP, variation of AF was greatly attenuated because of afterload effect.

From Kostiuk, Sagawa, and Shoukas , by permission of the American Heart Association, Inc.


Figure 36.

Changes in reservoir blood volume (top panel) caused by ΔCSP (2nd panel). Reservoir and constant‐flow pump inserted between caval veins and right atrium of dog. Constant inferior vena caval pressure (4th panel) and systemic perfusion flow (5th panel), but carotid sinus reflex was allowed to change SAP (3rd panel). ‐ ‐ ‐ ‐ ‐ On top panel reconstructs original trace (———), which had to be biased for maximum recording sensitivity. Increase in reservoir blood volume signifies decrease in blood volume in lumped venous capacity vessel in excess of passively increased blood volume in lumped arterial bed due to reflex increase in arterial pressure.

From Shoukas and Sagawa , by permission of the American Heart Association, Inc.


Figure 37.

Apparent arterial pressure‐flow relation (o———o) when CO is changed under closed‐loop control by arterial baroreflex. Variety of rectilinear resistances under open‐loop baroreflex (parametric) control (‐ ‐ ‐ ‐). In closed‐loop condition, pressure‐flow relation shifts from 1 line to another as reflex control varies with arterial pressure. P0, pressure axis intercept of extrapolated linear resistance; P0,a, apparent arterial pressure at 0 flow from extrapolation of quasi‐linear range of pressure‐flow relation curves under closed‐loop reflex control.

Based on data from Sagawa and Eisner


Figure 38.

Regional resistances as function of CSP under different anesthetics. □, α‐Chloralose (80 mg/kg, n = 14); Δ, pentobarbital sodium (25 mg/kg, n = 14); •, halothane (1%, n = 12). Bars, SEM. All data from vagotomized dogs. Different scales for ordinate.

From Cox and Bagshaw
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Kiichi Sagawa. Baroreflex Control of Systemic Arterial Pressure and Vascular Bed. Compr Physiol 2011, Supplement 8: Handbook of Physiology, The Cardiovascular System, Peripheral Circulation and Organ Blood Flow: 453-496. First published in print 1983. doi: 10.1002/cphy.cp020314