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Reflex Responses to Chemoreceptor Stimulation

Robert S. Fitzgerald, Sukhamay Lahiri

10.1002/cphy.cp030210

Source: Supplement 11: Handbook of Physiology, The Respiratory System, Control of Breathing

Originally published: 1986

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Historical Background
1.1 Beginnings
1.2 New Discoveries
2 Peripheral Chemoreceptors and Their Stimuli
2.1 Carotid Body Chemoreceptors
2.2 Aortic Body Chemoreceptors
2.3 Pulmonary Chemoreceptors
3 Conditioning of Afferent Stimulus from Peripheral Chemoreceptors
3.1 Receptor Responses to Environmental and Other Factors
3.2 Autonomic Input Into Carotid Body
3.3 Stimulus Profile
3.4 Timing of Stimulus
3.5 Other Afferent Stimuli Modifying Chemoreceptor Input
3.6 Effect of Central Nervous System Milieu on Neural Activity from Peripheral Chemoreceptors
4 Reflex Responses of Ventilatory Apparatus to Peripheral Chemoreceptor Stimulation
4.1 Factors Affecting Efferent Component of Response
4.2 Reflex Response of Static Lung Volumes
4.3 Reflex Dynamic Ventilatory Responses
4.4 Other Elements of Ventilatory Apparatus
4.5 Glomectomy and Asthma
5 Reflex Cardiovascular Responses to Peripheral Chemoreceptor Stimulation
5.1 Brain Stem and Autonomic Neural Output to Cardiovascular System
5.2 Reflex Changes in Cardiovascular Mechanical Variables
6 Other Reflex Responses to Peripheral Chemoreceptor Stimulation
6.1 Adrenal Cortex
6.2 Release of Norepinephrine From the Heart
6.3 Erythropoiesis
Figure 1. Figure 1.

Title pages of 3 earliest dissertations on the carotid body. Though Albrecht von Haller presided at first 2 presentations, actual presenters were Hartwig Taube (left panel) and Matthias Berckelmann (middle panel). Right panel: 1797 edition by Ernst Philipp Andersch of his father's earlier work that had been lost.

From Pick 356
Figure 2. Figure 2.

Gross structure and response of carotid body as presented by C. J. F. Heymans in his Nobel Lecture of December 12, 1945. He received the Nobel Prize in Physiology or Medicine in 1938 “for the discovery of the role played by the sinus and aortic mechanisms in the regulation of respiration.” Left panel: neural pathways from cardioaortic and carotid sinus zones. Right panel: pneumogram of chloralose‐anesthetized dog (A) and of blood pressure and cardiac rate (B). Time is in 3‐s intervals. Acetylcholine (0.1 mg) applied to chemoreceptors of glomus caroticum resulted in marked reflex hyperpnea and bradycardia.

From Heymans 198
Figure 3. Figure 3.

Single‐fiber chemoreceptor response to decreases in arterial O2 content in cat. Left panel: mean response (±SE) of 4 single carotid chemoreceptor fibers to decreases in O2 content of arterial blood (; vol %) achieved by either decreasing fraction of inspired O2 (hypoxic hypoxia) or by adding CO to the inspirate (CO hypoxia). Inset: response [impulses/s (IPS)] of single carotid chemoreceptor fiber to lowering of by adding CO. CTL, control; , partial pressure of O2 in arterial blood (mmHg); COHb, percent carboxyhemoglobin; , partial pressure of CO2 in arterial blood (mmHg); pHa, negative logarithm of H+ concentration in arterial blood; FBP, femoral arterial blood pressure (mmHg); A–D: blood samples taken at various times; RA, room air. To assure that fiber was not dead, cat was ventilated on 10% O2; fiber showed brisk increase in neural activity. Right panel: mean response of 3 single aortic chemoreceptor fibers to decreases in achieved as with carotid body responses. Though aortic body's response is larger in percent of control, control IPS is usually much lower for aortic fibers than for carotid fibers (cf. insets and Fig. 7). Inset: response of single aortic chemoreceptor fiber to lowering of by adding CO.

Courtesy of R. S. Fitzgerald and G. A. Dehghani
Figure 4. Figure 4.

Effects on single‐fiber activity of increases in at decreasing levels of . At low levels of , neural activity approaches zero regardless of value. On the assumption that normocapnia for cats is = 34–37 Torr, data show that response is linear as is lowered and that multiplicative interaction of O2 and CO2 still operates at these values.

