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Arterial Chemoreceptors

Carlos Eyzaguirre, Robert S. Fitzgerald, Sukhamay Lahiri, Patricio Zapata

10.1002/cphy.cp020316

Source: Supplement 8: Handbook of Physiology, The Cardiovascular System, Peripheral Circulation and Organ Blood Flow

Originally published: 1983

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Anatomical Features
2 Physiological Characteristics
2.1 Arterial Chemoreceptors in the Fetus and Newborn
2.2 Peripheral Chemoreceptors in Adult Human Beings and Animals
2.3 Adaptation of Response to CO2
2.4 Oscillations of Chemoreceptor Activity
2.5 Effects of Exercise
2.6 Blood Pressure, Perfusate Flow, and Carotid Chemoreceptor Activity
2.7 Oxygen Consumption and Tissue PO2
2.8 Oxygen Content and Po2 in Chemoreceptor Excitation
2.9 Anesthetics and Peripheral Chemoreceptor Activity
2.10 Effects of Temperature and Tonicity
2.11 Pathophysiology of Peripheral Chemoreceptors
3 Cellular Electrophysiological Mechanisms
3.1 Glomus Cells
4 Possible Role of Carotid Body Cells as Conditioners or Inductors of Chemosensory Activity
5 Chemosensory Nerve Endings
5.1 Mass Receptor Potential of Chemoreceptors
5.2 Recordings From Single Nerve Fibers and Endings
6 Hypotheses of Chemoreception
6.1 Hypotheses Involving Neurotransmitters
6.2 Mechanoreceptor Hypotheses
6.3 Hypotheses Based on pH Sensitivity
6.4 Protein Receptor Hypothesis
6.5 Metabolic Hypothesis
7 Interactions Between Nerves and Carotid Body Cells
7.1 Result of Efferent Input on Carotid Body Afferent Output
7.2 Physiological Significance of Carotid Nerve Efferent Input
8 Peripheral Chemoreceptors and Cardiovascular Control
8.1 Chemoreceptor Stimulation in the Normoxic Animal
8.2 Perfusion of Isolated Aortic and Carotid Chemoreceptors with Hypoxic and Hypoxic‐Hypercapnic Blood
8.3 Cardiovascular Adjustments to Acute Systemic Hypoxia
8.4 Chemoreceptor Activity and Regional Blood Flow
9 Conclusions
Figure 1. Figure 1.

Synaptic connections in rat carotid body. Note membrane densifications at different synaptic sites. Heavier densification (δ) is generally considered to be located in presynaptic side of junction. Glomus cells have dense‐core vesicles and are interconnected by synapses. Afferent nerve endings apposed to cells have relatively large synaptic vesicles and few large dense‐core vesicles. Some regions of afferent nerve endings are presynaptic to cells, some postsynaptic, and some form reciprocal synapses. Where cell bodies are presynaptic, large dense‐core vesicles and synaptic vesicles accumulate in equal concentration near synaptic junction, but synaptic vesicles predominate where cell processes are presynaptic.

Adapted from McDonald and Mitchell 412
Figure 2. Figure 2.

Effect of occluding the umbilical cord (for period indicated by bar on time marker) on aortic chemoreceptor activity in sheep fetal preparation. Neurograms A and B obtained when indicated on rate‐meter trace.

From Ponte and Purves 495
Figure 3. Figure 3.

Isometric diagram. Changes in carotid nerve impulses of cat during inhalation of different O2 mixtures at various ventilation levels. X axis, alveolar CO2 pressure (PCO2); Y axis, concentration of inhaled oxygen; Z axis, frequency of carotid nerve impulses; discharges mainly from chemoreceptors. Mean arterial pressure 140–170 Torr throughout series: O2‐impulse curves obtained during inhalation of O2 (100%‐10%) at different PCO2 levels; CO2‐impulse curves obtained at different PCO2 levels during inhalation of 100% O2 and of 50%, 20%, and 10% O2 in N2.

From Eyzaguirre and Lewin 176
Figure 4. Figure 4.

Chemoreceptor sensitivity to hypoxia. Effect of arterial O2 pressure (PaO2) levels on slopes of CO2‐response (mean ± SE) curves of aortic (▪) and carotid (•) cat chemoreceptors (n = 15).

