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Junctional Transmission in Smooth Muscle and the Autonomic Nervous System

Mollie E. Holman, G. D. S. Hirst

10.1002/cphy.cp010112

Source: Supplement 1: Handbook of Physiology, The Nervous System, Cellular Biology of Neurons

Originally published: 1977

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Autonomic Synapses
1.1 Electrical Properties of Ganglion Cells
1.2 Regenerative Electrical Activity in Autonomic Neurons
1.3 Release of Acetylcholine at Autonomic Synapses
1.4 Postsynaptic Action of Acetylcholine
1.5 Slow Synaptic Potentials
1.6 Presynaptic Changes in the Release of Acetylcholine
1.7 Auto inhibitory Phenomena
1.8 Varying Synaptic Actions of Different Presynaptic Fibers
2 Autonomic Nerve‐Smooth Muscle Transmission
2.1 Electrical Properties of Smooth Muscle
2.2 Action Potentials in Smooth Muscle
2.3 Classification of Smooth Muscles
2.4 Some Comments on the Innervation of Smooth Muscle
2.5 Transmission of Excitation
2.6 Transmission of Inhibition
2.7 null
3 Cellular Basis of some of the Actions of the Autonomic Nervous System
4 Conclusion
Figure 1. Figure 1.

A: relationship between steady‐state current and change in membrane potential in a ganglion cell of Auerbach's plexus. B: sample of records showing the effect of current pulses (lower trace) on membrane potential (upper trace). C: change in membrane potential in response to a small hyperpolarizing current pulse, used to determine the time constant of the cell.

From Holman et al. 150
Figure 2. Figure 2.

Effect of increasing depolarizing current pulses (lower traces) applied through an intracellular electrode on membrane potential and spike frequency (upper traces) in guinea pig inferior mesenteric ganglion cell.

From Crowcroft & Szurszewski 80
Figure 3. Figure 3.

Similar time course and amplitude of a spontaneous miniature excitatory postsynaptic potential (min e.p.s.p.) and of an ACh‐evoked response (the records are from different cells). As a rule the times to peak of miniature excitatory postsynaptic potentials were 5–8 ms, and the fastest ACh potentials had a range of 7–10 ms; frog parasympathetic ganglion cell.

From Dennis et al. 87
Figure 4. Figure 4.

A: extracellular record from a ganglion cell (frog sympathetic chain) of response to orthodromic stimulation. P, presynaptic nerve terminal response; S, synaptic response; C, action potential generated in the ganglion cell.B: intracellular recording from chick ciliary ganglion cell. Orthodromic response, recorded from cell during hyperpolarization, consisting of a coupling potential (apparently coincident with the action potential in the presynaptic terminal) followed by a synaptic potential. The coupling potential precedes the synaptic potential by 1.8 ms. [From Martin & Pilar 215.]

From Ginsborg 123
Figure 5. Figure 5.

Responses recorded on the same sweep to ACh and to synaptic stimulation (nerve). Dotted line indicates zero potential; frog sympathetic ganglion cell.

From Dennis et al. 87
Figure 6. Figure 6.

Conductance change during an ESP recorded from a chick ciliary ganglion cell. Solid lines: tracings of ESP's from cell with coupling potential (A) and from cell without (B). Dashed lines: shunt conductance during EPSP, calculated from relation G = (V – E + τV′)/‐RV, where E is the membrane potential in the absence of an ESP; V, the membrane potential at any given time; V′, the rate of change of V; if, the resting cell resistance; and τ, the resting membrane time constant. In A, R = 15 MΩ, τ = 1.5 ms; in B, R = 45 MΩ, τ = 2.0 ms.

From Martin & Pilar 215
Figure 7. Figure 7.

Action potentials recorded from cells of Auerbach's plexus (guinea pig small intestine) in response to intracellular stimulation. Recordings were made at different speeds. B‐D show the prolonged afterhyperpolarization characteristic of about one‐third of the cells recorded from in this preparation.

From Hirst et al. 137
Figure 8. Figure 8.