From Lahiri and DeLaney 254
Figure 5. Figure 5.

Statistical analyses of responses of 10 recordings from carotid chemoreceptors (left panel) and 5 recordings from aortic chemoreceptors (right panel) to increases in at different levels of . The 3 levels (torr) for carotid responses were (bottom to top) >470, 105–115, and 54–58. The 4 levels (mmHg) for aortic responses were (bottom to top) >350, 93–97, 66–71, and 39–45. Responses were analyzed first by analysis of variance, and then individual means were grouped by Duncan new multiple‐range technique. All points within brackets (a–g or a–e) are statistically indistinguishable from each other but are significantly different (P < 0.05) from all other points on graph.

From Fitzgerald and Dehghani 142
Figure 6. Figure 6.

Effect of intravenous dopamine infusion on responses of carotid chemoreceptors to changes in . Each data point, mean of 2 measurements from 1 cat. Variability between measurements was small. Total count of 3 afferents is plotted against . •, ▴, Without dopamine; Δ, effect of dopamine at = 24.9 mmHg. Dopamine suppressed stimulation of chemoreceptors by hypoxia.

From Lahiri et al. 264
Figure 7. Figure 7.

Single aortic and carotid chemoreceptor steady‐state responses to changes in at constant level of . Recordings were simultaneous in each of 4 cats. It is apparent that in the same animal the response of aortic chemoreceptors to hypoxia or hypercapnia is less than that of carotid chemoreceptors.

From Lahiri et al. 262
Figure 8. Figure 8.

Effect of induced hypotension on aortic and carotid chemoreceptor activity. Top to bottom: tracheal ; arterial blood pressure (Psa); tracheal ; aortic body chemoreceptor activity, imp · s−1 and impulses; carotid body chemoreceptor activity, imp · s−1 and impulses.

Courtesy of S. Lahiri
Figure 9. Figure 9.

Variables in ventilatory responses to hypercapnia and hypoxia. Slope was derived from providing subjects with 7% CO2 in O2 for rebreathing. Ventilation was measured against rising . Isocapnic hypoxia was generated by manipulating gas mixtures. Variable A is from = + A/( −30), where is the inspired minute volume and is at high where extrapolated slope approaches zero; it represents shape of hyperbolic curve and therefore hypoxic response. The greater the response, the higher the A value 236. Height of bar equals mean value. Thiopental provoked parallel depression of hypoxic and hypercapnic responses. Halothane totally abolished response to hypoxia.

From Knill et al. 235
Figure 10. Figure 10.

Dose response (mean ± SE) of awake goats to intravenous bolus injections of dopamine at various inspired O2 levels (). Responses are expressed as ventilation ratio (VR30), which is minute ventilation () measured over 30‐s period after injection divided by control . Responses when was 0.14 and 1.0 were significantly lower than when was 0.21.

From Bisgard et al. 45
Figure 11. Figure 11.

Ventilatory response of anesthetized dog to 3 different stimulus profiles bilaterally delivered to carotid bodies from infusion apparatus. Left panel: response (VR, ventilation ratio) at given point during stimulus application divided by control level of ventilation to step decrease in (•, = 35.3 mmHg; ○, = 43.6 mmHg). Middle panel: VR and mean during ramp perfusion into common carotid arteries. Right panel: mean VR during 6‐s hypoxic pulse train perfusions into common carotid arteries (3 s at 32.5 mmHg and 3 s at 92.4 mmHg). These response patterns differ from patterns produced by same 3 techniques with hypercapnic blood.

From Dutton et al. 117
Figure 12. Figure 12.

Examples of carotid chemoreceptor fiber activity in cat showing little periodicity in its autocorrelogram but distinct rhythmicity when cross‐correlated with respiratory cycle reference pulse. Breathing mixture was air in A, 8% CO2 + 20% O2 + 72% N2 in B, and 10% O2 + 90% N2 in C. With hypercapnia, degree of variation of cross‐correlogram pattern decreases, whereas with hypoxia there is still significant respiratory modulation even though mean interspike interval is approximately that obtained with hypercapnia. Hence, when CO2 is steady but O2 is fluctuating, chemoreceptor discharge is steady, but when CO2 is fluctuating and O2 is steady, chemoreceptor discharge fluctuates.