From Lahiri et al. 363
Figure 5. Figure 5.

Response (mean ± SE) of carotid and aortic bodies to increasing levels of PaCO2 at decreasing levels of PaO2 (bottom to top). Carotid sinus nerve (n = 10) and aortic depressor nerve (n = 6) single‐ or few‐fiber preparations from the cat. ×, Hyperoxia; ⋄, normoxia; •, mild hypoxia; ○, moderate hypoxia.

Adapted from Fitzgerald and Dehghani 196
Figure 6. Figure 6.

Responses of cat carotid chemoreceptor fibers to changes in end‐tidal CO2 pressure (PETCO2) during hyperoxia (A) and hypoxia (B). Increases in PETCO2 were followed by increases in discharge rates during both hyperoxia and hypoxia. PSA, systemic arterial pressure.

From Lahiri et al. (ref. 364 and unpublished observations)
Figure 7. Figure 7.

Aortic (Δ) and carotid (•) chemoreceptor responses of cat to carboxyhemoglobinemia during normoxia (PaO2 = 82–98 Torr; mean ± SE; n = 10). Only aortic chemoreceptors were stimulated.

From Lahiri et al. 367
Figure 8. Figure 8.

Arrhenius plots for single (different) chemoreceptor fibers of cat carotid body in vitro under different conditions. A: line 2, regression under 50% O2 in N2; line 1, under 100% O2; line 3, under air. B: line 1, under 50% O2 in N2, pH 7.43; line 2, under 6% CO2) 50% O2, and 44% N2, pH 7.43; line 3, under 6% CO2, 50% O2, and 44% N2, pH 6. C: line 1, under 50% O2 in N2, pH 7.43; line 2, under 6% CO2, 50% O2, and 44% N2, pH 6; line 3, after 50% O2 in N2, pH 6. D: under 100% O2; line 2, pH 7.43; line 3, pH 7.0; line 1, pH 7.8. Ordinates, impulses per second; abscissae, 105 × reciprocal of absolute temperature (T−l).

From Gallego et al. 211
Figure 9. Figure 9.

Results obtained with solutions of different osmolalities on single unit discharge in separate experiments on cat carotid bodies in vitro. Ordinate: mean change in discharge frequency (ΔF) ± SEM; +, increase; −, decrease. Abscissa: changes in osmolality (Δ mOsm); 304 mOsm, mean control osmolality. Bars, ΔF; vertical lines, SE; *P < 0.05; circled numbers above and below bars, number of observations. Right, changes during application of hyperosmotic solutions at constant [Na+]o = 154 mM; left, changes in frequency induced by hyposomotic solutions at constant [Na+]o = 112 mM.

From Gallego et al. 211
Figure 10. Figure 10.

Effect of drop in temperature from 37°C to 32°C on membrane potential (MP) and input resistance (Ro) of single glomus cell and on sensory discharge frequency of whole carotid nerve. Cat carotid body preparation in vitro. Upper trace, intracellular recording; middle trace, sensory discharge frequency; lower trace, temperature.

From Baron and Eyzaguirre 32
Figure 11. Figure 11.

Reversal potential of cooling effect on cat carotid body in vitro. A: intracellular recording of MP and Ro before, during, and after cooling at normal resting potential. B: MP artificially displaced toward positive values by injecting direct outward current (0.4 nA) through recording electrode. Note cell repolarization and loss of Ro during cooling. C: cell artificially hyperpolarized by injecting direct inward current (0.1 nA) through micropipette. Note larger depolarization during cooling. In all 3 cases, potential of cell tends to reach same level. D: temperature record.

From Baron and Eyzaguirre 32
Figure 12. Figure 12.

Effects of solutions of different osmolalities on MP and Ro of glomus cells of cat carotid body in vitro. Right, hyperosmotic solutions, [Na+]o = 154 mM; left, hyposmotic solutions, [Na+]o = 112 mM. Open bars, changes in MP; shaded bars, changes in Ro; *P < 0.05; circled numbers above bars, number of observations. Left ordinate (MP): +, depolarization; −, hyperpolarization. Right ordinate (Ro): +, increase; −, decrease. Abscissa: changes in osmolality (± mOsm); 302 mOsm, mean control osmolarity.