Apparatus devised by Abe & Tomita 2 to study the cable properties of smooth muscle. Stimulating electrodes consist of silver plates (50 μm thick) with a small hole (ca. 1‐mm diameter) through which the preparation could be passed. The recording chamber is to the left of the stimulating electrodes. The surface of the stimulating electrode exposed to the recording chamber is coated with epoxy resin. Stimulating electrodes are about 1 cm apart. Relative values of the intensity of the stimulating current can be obtained from the voltage gradient measured between electrodes placed within the stimulating compartment. Each compartment is irrigated separately. V records membrane potential and I records relative values of current intensity.
Figure 9. Figure 9.

Equivalent circuit for an electrical cable. The membrane of a small segment of cable (δx) is modeled by δrm and δcm in parallel and the internal resistance between neighboring segments by δri; rm is the resistance and cm the capacitance of the membrane, and ri is the internal resistance of a 1‐cm length of cable. Thus rm (Ω · cm) = Rm /πd and rj, (Ω/cm) = 4Ri/πd2 where d is the diameter of the fiber, Rm is the resistance of 1 cm2 of membrane, and Ri is the specific resistivity of the core of the cable.
Figure 10. Figure 10.

Model of a smooth muscle bundle. The bundle is delimited by a connective tissue sheath (the perimysium) in which each smooth muscle cell (black cylinder) is surrounded by a number of cells, to some of which it is coupled (lines). In the electrical model, each cell had an impedance made up of the parallel resistance (rm) and capacitance (cm) of the cell membrane (rm = 3.6 × 109 Ωcm = 2.8 × 10−11 F), and each coupling was represented by a simple resistance (rb = 7 × 107 Ω); each cell coupled with 4 cells in the transverse (or radial) plane through the bundle and with 2 cells in the longitudinal direction along the bundle.

From Bennett 22
Figure 11. Figure 11.

Response of an electrical model of the smooth muscle cell bundle to different patterns of current injection. A: temporal distribution of potential in the model resulting from current injection from either an intracellular electrode (upper curves) or an extracellular ring electrode (lower curves); (•), experimental results for these kinds of current injection in the guinea pig vas deferens; the number of couplings distant from the point of current injection is given for each curve. B: spatial distribution of potential in the model resulting from current injection from either an intracellular electrode (lower curve) or an extracellular ring electrode (upper curve); (•), experimental results for extracellular current injection in the guinea pig vas deferens; the experimental results for intracellular current injection are for the cat duodenum, assuming a cell diameter of 5 μm

Data for B from Kobayashi et al. 176; figures from Bennett 22
Figure 12. Figure 12.

Small nerve trunk from the serous coat of the vas deferens. Schwann cells (s), containing numerous unmyelinated axons (a), lie in a reinforcing sheath of collagen filaments (c) within a complete investment of perineurium, which in this case is only a single layer of attenuated, squamous, capsular sheet cells. Some of the axons are in contact with each other. A few of the axons contain clumps of synaptic vesicles (v); others contain neurofilaments. Fixation delayed 4 min. × 7,000.

From Merrillees et al. 229
Figure 13. Figure 13.

Effects of TEA (1.1 × 10−3 g/ml) on the membrane activity of antral smooth muscle (guinea pig). The records were taken from the same muscle cell throughout the experiment. The recording electrode was placed at 0.2‐mm distance from the stimulating plate of a set‐up similar to that of Fig. 8. Note that small depolarizing potentials of the membrane could be recorded by applications of the middle grade of the inward current pulses to the tissue. Presumably the spike elicited at the distant stimulating electrode could be recorded by the intracellular electrode because of electrotonic conduction.

From Ito et al. 159
Figure 14. Figure 14.

Peripheral extensions of a single postganglionic axon (Ax). Preterminal branches (P.T. Ax) eventually give rise to varicose terminal axons (T.Ax).
Figure 15. Figure 15.

Profile of an adrenergic neuron deeply embedded in a smooth muscle cell. Muscle cell processes form a mesaxon‐like slit (arrow). Guinea pig vas deferens after treatment with 5‐hydroxydopamine. × 17,500.