From Gehrich and Moore 161
Figure 13. Figure 13.

Effect of injections of saline equilibrated with 100% CO2 given via catheter in root of aorta in 1 cat. A: small injection causes brief drop in pH of similar amplitude to spontaneous respiratory oscillations. Though sinus nerve discharge appears unaffected, tidal volume (inspiration upward) is increased for 1 breath. B: large injection causes drop in pH 4–5 times amplitude of pH oscillations and causes obvious burst of firing and increase in 1 inspiratory tidal volume. Vertical divisions are seconds.

From Band et al. 15
Figure 14. Figure 14.

Summary of data from 3 dogs showing effect on breathing rhythm of progressively removing afferent inputs. f, Breathing frequency; Te, expiratory duration that includes duration of expiratory airflow (1–3 s) and expiratory pause; Ti, duration of inspiration; W, wakefulness; S, slow‐wave sleep; S + VB, slow‐wave sleep and vagal blockade; S + VB + O2, slow‐wave sleep, vagal blockade, and 1 breath of 100% O2. Open columns, normal pH; stippled columns, during chronic metabolic alkalosis. For W, S, and S + VB, values are means ± 1 SE of 10–30 separate studies. Values in each study were based on means of 8–10 consecutive breaths. Values for S + VB + O2 are means ± 1 SE of 11–18 individual measurements of first respiratory cycle after inhalation of 100% O2. For S + VB + O2 during alkalosis, n = 4 for dog 1 and n = 1 for dog 2. Largest depression in f in sequence of maneuvers comes when sleeping animal had vagi blocked, whereas ventilation did not follow this pattern.

From Sullivan et al. 412
Figure 15. Figure 15.

Relationship between carotid perfusion pressure (which reflects degree of baroreceptor activity) and minute ventilation during stimulation of carotid chemoreceptors with 1 and 3 μg nicotine and no chemoreceptor stimulation. Slopes were calculated from data from 8 dogs with vagus nerves intact. Slopes during chemoreceptor stimulation are significantly different from slope with no chemoreceptor stimulation.

From Heistad et al. 193
Figure 16. Figure 16.

Mean (±SE) ventilatory responses (V, Vt, and changes in these variables) of 6 cats with left carotid sinus nerve sectioned and right carotid artery perfused with autologous blood from extracorporeal circuit with of 33 mmHg and of either 42.8 mmHg (•) or 21.4 mmHg (○); 3 different gas mixtures were presented for inspiration, creating 3 different values. P, significance of differences between observed responses at different levels of . Vt,hyp and Vhyp, Vt and ventilation during hypoxic perfusion of right carotid body; Vt,contr and Vcontr, Vt and ventilation during control conditions. At of 30.6 mmHg, stimulation of right carotid body with hypoxic blood ( of 42.8 mmHg) could increase Vt by 34% and V by 42%. However, when rose to 54.7 mmHg and central chemoreceptor afferent activity increased so that Vt increased from 18.4 to 35.9 ml/kg and V increased from 432.4 to 906.0 ml · kg−1 · min−1, then hypoxic perfusion of right carotid body could further increase Vt by only 8% and V by only 8%. This is considered a hypoadditive interaction of respiratory drives at level of integration of peripheral and central afferents.

From Giese et al. 165
Figure 17. Figure 17.

Response curves of ventilation, tidal volume, and frequency to increases in at decreasing levels of before and after bilateral coagulation of brain stem intermediate chemosensitive area S in 2 cats. Values for partial pressures are in kPa (1 kPa ≈ 7.5 mmHg). Data show that peripheral chemoreceptor drive can guarantee ventilation in absence of central chemosensitive drive, but interaction between central and peripheral chemoreceptor activity affects ventilation by greatly increasing slope of response to increases in , clearly a multiplicative interaction between peripheral and central chemosensitivities. Before coagulation, ventilatory response of cat to increasing on background of decreasing is not clearly multiplicative. One cat (bottom graph) seems to show multiplicative interaction between O2 and CO2 whereas the other cat (top graph) does not.

From Schlaefke et al. 393
Figure 18. Figure 18.

Effect of acute hypoxia on functional residual capacity (FRC) in intact rats, rats with vagal blockade, and rats with vagal and carotid body nerve blockade.

From Barer et al. 19
Figure 19. Figure 19.