From Gallego et al. 211
Figure 13. Figure 13.

Effects of acetylcholine (ACh; 50 μg) on glomus cell of cat carotid body in vitro impaled with micropipette filled with 6% Procion yellow. Cell identified after ejecting dye from pipette. Upper trace, ΔMP induced by drug; middle trace, lower trace, high‐gain AC recording of voltage noise from same cell. 1, Base‐line noise (before ACh); 2, noise recorded near peak of drug‐induced depolarization.

From Hayashida and Eyzaguirre 256
Figure 14. Figure 14.

Effect of asphyxia and N2 inhalation on sensory discharges recorded from superior laryngeal nerve (SLN) filament innervating cat carotid body; carotid nerve anastomosed to SLN 131 days prior to experiment. A : control discharge during spontaneous inhalation of room air. B : discharge during peak of asphyxic effect elicited by tracheal occlusion. C: discharge several seconds after B and during inhalation of room air. D: discharge during peak of effect induced by inhalation of 100% N2. Bottom: frequency changes induced by asphyxia and 100% N2. Effects elicited between arrows.

From Zapata et al. 594
Figure 15. Figure 15.

Dose‐peak‐response curves constructed after intravenous injections of different doses of NaCN and nicotine. Bilateral carotid nerve recordings from cat. A, B: carotid nerve crushed 6 days before. ▪, Responses of nerve crushed close to glomus; ▪, responses of nerve crushed far from glomus. C, D: different experiment. •, Response of normal nerve; ○, response of nerve crushed in its middle 6 days before.

From Zapata et al. 597
Figure 16. Figure 16.

Labeled carotid nerve terminal of cat carotid body examined with ultrastructural autoradiography 6 days after treating petrosal ganglion with [3H] proline. Note silver grains over nerve ending after radioactive material was transported through sensory nerves.

Courtesy of S. J. Fidone, P. Zapata, and L. J. Stensaas (see also ref. 194)
Figure 17. Figure 17.

Local nature of slow negative (mass receptor) potential induced by intra‐arterial injections of ACh in cat. Upper traces, recording from nerve; lower traces, sensory discharge frequency. Injections made at arrows. A: slow potential elicited by intra‐arterial injection of 5 μg of ACh, recorded from filament of carotid nerve with proximal electrode placed at entrance of nerve into glomus. B: proximal electrode at 0.7 mm from glomus; distal electrode remained stationary near cut end of nerve.

From Eyzaguirre et al. 182
Figure 18. Figure 18.

Impalement of single nerve ending yields spontaneous depolarizing potentials (SDPs) that (if large enough) seem to evoke sensory discharges in cat carotid body in vitro. Smaller SDPs do not appear to give rise to action potentials. Terminal was invaded by spikes originating elsewhere. Nerve ending identified by intracellular staining after recording.

From Hayashida et al. 257
Figure 19. Figure 19.

Effects of physostigmine (eserine) and mecamylamine on Loewi‐type effect in cat carotid bodies in vitro. Inset, experimental situation. Locke solution equilibrated with 50% O2 in N2, pH 7.45 at 35°C, flowing at 0.6 ml/min under paraffin oil. Donor carotid body (1) is upstream and separated from downstream detector carotid body preparation (2) by 17 mm. Direction of flow indicated by horizontal arrows. Electrical current (60 μA DC) applied for 60 s to carotid body 1 (vertical arrows) and sensory discharges recorded from carotid nerve of preparation 2. A : preparations bathed in normal Locke solution. B: preparations bathed with physostigmine‐Locke solution (10−6 g/ml eserine salicylate) for 60 min. C: mecamylamine HCl (10−4 g/ml) added to physostigmine‐Locke solution for 60 min. Each point shows mean frequency recorded during 60 s.

From Eyzaguirre and Zapata 186
Figure 20. Figure 20.

Catecholamine biosynthesis in rat carotid body. Circles, pools of tyrosine (TYR), dihydroxyphenylalanine (DOPA), dopamine (DA), norepinephrine (NA), and epinephrine (AD). Arrows, enzymes involved: tyrosine hydroxylase (TH), aromatic L‐amino acid decarboxylase (AAAD), dopamine β‐hydroxylase (DBH), and phenylethanolamine N‐methyltransferase (PNMT); widths indicate activities of enzymes. Columns: immunofluorescence intensity in glomus cells, enzymatic activity, contents, and changes induced by hypoxia.