From Furness & Iwayama 113
Figure 16. Figure 16.

Intracellular recordings of spontaneous potentials in smooth muscle cells of the guinea pig vas deferens. Records from 3 different preparations (A‐C). Note the difference in time scales.

From Burnstock & Holman 58
Figure 17. Figure 17.

A: records from guinea pig vas deferens of spontaneous and evoked EJP's. In a and b, the hypogastric nerve was stimulated at 1/s; c shows spontaneous EJP's. B: records from guinea pig vas deferens illustrating the time course of subthreshold EJP's. Frequency of stimulation: 0.1, 0.25, 0.5, 0.75, and 1.0/s as indicated. [From Burnstock et al. 61.]

From Burnstock & Holman 58
Figure 18. Figure 18.

A: theoretical junction potential resulting from a transmitter released from close‐contact varicosities. Potential change developed in all cells throughout the smooth muscle syncytium if 20% of the cells undergo a shunt conductance change (dashed line). This shunt conductance change has the same time course as a spontaneous miniature potential and represents the conductance change occurring at a close‐contact varicosity during transmitter release. B: theoretical junction potential resulting from transmitter released from small axon bundles. Potential change developed in cells at the center (large potential) and surface (smaller potential) of a 50‐μm diameter smooth muscle bundle during the diffusion of transmitter out of the bundle; the time course of the shunt conductance change in cells at the center of the bundle caused by the action of the diffusing transmitter is also shown by a broken line. It is suggested that this junction potential is due to transmitter released from small axon bundles in the muscle.

From Bennett 20
Figure 19. Figure 19.

Records from mouse vas deferens. Spontaneous EJP's recorded by superimposing up to 10 sweeps. Record bottom right shows EJP in response to hypogastric nerve stimulation (arrow); 100‐ms calibration bar applies only for this record.

From Holman 148
Figure 20. Figure 20.

Records taken with sucrose gap from a strip of rabbit portal vein. A: normal activity. B: response to stimulation of intramural noradrenergic nerves (horizontal bar). C: response to 0.1 μg norepinephrine (noradrenaline, NA).

From Holman 148
Figure 21. Figure 21.

Intracellular recording showing, in the same cell, EJP's in response to transmural stimulation at (A) 0.6 pulses/s, (B) 1.0 pulses/s, (C) 2.0 pulses/s, (D) 3.6 pulses/s. Note facilitation of the first 6–10 EJP's and the spontaneous fluctuation in amplitude of EJP's, particularly with higher stimulation frequencies. Pigeon gizzard. Vertical calibration, 40 mV; horizontal calibrations, 1 s.

From Bennett 26
Figure 22. Figure 22.

Responses recorded from guinea pig taenia coli to stimulation of intramural inhibitory nerves in the presence of atropine (10−7 g/ml).

From Bennett 17
Figure 23. Figure 23.

Mean IJP amplitude expressed as a percentage of the maximum, against [Ca2+)0. Upper 3 curves (▪, •, ▴) were obtained from separate experiments. The fourth curve (○) shows the effect of decreasing [Ca2+]0 while holding [Mg2+]0 constant at a higher value of 10 mM. Note that at 0.25 mM [Ca2+]., the IJP was abolished. SE for any one point never exceeded 5%.

From J. Weinrich, unpublished observations
Figure 24. Figure 24.

Intracellular potentials recorded from a preganglionic neuron (T3 and T4 level) activated by one (A) and two (B and C) antidromic stimuli. Arrow in A indicates the onset of the slow phase of repolarization (see text). Records B and C indicate that, when the interval between stimuli is sufficiently short, the antidromic action potential conducted up the axon may fail to depolarize the soma to a level at which an action potential is initiated. Time marker, 5 ms; amplitude; 50 mV.

From De Molina Fernandez et al. 86
Figure 25. Figure 25.