Ventilatory response to NaCN (50 μg/kg iv) before (C) and after peripheral chemoreceptor denervation in 6 unanesthetized ponies. Limited but significant recovery of function was present 22 mo after denervation. Sectioning of aortic nerve nearly eliminated ventilatory response to NaCN. Data strongly suggest that aortic chemoreceptors become functional in time‐dependent manner after carotid body denervation.

From Bisgard et al. 44
Figure 20. Figure 20.

Ventilation per kilogram body wt as function of . Mean values (±SD) of 4 cats in which response curves were determined before (Δ) and after (∇) vagotomy and after subsequent sinus neurotomy (□). Slope was reduced 42.3% with no significant change in intercept.

From Berkenbosch et al. 31
Figure 21. Figure 21.

Cat under light pentobarbitone anesthesia. A: transient effects on Ti‐vs.‐Vt relationships showing typical hysteresis loop in response to step changes in inhaled CO2 concentration. B: Corresponding Ti‐vs.‐Te relationships. +, Response to rapid increase in from 3.8% (breathing air) to 8.0%; Δ, response back to breathing air with alveolar CO2 fraction () of 3.8%.

From Bradley et al. 62
Figure 22. Figure 22.

Relationship between inspiratory duration (Ti) and tidal volume (Vt) resulting from 3 types of respiratory drive in cat with intact vagus nerves, o, Rebreathing O2; x, rebreathing air; Δ, breathing 8% O2 in N2. In each case, cat was first hyperventilated to apnea. These findings are similar to those of Hey et al. 197 in humans. Differential changes in functional residual capacity (FRC) would likely provoke a different pattern in Ti‐vs.‐Vt responses to stimuli; in humans, FRC has been reported to increase with both hypoxia and hypercapnia 157; it has not been measured in cats during hypercapnia.

From Widdicombe and Winning 439
Figure 23. Figure 23.

Comparison of changes in activity of individual medullary and pontine respiratory units in response to equivalent changes in peak integrated phrenic activity resulting from carotid chemoreceptor stimulation and hyperoxic hypercapnia. For each graph, percentage alterations in number of spikes per respiratory cycle (f/cycle) achieved during hyperoxic hypercapnia are plotted vs. f/cycle changes obtained for same respiratory unit during carotid chemoreceptor stimulation. Stippled and cross‐hatched areas contain units for which f/cycle was unaltered or declined during carotid chemoreception stimulation and hypercapnia, respectively. Lines of identity are drawn in each panel. D‐I, dorsal respiratory nucleus inspiratory units; V‐I, ventral respiratory nucleus inspiratory units; V‐E, ventral respiratory nucleus expiratory units.

From St. John 383
Figure 24. Figure 24.

Effects of hypercapnia and hypoxia on relationship between minute phrenic activity (neural analogue of minute ventilation) and peak phrenic activity in nerve bursts (top panel), expiratory time (middle panel), and inspiratory time (bottom panel) in absence of breathing movements in representative animal with intact vagi. At any level of minute phrenic activity above base‐line level, peak phrenic activity and inspiratory and expiratory times were all less, and therefore respiratory frequency was greater during hypoxia than during hypercapnia.

From Ledlie et al. 272
Figure 25. Figure 25.

Effect of carotid chemoreceptor stimulation on cardiac sympathetic nerve discharge. At signal, 0.2 ml of NaCN solution (50 μ/ml) was injected through right thyroid artery in open‐chest cat anesthetized with urethan (250 mg/kg) and chloralose (30 mg/kg) and artificially ventilated. A: control; B: heart electrically paced, blood pressure kept artificially constant. Top trace to bottom trace: BP, arterial blood pressure; MBP, mean arterial pressure; HR, heart rate; CSN, integrated discharge activity of right cardiac sympathetic nerve; PhN, activity of right phrenic nerve.

From Montarolo et al. 324
Figure 26. Figure 26.

Effects in cat with intact vagi on sympathetic nerve activities of chemoreceptor stimulation by injection of lobeline (10 μg) into carotid sinus (CS) of either side. A and B: normal conditions. C and D: blood pressure kept constant by stabilization; blood volume (BV) is shown as grams moved from circulatory system to extracorporeal reservoir in maintaining blood pressure constant. BP, femoral artery pressure; HR, heart rate; L‐VNA (R‐VNA), integrated activity from left (right) sympathetic vertebral nerve; L‐CNA (R‐CNA), integrated activity from left (right) sympathetic cardiac nerve; CS‐chemoreceptors, carotid body. Selective stimulation of a given carotid body provokes inhibition of activity in ipsilateral cardiac nerve.