Data from Bolme et al. 71, Hanbauer et al. 247, and Hellström, and co‐workers 265,266
Figure 21. Figure 21.

Dopamine (DA) and norepinephrine (NA) contents in rat carotid body. Areas of large circles, proportional content detected under basal conditions. Upward and downward arrows, increase or decrease in content produced by different conditions. Thin horizontal arrows, moderate but statistically significant effects. Thick horizontal arrows, pronounced effects. Small circles, no effects. Responses to a precursor (L‐dopa), a monoamine oxidase (MAO) inhibitor (pargyline), 1 wk of carotid neurotomy, chronic treatment with a glucocorticoid (dexamethasone) and a mineralocorticoid (doca), 15–60 min of hypoxia, 2‐h administration of reserpine and a DBH inhibitor [diethyldithiocarbamate (DDC)], 1 wk of sympathectomy, and chronic administration of 6‐hydroxydopamine (6‐OHDA).

Data from Hanbauer and Hellström 245, Hellström et al. 264, and Hellström and Kaslow 266
Figure 22. Figure 22.

Dose‐response curves of cat carotid chemoreceptors to haloperidol at various levels of PaO2. Haloperidol had little effect during hyperoxia when receptor activity was slight. At lower PaO2 levels, as activity increased, haloperidol had an augmenting effect. At all levels of PaO2, saturation dose of haloperidol appeared to be 1 mg/kg. At each dose of haloperidol, responses to steady‐state PaO2 were obtained systematically. Each data point represents 1 measurement.

From Lahiri et al. 370
Figure 23. Figure 23.

Effects of close intra‐arterial injections of Met‐enkephalin before (A) and after (B) naloxone on cat carotid nerve discharges. Met‐enkephalin inhibited chemoreceptor activity, and this effect was blocked by naloxone. PSA, systemic arterial pressure.

From Pokorski and Lahiri 494
Figure 24. Figure 24.

Inhibition of cat carotid chemoreceptor response to PaO2 by oligomycin. Carotid chemoreceptor steady‐state responses to PaO2 at constant PaCO2 in 8 experiments before (A) and after (B) oligomycin (50–500 μg ia). Open symbols in B correspond to closed symbols in A. Chemoreceptor activity is insensitive to PaO2 level after oligomycin.

From Mulligan et al. 449
Figure 25. Figure 25.

Responses of carotid chemoreceptor afferent of cat to similar changes in PETCO2, before (A, C) and after (B, D) oligomycin (200 μg ia) during hyperoxia (PaO2 > 400 Torr). After oligomycin there were appreciable overshoots and undershoots in activity.

From Mulligan et al. 448
Figure 26. Figure 26.

Temporal separation of aortic and carotid body stimulation in the dog. A: nicotine injected through catheter placed in aorta (just beyond aortic valves) reaches aortic bodies within 1 s; coils of plastic tubing inserted in common carotids delay nicotine from reaching carotid bodies for 75 s. B: nicotine injected at 2 stimulates aortic bodies and causes tachycardia and hypertension. Neuromuscular blocking agent (succinylcholine) injected at 1 produces apnea and eliminates effects of hyperventilation. Top tracing, respiratory air flow; bottom tracing, carotid blood pressure. At 3, 45 s was deleted to save space; A.A., ascending aorta.

From Comroe and Mortimer 99
Figure 27. Figure 27.

Effects of carotid chemoreceptor stimulation on phasic and mean arterial pressure, left ventricular (LV) pressure, dP/dt, LV diameter, respiration (monitored by pneumograph), and heart rate in dog with spontaneous respiration (left panel). Heart rate remained constant, but chemoreceptor stimulation markedly increased respiration with increase in aortic and LV pressures and in dP/dt. With ventilation controlled (right panel), same carotid chemoreceptor stimulus induced larger increase in pressures and dP/dt.

From Vatner and Rutherford 554
Figure 28. Figure 28.