Intracellular potentials recorded from tonically active neurons (1, 2, and 3) of the rabbit's superior cervical ganglion. 1: upper trace, respiration with upward movement indicating inspiration; middle trace, blood pressure; lower trace, intracellular potentials. Lower record shows intracellular potentials of the same neuron recorded with a faster movement of a film. 2: upper trace, respiration; lower trace, intracellular potentials. 3: intracellular potentials. Calibration, 50 mV; time mark on each record, 0.5 s.

From Skok 260
Figure 26. Figure 26.

Intracellular recording from a cell in the guinea pig inferior mesenteric ganglion before, during, and after cutting the lumbar colonic nerves. Moment of cutting the nerve is indicated by a dot.

From Crowcroft et al. 79
Figure 27. Figure 27.

Effect of periarterial nerve stimulation on transmission in the myenteric plexus of guinea pig small intestine. Upper trace in each illustration records the membrane potential of a myenteric neuron at slow scan speed (horizontal calibration, 200 ms); lower traces show ESP's evoked by single transmural stimulus applied across the plexus flap close to the recording site (horizontal calibration, 50 ms). In A and C, recorded 15 s before and 15 s after B, respectively, it can be seen that such stimuli each evoked 2 ESP's. However, when the periarterial nerves leading to the plexus flap were stimulated (solid bar, 10 Hz for 1 s) just before a transmural stimulus (B) only a single ESP was evoked. Vertical calibration bar (10 mV) applies to each trace.

From Hirst & McKirdy 139
Figure 28. Figure 28.

Intracellular recordings from 3 different myenteric neurons (A‐C). Each neuron was situated in a flap of myenteric plexus attached to the aboral end of an intestinal segment [see 138]. Calibration bars, 40 mV and 200 ms.

From Hirst & McKirdy 138
Figure 29. Figure 29.

Intracellular recording from circular layer of smooth muscle showing the sequence of changes in membrane potential evoked by a brief distension (1‐s duration; distending volume, 0.3 ml; recording site, 3 cm aboral to balloon). Upper trace was made using drug‐free solution. An IJP followed by an EJP was recorded. Lower trace was made 8 min after changing to atropine solution (2 × 10−7 g/ml); it can be seen that the EJP has been abolished.

From Hirst, Holman, and McKirdy 136


Figure 1.

A: relationship between steady‐state current and change in membrane potential in a ganglion cell of Auerbach's plexus. B: sample of records showing the effect of current pulses (lower trace) on membrane potential (upper trace). C: change in membrane potential in response to a small hyperpolarizing current pulse, used to determine the time constant of the cell.

From Holman et al. 150


Figure 2.

Effect of increasing depolarizing current pulses (lower traces) applied through an intracellular electrode on membrane potential and spike frequency (upper traces) in guinea pig inferior mesenteric ganglion cell.

From Crowcroft & Szurszewski 80


Figure 3.

Similar time course and amplitude of a spontaneous miniature excitatory postsynaptic potential (min e.p.s.p.) and of an ACh‐evoked response (the records are from different cells). As a rule the times to peak of miniature excitatory postsynaptic potentials were 5–8 ms, and the fastest ACh potentials had a range of 7–10 ms; frog parasympathetic ganglion cell.

From Dennis et al. 87


Figure 4.

A: extracellular record from a ganglion cell (frog sympathetic chain) of response to orthodromic stimulation. P, presynaptic nerve terminal response; S, synaptic response; C, action potential generated in the ganglion cell.B: intracellular recording from chick ciliary ganglion cell. Orthodromic response, recorded from cell during hyperpolarization, consisting of a coupling potential (apparently coincident with the action potential in the presynaptic terminal) followed by a synaptic potential. The coupling potential precedes the synaptic potential by 1.8 ms. [From Martin & Pilar 215.]

From Ginsborg 123


Figure 5.

Responses recorded on the same sweep to ACh and to synaptic stimulation (nerve). Dotted line indicates zero potential; frog sympathetic ganglion cell.

From Dennis et al. 87


Figure 6.