From Kollai et al. 240
Figure 27. Figure 27.

Changes in plasma ACTH and in Cortisol secretion rate at 10 and 20 min after exposure to hypoxia (mean ± SE). svHH, spontaneous ventilation and normocapnic hypoxic hypoxia; cvHH, controlled ventilation and normocapnic hypoxic hypoxia; cvCOH, controlled ventilation and normocapnic CO hypoxia; HH on COH, hypoxic hypoxia with elevated carboxyhemoglobin. Circles above bars, significant response (P < 0.05). Group I (control, n = 6) was maintained normoxic and normocapnic and showed no changes in any variables.

From Raff et al. 369


Figure 1.

Title pages of 3 earliest dissertations on the carotid body. Though Albrecht von Haller presided at first 2 presentations, actual presenters were Hartwig Taube (left panel) and Matthias Berckelmann (middle panel). Right panel: 1797 edition by Ernst Philipp Andersch of his father's earlier work that had been lost.

From Pick 356


Figure 2.

Gross structure and response of carotid body as presented by C. J. F. Heymans in his Nobel Lecture of December 12, 1945. He received the Nobel Prize in Physiology or Medicine in 1938 “for the discovery of the role played by the sinus and aortic mechanisms in the regulation of respiration.” Left panel: neural pathways from cardioaortic and carotid sinus zones. Right panel: pneumogram of chloralose‐anesthetized dog (A) and of blood pressure and cardiac rate (B). Time is in 3‐s intervals. Acetylcholine (0.1 mg) applied to chemoreceptors of glomus caroticum resulted in marked reflex hyperpnea and bradycardia.

From Heymans 198


Figure 3.

Single‐fiber chemoreceptor response to decreases in arterial O2 content in cat. Left panel: mean response (±SE) of 4 single carotid chemoreceptor fibers to decreases in O2 content of arterial blood (; vol %) achieved by either decreasing fraction of inspired O2 (hypoxic hypoxia) or by adding CO to the inspirate (CO hypoxia). Inset: response [impulses/s (IPS)] of single carotid chemoreceptor fiber to lowering of by adding CO. CTL, control; , partial pressure of O2 in arterial blood (mmHg); COHb, percent carboxyhemoglobin; , partial pressure of CO2 in arterial blood (mmHg); pHa, negative logarithm of H+ concentration in arterial blood; FBP, femoral arterial blood pressure (mmHg); A–D: blood samples taken at various times; RA, room air. To assure that fiber was not dead, cat was ventilated on 10% O2; fiber showed brisk increase in neural activity. Right panel: mean response of 3 single aortic chemoreceptor fibers to decreases in achieved as with carotid body responses. Though aortic body's response is larger in percent of control, control IPS is usually much lower for aortic fibers than for carotid fibers (cf. insets and Fig. 7). Inset: response of single aortic chemoreceptor fiber to lowering of by adding CO.

Courtesy of R. S. Fitzgerald and G. A. Dehghani


Figure 4.

Effects on single‐fiber activity of increases in at decreasing levels of . At low levels of , neural activity approaches zero regardless of value. On the assumption that normocapnia for cats is = 34–37 Torr, data show that response is linear as is lowered and that multiplicative interaction of O2 and CO2 still operates at these values.

From Lahiri and DeLaney 254


Figure 5.

Statistical analyses of responses of 10 recordings from carotid chemoreceptors (left panel) and 5 recordings from aortic chemoreceptors (right panel) to increases in at different levels of . The 3 levels (torr) for carotid responses were (bottom to top) >470, 105–115, and 54–58. The 4 levels (mmHg) for aortic responses were (bottom to top) >350, 93–97, 66–71, and 39–45. Responses were analyzed first by analysis of variance, and then individual means were grouped by Duncan new multiple‐range technique. All points within brackets (a–g or a–e) are statistically indistinguishable from each other but are significantly different (P < 0.05) from all other points on graph.

From Fitzgerald and Dehghani 142


Figure 6.