Hemodynamics during hypoxic hypoxia (HH) and CO hypoxia (COH) before and after carotid body resection (CBR) in the dog. A: Cao2, arterial oxygen content; Pa, mean arterial pressure; Q, cardiac output; TPR, total peripheral resistance. B: HR, heart rate; SV, stroke volume; PLA, mean left atrial pressure; SW/PLA, stroke work. Bars, ±; SD.

From Sylvester et al. 544
Figure 29. Figure 29.

Effect of arterial hypoxia on systemic hemodynamics in conscious dogs. C, control room‐air breathing; H, hypoxia; brackets, ± SEM.

From Krasney and Koehler 346
Figure 30. Figure 30.

Cardiac responses to arterial hypoxia in conscious dogs. C, control room‐air breathing; H, hypoxia; brackets, ±; SEM.

From Krasney and Koehler 346
Figure 31. Figure 31.

Responses to aortic injection of cyanide (CN) before (left) and after (right) acute carotid denervation and bilateral vagotomy in dog. Changes in perfusion pressure (PP) in gracilis muscle and paw were abolished, indicating that reflex responses in muscle and paw were triggered by afferent impulses from carotid sinus area and aortic arch. Changes in systemic arterial pressure (SAP) were small and persisted with some modification after denervation.

From Calvelo et al. 79, by permission of the American Heart Association, Inc


Figure 1.

Synaptic connections in rat carotid body. Note membrane densifications at different synaptic sites. Heavier densification (δ) is generally considered to be located in presynaptic side of junction. Glomus cells have dense‐core vesicles and are interconnected by synapses. Afferent nerve endings apposed to cells have relatively large synaptic vesicles and few large dense‐core vesicles. Some regions of afferent nerve endings are presynaptic to cells, some postsynaptic, and some form reciprocal synapses. Where cell bodies are presynaptic, large dense‐core vesicles and synaptic vesicles accumulate in equal concentration near synaptic junction, but synaptic vesicles predominate where cell processes are presynaptic.

Adapted from McDonald and Mitchell 412


Figure 2.

Effect of occluding the umbilical cord (for period indicated by bar on time marker) on aortic chemoreceptor activity in sheep fetal preparation. Neurograms A and B obtained when indicated on rate‐meter trace.

From Ponte and Purves 495


Figure 3.

Isometric diagram. Changes in carotid nerve impulses of cat during inhalation of different O2 mixtures at various ventilation levels. X axis, alveolar CO2 pressure (PCO2); Y axis, concentration of inhaled oxygen; Z axis, frequency of carotid nerve impulses; discharges mainly from chemoreceptors. Mean arterial pressure 140–170 Torr throughout series: O2‐impulse curves obtained during inhalation of O2 (100%‐10%) at different PCO2 levels; CO2‐impulse curves obtained at different PCO2 levels during inhalation of 100% O2 and of 50%, 20%, and 10% O2 in N2.

From Eyzaguirre and Lewin 176


Figure 4.

Chemoreceptor sensitivity to hypoxia. Effect of arterial O2 pressure (PaO2) levels on slopes of CO2‐response (mean ± SE) curves of aortic (▪) and carotid (•) cat chemoreceptors (n = 15).

From Lahiri et al. 363


Figure 5.

Response (mean ± SE) of carotid and aortic bodies to increasing levels of PaCO2 at decreasing levels of PaO2 (bottom to top). Carotid sinus nerve (n = 10) and aortic depressor nerve (n = 6) single‐ or few‐fiber preparations from the cat. ×, Hyperoxia; ⋄, normoxia; •, mild hypoxia; ○, moderate hypoxia.

Adapted from Fitzgerald and Dehghani 196


Figure 6.

Responses of cat carotid chemoreceptor fibers to changes in end‐tidal CO2 pressure (PETCO2) during hyperoxia (A) and hypoxia (B). Increases in PETCO2 were followed by increases in discharge rates during both hyperoxia and hypoxia. PSA, systemic arterial pressure.

From Lahiri et al. (ref. 364 and unpublished observations)


Figure 7.

Aortic (Δ) and carotid (•) chemoreceptor responses of cat to carboxyhemoglobinemia during normoxia (PaO2 = 82–98 Torr; mean ± SE; n = 10). Only aortic chemoreceptors were stimulated.

From Lahiri et al. 367


Figure 8.