Conductance change during an ESP recorded from a chick ciliary ganglion cell. Solid lines: tracings of ESP's from cell with coupling potential (A) and from cell without (B). Dashed lines: shunt conductance during EPSP, calculated from relation G = (V – E + τV′)/‐RV, where E is the membrane potential in the absence of an ESP; V, the membrane potential at any given time; V′, the rate of change of V; if, the resting cell resistance; and τ, the resting membrane time constant. In A, R = 15 MΩ, τ = 1.5 ms; in B, R = 45 MΩ, τ = 2.0 ms.

From Martin & Pilar 215


Figure 7.

Action potentials recorded from cells of Auerbach's plexus (guinea pig small intestine) in response to intracellular stimulation. Recordings were made at different speeds. B‐D show the prolonged afterhyperpolarization characteristic of about one‐third of the cells recorded from in this preparation.

From Hirst et al. 137


Figure 8.

Apparatus devised by Abe & Tomita 2 to study the cable properties of smooth muscle. Stimulating electrodes consist of silver plates (50 μm thick) with a small hole (ca. 1‐mm diameter) through which the preparation could be passed. The recording chamber is to the left of the stimulating electrodes. The surface of the stimulating electrode exposed to the recording chamber is coated with epoxy resin. Stimulating electrodes are about 1 cm apart. Relative values of the intensity of the stimulating current can be obtained from the voltage gradient measured between electrodes placed within the stimulating compartment. Each compartment is irrigated separately. V records membrane potential and I records relative values of current intensity.



Figure 9.

Equivalent circuit for an electrical cable. The membrane of a small segment of cable (δx) is modeled by δrm and δcm in parallel and the internal resistance between neighboring segments by δri; rm is the resistance and cm the capacitance of the membrane, and ri is the internal resistance of a 1‐cm length of cable. Thus rm (Ω · cm) = Rm /πd and rj, (Ω/cm) = 4Ri/πd2 where d is the diameter of the fiber, Rm is the resistance of 1 cm2 of membrane, and Ri is the specific resistivity of the core of the cable.



Figure 10.

Model of a smooth muscle bundle. The bundle is delimited by a connective tissue sheath (the perimysium) in which each smooth muscle cell (black cylinder) is surrounded by a number of cells, to some of which it is coupled (lines). In the electrical model, each cell had an impedance made up of the parallel resistance (rm) and capacitance (cm) of the cell membrane (rm = 3.6 × 109 Ωcm = 2.8 × 10−11 F), and each coupling was represented by a simple resistance (rb = 7 × 107 Ω); each cell coupled with 4 cells in the transverse (or radial) plane through the bundle and with 2 cells in the longitudinal direction along the bundle.

From Bennett 22


Figure 11.

Response of an electrical model of the smooth muscle cell bundle to different patterns of current injection. A: temporal distribution of potential in the model resulting from current injection from either an intracellular electrode (upper curves) or an extracellular ring electrode (lower curves); (•), experimental results for these kinds of current injection in the guinea pig vas deferens; the number of couplings distant from the point of current injection is given for each curve. B: spatial distribution of potential in the model resulting from current injection from either an intracellular electrode (lower curve) or an extracellular ring electrode (upper curve); (•), experimental results for extracellular current injection in the guinea pig vas deferens; the experimental results for intracellular current injection are for the cat duodenum, assuming a cell diameter of 5 μm

Data for B from Kobayashi et al. 176; figures from Bennett 22


Figure 12.

Small nerve trunk from the serous coat of the vas deferens. Schwann cells (s), containing numerous unmyelinated axons (a), lie in a reinforcing sheath of collagen filaments (c) within a complete investment of perineurium, which in this case is only a single layer of attenuated, squamous, capsular sheet cells. Some of the axons are in contact with each other. A few of the axons contain clumps of synaptic vesicles (v); others contain neurofilaments. Fixation delayed 4 min. × 7,000.

From Merrillees et al. 229


Figure 13.