Effect of intravenous dopamine infusion on responses of carotid chemoreceptors to changes in . Each data point, mean of 2 measurements from 1 cat. Variability between measurements was small. Total count of 3 afferents is plotted against . •, ▴, Without dopamine; Δ, effect of dopamine at = 24.9 mmHg. Dopamine suppressed stimulation of chemoreceptors by hypoxia.

From Lahiri et al. 264


Figure 7.

Single aortic and carotid chemoreceptor steady‐state responses to changes in at constant level of . Recordings were simultaneous in each of 4 cats. It is apparent that in the same animal the response of aortic chemoreceptors to hypoxia or hypercapnia is less than that of carotid chemoreceptors.

From Lahiri et al. 262


Figure 8.

Effect of induced hypotension on aortic and carotid chemoreceptor activity. Top to bottom: tracheal ; arterial blood pressure (Psa); tracheal ; aortic body chemoreceptor activity, imp · s−1 and impulses; carotid body chemoreceptor activity, imp · s−1 and impulses.

Courtesy of S. Lahiri


Figure 9.

Variables in ventilatory responses to hypercapnia and hypoxia. Slope was derived from providing subjects with 7% CO2 in O2 for rebreathing. Ventilation was measured against rising . Isocapnic hypoxia was generated by manipulating gas mixtures. Variable A is from = + A/( −30), where is the inspired minute volume and is at high where extrapolated slope approaches zero; it represents shape of hyperbolic curve and therefore hypoxic response. The greater the response, the higher the A value 236. Height of bar equals mean value. Thiopental provoked parallel depression of hypoxic and hypercapnic responses. Halothane totally abolished response to hypoxia.

From Knill et al. 235


Figure 10.

Dose response (mean ± SE) of awake goats to intravenous bolus injections of dopamine at various inspired O2 levels (). Responses are expressed as ventilation ratio (VR30), which is minute ventilation () measured over 30‐s period after injection divided by control . Responses when was 0.14 and 1.0 were significantly lower than when was 0.21.

From Bisgard et al. 45


Figure 11.

Ventilatory response of anesthetized dog to 3 different stimulus profiles bilaterally delivered to carotid bodies from infusion apparatus. Left panel: response (VR, ventilation ratio) at given point during stimulus application divided by control level of ventilation to step decrease in (•, = 35.3 mmHg; ○, = 43.6 mmHg). Middle panel: VR and mean during ramp perfusion into common carotid arteries. Right panel: mean VR during 6‐s hypoxic pulse train perfusions into common carotid arteries (3 s at 32.5 mmHg and 3 s at 92.4 mmHg). These response patterns differ from patterns produced by same 3 techniques with hypercapnic blood.

From Dutton et al. 117


Figure 12.

Examples of carotid chemoreceptor fiber activity in cat showing little periodicity in its autocorrelogram but distinct rhythmicity when cross‐correlated with respiratory cycle reference pulse. Breathing mixture was air in A, 8% CO2 + 20% O2 + 72% N2 in B, and 10% O2 + 90% N2 in C. With hypercapnia, degree of variation of cross‐correlogram pattern decreases, whereas with hypoxia there is still significant respiratory modulation even though mean interspike interval is approximately that obtained with hypercapnia. Hence, when CO2 is steady but O2 is fluctuating, chemoreceptor discharge is steady, but when CO2 is fluctuating and O2 is steady, chemoreceptor discharge fluctuates.

From Gehrich and Moore 161


Figure 13.

Effect of injections of saline equilibrated with 100% CO2 given via catheter in root of aorta in 1 cat. A: small injection causes brief drop in pH of similar amplitude to spontaneous respiratory oscillations. Though sinus nerve discharge appears unaffected, tidal volume (inspiration upward) is increased for 1 breath. B: large injection causes drop in pH 4–5 times amplitude of pH oscillations and causes obvious burst of firing and increase in 1 inspiratory tidal volume. Vertical divisions are seconds.

From Band et al. 15


Figure 14.