Arrhenius plots for single (different) chemoreceptor fibers of cat carotid body in vitro under different conditions. A: line 2, regression under 50% O2 in N2; line 1, under 100% O2; line 3, under air. B: line 1, under 50% O2 in N2, pH 7.43; line 2, under 6% CO2) 50% O2, and 44% N2, pH 7.43; line 3, under 6% CO2, 50% O2, and 44% N2, pH 6. C: line 1, under 50% O2 in N2, pH 7.43; line 2, under 6% CO2, 50% O2, and 44% N2, pH 6; line 3, after 50% O2 in N2, pH 6. D: under 100% O2; line 2, pH 7.43; line 3, pH 7.0; line 1, pH 7.8. Ordinates, impulses per second; abscissae, 105 × reciprocal of absolute temperature (T−l).

From Gallego et al. 211


Figure 9.

Results obtained with solutions of different osmolalities on single unit discharge in separate experiments on cat carotid bodies in vitro. Ordinate: mean change in discharge frequency (ΔF) ± SEM; +, increase; −, decrease. Abscissa: changes in osmolality (Δ mOsm); 304 mOsm, mean control osmolality. Bars, ΔF; vertical lines, SE; *P < 0.05; circled numbers above and below bars, number of observations. Right, changes during application of hyperosmotic solutions at constant [Na+]o = 154 mM; left, changes in frequency induced by hyposomotic solutions at constant [Na+]o = 112 mM.

From Gallego et al. 211


Figure 10.

Effect of drop in temperature from 37°C to 32°C on membrane potential (MP) and input resistance (Ro) of single glomus cell and on sensory discharge frequency of whole carotid nerve. Cat carotid body preparation in vitro. Upper trace, intracellular recording; middle trace, sensory discharge frequency; lower trace, temperature.

From Baron and Eyzaguirre 32


Figure 11.

Reversal potential of cooling effect on cat carotid body in vitro. A: intracellular recording of MP and Ro before, during, and after cooling at normal resting potential. B: MP artificially displaced toward positive values by injecting direct outward current (0.4 nA) through recording electrode. Note cell repolarization and loss of Ro during cooling. C: cell artificially hyperpolarized by injecting direct inward current (0.1 nA) through micropipette. Note larger depolarization during cooling. In all 3 cases, potential of cell tends to reach same level. D: temperature record.

From Baron and Eyzaguirre 32


Figure 12.

Effects of solutions of different osmolalities on MP and Ro of glomus cells of cat carotid body in vitro. Right, hyperosmotic solutions, [Na+]o = 154 mM; left, hyposmotic solutions, [Na+]o = 112 mM. Open bars, changes in MP; shaded bars, changes in Ro; *P < 0.05; circled numbers above bars, number of observations. Left ordinate (MP): +, depolarization; −, hyperpolarization. Right ordinate (Ro): +, increase; −, decrease. Abscissa: changes in osmolality (± mOsm); 302 mOsm, mean control osmolarity.

From Gallego et al. 211


Figure 13.

Effects of acetylcholine (ACh; 50 μg) on glomus cell of cat carotid body in vitro impaled with micropipette filled with 6% Procion yellow. Cell identified after ejecting dye from pipette. Upper trace, ΔMP induced by drug; middle trace, lower trace, high‐gain AC recording of voltage noise from same cell. 1, Base‐line noise (before ACh); 2, noise recorded near peak of drug‐induced depolarization.

From Hayashida and Eyzaguirre 256


Figure 14.

Effect of asphyxia and N2 inhalation on sensory discharges recorded from superior laryngeal nerve (SLN) filament innervating cat carotid body; carotid nerve anastomosed to SLN 131 days prior to experiment. A : control discharge during spontaneous inhalation of room air. B : discharge during peak of asphyxic effect elicited by tracheal occlusion. C: discharge several seconds after B and during inhalation of room air. D: discharge during peak of effect induced by inhalation of 100% N2. Bottom: frequency changes induced by asphyxia and 100% N2. Effects elicited between arrows.

From Zapata et al. 594


Figure 15.

Dose‐peak‐response curves constructed after intravenous injections of different doses of NaCN and nicotine. Bilateral carotid nerve recordings from cat. A, B: carotid nerve crushed 6 days before. ▪, Responses of nerve crushed close to glomus; ▪, responses of nerve crushed far from glomus. C, D: different experiment. •, Response of normal nerve; ○, response of nerve crushed in its middle 6 days before.

From Zapata et al. 597


Figure 16.

Labeled carotid nerve terminal of cat carotid body examined with ultrastructural autoradiography 6 days after treating petrosal ganglion with [3H] proline. Note silver grains over nerve ending after radioactive material was transported through sensory nerves.

Courtesy of S. J. Fidone, P. Zapata, and L. J. Stensaas (see also ref. 194)


Figure 17.

Local nature of slow negative (mass receptor) potential induced by intra‐arterial injections of ACh in cat. Upper traces, recording from nerve; lower traces, sensory discharge frequency. Injections made at arrows. A: slow potential elicited by intra‐arterial injection of 5 μg of ACh, recorded from filament of carotid nerve with proximal electrode placed at entrance of nerve into glomus. B: proximal electrode at 0.7 mm from glomus; distal electrode remained stationary near cut end of nerve.

From Eyzaguirre et al. 182


Figure 18.

Impalement of single nerve ending yields spontaneous depolarizing potentials (SDPs) that (if large enough) seem to evoke sensory discharges in cat carotid body in vitro. Smaller SDPs do not appear to give rise to action potentials. Terminal was invaded by spikes originating elsewhere. Nerve ending identified by intracellular staining after recording.

From Hayashida et al. 257


Figure 19.

Effects of physostigmine (eserine) and mecamylamine on Loewi‐type effect in cat carotid bodies in vitro. Inset, experimental situation. Locke solution equilibrated with 50% O2 in N2, pH 7.45 at 35°C, flowing at 0.6 ml/min under paraffin oil. Donor carotid body (1) is upstream and separated from downstream detector carotid body preparation (2) by 17 mm. Direction of flow indicated by horizontal arrows. Electrical current (60 μA DC) applied for 60 s to carotid body 1 (vertical arrows) and sensory discharges recorded from carotid nerve of preparation 2. A : preparations bathed in normal Locke solution. B: preparations bathed with physostigmine‐Locke solution (10−6 g/ml eserine salicylate) for 60 min. C: mecamylamine HCl (10−4 g/ml) added to physostigmine‐Locke solution for 60 min. Each point shows mean frequency recorded during 60 s.

From Eyzaguirre and Zapata 186


Figure 20.

Catecholamine biosynthesis in rat carotid body. Circles, pools of tyrosine (TYR), dihydroxyphenylalanine (DOPA), dopamine (DA), norepinephrine (NA), and epinephrine (AD). Arrows, enzymes involved: tyrosine hydroxylase (TH), aromatic L‐amino acid decarboxylase (AAAD), dopamine β‐hydroxylase (DBH), and phenylethanolamine N‐methyltransferase (PNMT); widths indicate activities of enzymes. Columns: immunofluorescence intensity in glomus cells, enzymatic activity, contents, and changes induced by hypoxia.

Data from Bolme et al. 71, Hanbauer et al. 247, and Hellström, and co‐workers 265,266


Figure 21.

Dopamine (DA) and norepinephrine (NA) contents in rat carotid body. Areas of large circles, proportional content detected under basal conditions. Upward and downward arrows, increase or decrease in content produced by different conditions. Thin horizontal arrows, moderate but statistically significant effects. Thick horizontal arrows, pronounced effects. Small circles, no effects. Responses to a precursor (L‐dopa), a monoamine oxidase (MAO) inhibitor (pargyline), 1 wk of carotid neurotomy, chronic treatment with a glucocorticoid (dexamethasone) and a mineralocorticoid (doca), 15–60 min of hypoxia, 2‐h administration of reserpine and a DBH inhibitor [diethyldithiocarbamate (DDC)], 1 wk of sympathectomy, and chronic administration of 6‐hydroxydopamine (6‐OHDA).

Data from Hanbauer and Hellström 245, Hellström et al. 264, and Hellström and Kaslow 266


Figure 22.

Dose‐response curves of cat carotid chemoreceptors to haloperidol at various levels of PaO2. Haloperidol had little effect during hyperoxia when receptor activity was slight. At lower PaO2 levels, as activity increased, haloperidol had an augmenting effect. At all levels of PaO2, saturation dose of haloperidol appeared to be 1 mg/kg. At each dose of haloperidol, responses to steady‐state PaO2 were obtained systematically. Each data point represents 1 measurement.