Summary of data from 3 dogs showing effect on breathing rhythm of progressively removing afferent inputs. f, Breathing frequency; Te, expiratory duration that includes duration of expiratory airflow (1–3 s) and expiratory pause; Ti, duration of inspiration; W, wakefulness; S, slow‐wave sleep; S + VB, slow‐wave sleep and vagal blockade; S + VB + O2, slow‐wave sleep, vagal blockade, and 1 breath of 100% O2. Open columns, normal pH; stippled columns, during chronic metabolic alkalosis. For W, S, and S + VB, values are means ± 1 SE of 10–30 separate studies. Values in each study were based on means of 8–10 consecutive breaths. Values for S + VB + O2 are means ± 1 SE of 11–18 individual measurements of first respiratory cycle after inhalation of 100% O2. For S + VB + O2 during alkalosis, n = 4 for dog 1 and n = 1 for dog 2. Largest depression in f in sequence of maneuvers comes when sleeping animal had vagi blocked, whereas ventilation did not follow this pattern.

From Sullivan et al. 412


Figure 15.

Relationship between carotid perfusion pressure (which reflects degree of baroreceptor activity) and minute ventilation during stimulation of carotid chemoreceptors with 1 and 3 μg nicotine and no chemoreceptor stimulation. Slopes were calculated from data from 8 dogs with vagus nerves intact. Slopes during chemoreceptor stimulation are significantly different from slope with no chemoreceptor stimulation.

From Heistad et al. 193


Figure 16.

Mean (±SE) ventilatory responses (V, Vt, and changes in these variables) of 6 cats with left carotid sinus nerve sectioned and right carotid artery perfused with autologous blood from extracorporeal circuit with of 33 mmHg and of either 42.8 mmHg (•) or 21.4 mmHg (○); 3 different gas mixtures were presented for inspiration, creating 3 different values. P, significance of differences between observed responses at different levels of . Vt,hyp and Vhyp, Vt and ventilation during hypoxic perfusion of right carotid body; Vt,contr and Vcontr, Vt and ventilation during control conditions. At of 30.6 mmHg, stimulation of right carotid body with hypoxic blood ( of 42.8 mmHg) could increase Vt by 34% and V by 42%. However, when rose to 54.7 mmHg and central chemoreceptor afferent activity increased so that Vt increased from 18.4 to 35.9 ml/kg and V increased from 432.4 to 906.0 ml · kg−1 · min−1, then hypoxic perfusion of right carotid body could further increase Vt by only 8% and V by only 8%. This is considered a hypoadditive interaction of respiratory drives at level of integration of peripheral and central afferents.

From Giese et al. 165


Figure 17.

Response curves of ventilation, tidal volume, and frequency to increases in at decreasing levels of before and after bilateral coagulation of brain stem intermediate chemosensitive area S in 2 cats. Values for partial pressures are in kPa (1 kPa ≈ 7.5 mmHg). Data show that peripheral chemoreceptor drive can guarantee ventilation in absence of central chemosensitive drive, but interaction between central and peripheral chemoreceptor activity affects ventilation by greatly increasing slope of response to increases in , clearly a multiplicative interaction between peripheral and central chemosensitivities. Before coagulation, ventilatory response of cat to increasing on background of decreasing is not clearly multiplicative. One cat (bottom graph) seems to show multiplicative interaction between O2 and CO2 whereas the other cat (top graph) does not.

From Schlaefke et al. 393


Figure 18.

Effect of acute hypoxia on functional residual capacity (FRC) in intact rats, rats with vagal blockade, and rats with vagal and carotid body nerve blockade.

From Barer et al. 19


Figure 19.

Ventilatory response to NaCN (50 μg/kg iv) before (C) and after peripheral chemoreceptor denervation in 6 unanesthetized ponies. Limited but significant recovery of function was present 22 mo after denervation. Sectioning of aortic nerve nearly eliminated ventilatory response to NaCN. Data strongly suggest that aortic chemoreceptors become functional in time‐dependent manner after carotid body denervation.

From Bisgard et al. 44


Figure 20.

Ventilation per kilogram body wt as function of . Mean values (±SD) of 4 cats in which response curves were determined before (Δ) and after (∇) vagotomy and after subsequent sinus neurotomy (□). Slope was reduced 42.3% with no significant change in intercept.

From Berkenbosch et al. 31


Figure 21.

Cat under light pentobarbitone anesthesia. A: transient effects on Ti‐vs.‐Vt relationships showing typical hysteresis loop in response to step changes in inhaled CO2 concentration. B: Corresponding Ti‐vs.‐Te relationships. +, Response to rapid increase in from 3.8% (breathing air) to 8.0%; Δ, response back to breathing air with alveolar CO2 fraction () of 3.8%.

From Bradley et al. 62


Figure 22.

Relationship between inspiratory duration (Ti) and tidal volume (Vt) resulting from 3 types of respiratory drive in cat with intact vagus nerves, o, Rebreathing O2; x, rebreathing air; Δ, breathing 8% O2 in N2. In each case, cat was first hyperventilated to apnea. These findings are similar to those of Hey et al. 197 in humans. Differential changes in functional residual capacity (FRC) would likely provoke a different pattern in Ti‐vs.‐Vt responses to stimuli; in humans, FRC has been reported to increase with both hypoxia and hypercapnia 157; it has not been measured in cats during hypercapnia.

From Widdicombe and Winning 439


Figure 23.

Comparison of changes in activity of individual medullary and pontine respiratory units in response to equivalent changes in peak integrated phrenic activity resulting from carotid chemoreceptor stimulation and hyperoxic hypercapnia. For each graph, percentage alterations in number of spikes per respiratory cycle (f/cycle) achieved during hyperoxic hypercapnia are plotted vs. f/cycle changes obtained for same respiratory unit during carotid chemoreceptor stimulation. Stippled and cross‐hatched areas contain units for which f/cycle was unaltered or declined during carotid chemoreception stimulation and hypercapnia, respectively. Lines of identity are drawn in each panel. D‐I, dorsal respiratory nucleus inspiratory units; V‐I, ventral respiratory nucleus inspiratory units; V‐E, ventral respiratory nucleus expiratory units.

From St. John 383


Figure 24.

Effects of hypercapnia and hypoxia on relationship between minute phrenic activity (neural analogue of minute ventilation) and peak phrenic activity in nerve bursts (top panel), expiratory time (middle panel), and inspiratory time (bottom panel) in absence of breathing movements in representative animal with intact vagi. At any level of minute phrenic activity above base‐line level, peak phrenic activity and inspiratory and expiratory times were all less, and therefore respiratory frequency was greater during hypoxia than during hypercapnia.

From Ledlie et al. 272


Figure 25.

Effect of carotid chemoreceptor stimulation on cardiac sympathetic nerve discharge. At signal, 0.2 ml of NaCN solution (50 μ/ml) was injected through right thyroid artery in open‐chest cat anesthetized with urethan (250 mg/kg) and chloralose (30 mg/kg) and artificially ventilated. A: control; B: heart electrically paced, blood pressure kept artificially constant. Top trace to bottom trace: BP, arterial blood pressure; MBP, mean arterial pressure; HR, heart rate; CSN, integrated discharge activity of right cardiac sympathetic nerve; PhN, activity of right phrenic nerve.

From Montarolo et al. 324


Figure 26.

Effects in cat with intact vagi on sympathetic nerve activities of chemoreceptor stimulation by injection of lobeline (10 μg) into carotid sinus (CS) of either side. A and B: normal conditions. C and D: blood pressure kept constant by stabilization; blood volume (BV) is shown as grams moved from circulatory system to extracorporeal reservoir in maintaining blood pressure constant. BP, femoral artery pressure; HR, heart rate; L‐VNA (R‐VNA), integrated activity from left (right) sympathetic vertebral nerve; L‐CNA (R‐CNA), integrated activity from left (right) sympathetic cardiac nerve; CS‐chemoreceptors, carotid body. Selective stimulation of a given carotid body provokes inhibition of activity in ipsilateral cardiac nerve.

From Kollai et al. 240


Figure 27.

Changes in plasma ACTH and in Cortisol secretion rate at 10 and 20 min after exposure to hypoxia (mean ± SE). svHH, spontaneous ventilation and normocapnic hypoxic hypoxia; cvHH, controlled ventilation and normocapnic hypoxic hypoxia; cvCOH, controlled ventilation and normocapnic CO hypoxia; HH on COH, hypoxic hypoxia with elevated carboxyhemoglobin. Circles above bars, significant response (P < 0.05). Group I (control, n = 6) was maintained normoxic and normocapnic and showed no changes in any variables.

From Raff et al. 369
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Article Title: Reflex Responses to Chemoreceptor Stimulation
 

How to Cite

Robert S. Fitzgerald, Sukhamay Lahiri. Reflex Responses to Chemoreceptor Stimulation. Compr Physiol 2011, Supplement 11: Handbook of Physiology, The Respiratory System, Control of Breathing: 313-362. First published in print 1986. doi: 10.1002/cphy.cp030210