From Lahiri et al. 370


Figure 23.

Effects of close intra‐arterial injections of Met‐enkephalin before (A) and after (B) naloxone on cat carotid nerve discharges. Met‐enkephalin inhibited chemoreceptor activity, and this effect was blocked by naloxone. PSA, systemic arterial pressure.

From Pokorski and Lahiri 494


Figure 24.

Inhibition of cat carotid chemoreceptor response to PaO2 by oligomycin. Carotid chemoreceptor steady‐state responses to PaO2 at constant PaCO2 in 8 experiments before (A) and after (B) oligomycin (50–500 μg ia). Open symbols in B correspond to closed symbols in A. Chemoreceptor activity is insensitive to PaO2 level after oligomycin.

From Mulligan et al. 449


Figure 25.

Responses of carotid chemoreceptor afferent of cat to similar changes in PETCO2, before (A, C) and after (B, D) oligomycin (200 μg ia) during hyperoxia (PaO2 > 400 Torr). After oligomycin there were appreciable overshoots and undershoots in activity.

From Mulligan et al. 448


Figure 26.

Temporal separation of aortic and carotid body stimulation in the dog. A: nicotine injected through catheter placed in aorta (just beyond aortic valves) reaches aortic bodies within 1 s; coils of plastic tubing inserted in common carotids delay nicotine from reaching carotid bodies for 75 s. B: nicotine injected at 2 stimulates aortic bodies and causes tachycardia and hypertension. Neuromuscular blocking agent (succinylcholine) injected at 1 produces apnea and eliminates effects of hyperventilation. Top tracing, respiratory air flow; bottom tracing, carotid blood pressure. At 3, 45 s was deleted to save space; A.A., ascending aorta.

From Comroe and Mortimer 99


Figure 27.

Effects of carotid chemoreceptor stimulation on phasic and mean arterial pressure, left ventricular (LV) pressure, dP/dt, LV diameter, respiration (monitored by pneumograph), and heart rate in dog with spontaneous respiration (left panel). Heart rate remained constant, but chemoreceptor stimulation markedly increased respiration with increase in aortic and LV pressures and in dP/dt. With ventilation controlled (right panel), same carotid chemoreceptor stimulus induced larger increase in pressures and dP/dt.

From Vatner and Rutherford 554


Figure 28.

Hemodynamics during hypoxic hypoxia (HH) and CO hypoxia (COH) before and after carotid body resection (CBR) in the dog. A: Cao2, arterial oxygen content; Pa, mean arterial pressure; Q, cardiac output; TPR, total peripheral resistance. B: HR, heart rate; SV, stroke volume; PLA, mean left atrial pressure; SW/PLA, stroke work. Bars, ±; SD.

From Sylvester et al. 544


Figure 29.

Effect of arterial hypoxia on systemic hemodynamics in conscious dogs. C, control room‐air breathing; H, hypoxia; brackets, ± SEM.

From Krasney and Koehler 346


Figure 30.

Cardiac responses to arterial hypoxia in conscious dogs. C, control room‐air breathing; H, hypoxia; brackets, ±; SEM.

From Krasney and Koehler 346


Figure 31.

Responses to aortic injection of cyanide (CN) before (left) and after (right) acute carotid denervation and bilateral vagotomy in dog. Changes in perfusion pressure (PP) in gracilis muscle and paw were abolished, indicating that reflex responses in muscle and paw were triggered by afferent impulses from carotid sinus area and aortic arch. Changes in systemic arterial pressure (SAP) were small and persisted with some modification after denervation.

From Calvelo et al. 79, by permission of the American Heart Association, Inc
References
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Article Title: Arterial Chemoreceptors
 

How to Cite

Carlos Eyzaguirre, Robert S. Fitzgerald, Sukhamay Lahiri, Patricio Zapata. Arterial Chemoreceptors. Compr Physiol 2011, Supplement 8: Handbook of Physiology, The Cardiovascular System, Peripheral Circulation and Organ Blood Flow: 557-621. First published in print 1983. doi: 10.1002/cphy.cp020316