Effects of TEA (1.1 × 10−3 g/ml) on the membrane activity of antral smooth muscle (guinea pig). The records were taken from the same muscle cell throughout the experiment. The recording electrode was placed at 0.2‐mm distance from the stimulating plate of a set‐up similar to that of Fig. 8. Note that small depolarizing potentials of the membrane could be recorded by applications of the middle grade of the inward current pulses to the tissue. Presumably the spike elicited at the distant stimulating electrode could be recorded by the intracellular electrode because of electrotonic conduction.

From Ito et al. 159


Figure 14.

Peripheral extensions of a single postganglionic axon (Ax). Preterminal branches (P.T. Ax) eventually give rise to varicose terminal axons (T.Ax).



Figure 15.

Profile of an adrenergic neuron deeply embedded in a smooth muscle cell. Muscle cell processes form a mesaxon‐like slit (arrow). Guinea pig vas deferens after treatment with 5‐hydroxydopamine. × 17,500.

From Furness & Iwayama 113


Figure 16.

Intracellular recordings of spontaneous potentials in smooth muscle cells of the guinea pig vas deferens. Records from 3 different preparations (A‐C). Note the difference in time scales.

From Burnstock & Holman 58


Figure 17.

A: records from guinea pig vas deferens of spontaneous and evoked EJP's. In a and b, the hypogastric nerve was stimulated at 1/s; c shows spontaneous EJP's. B: records from guinea pig vas deferens illustrating the time course of subthreshold EJP's. Frequency of stimulation: 0.1, 0.25, 0.5, 0.75, and 1.0/s as indicated. [From Burnstock et al. 61.]

From Burnstock & Holman 58


Figure 18.

A: theoretical junction potential resulting from a transmitter released from close‐contact varicosities. Potential change developed in all cells throughout the smooth muscle syncytium if 20% of the cells undergo a shunt conductance change (dashed line). This shunt conductance change has the same time course as a spontaneous miniature potential and represents the conductance change occurring at a close‐contact varicosity during transmitter release. B: theoretical junction potential resulting from transmitter released from small axon bundles. Potential change developed in cells at the center (large potential) and surface (smaller potential) of a 50‐μm diameter smooth muscle bundle during the diffusion of transmitter out of the bundle; the time course of the shunt conductance change in cells at the center of the bundle caused by the action of the diffusing transmitter is also shown by a broken line. It is suggested that this junction potential is due to transmitter released from small axon bundles in the muscle.

From Bennett 20


Figure 19.

Records from mouse vas deferens. Spontaneous EJP's recorded by superimposing up to 10 sweeps. Record bottom right shows EJP in response to hypogastric nerve stimulation (arrow); 100‐ms calibration bar applies only for this record.

From Holman 148


Figure 20.

Records taken with sucrose gap from a strip of rabbit portal vein. A: normal activity. B: response to stimulation of intramural noradrenergic nerves (horizontal bar). C: response to 0.1 μg norepinephrine (noradrenaline, NA).

From Holman 148


Figure 21.

Intracellular recording showing, in the same cell, EJP's in response to transmural stimulation at (A) 0.6 pulses/s, (B) 1.0 pulses/s, (C) 2.0 pulses/s, (D) 3.6 pulses/s. Note facilitation of the first 6–10 EJP's and the spontaneous fluctuation in amplitude of EJP's, particularly with higher stimulation frequencies. Pigeon gizzard. Vertical calibration, 40 mV; horizontal calibrations, 1 s.

From Bennett 26


Figure 22.

Responses recorded from guinea pig taenia coli to stimulation of intramural inhibitory nerves in the presence of atropine (10−7 g/ml).

From Bennett 17


Figure 23.

Mean IJP amplitude expressed as a percentage of the maximum, against [Ca2+)0. Upper 3 curves (▪, •, ▴) were obtained from separate experiments. The fourth curve (○) shows the effect of decreasing [Ca2+]0 while holding [Mg2+]0 constant at a higher value of 10 mM. Note that at 0.25 mM [Ca2+]., the IJP was abolished. SE for any one point never exceeded 5%.

From J. Weinrich, unpublished observations


Figure 24.

Intracellular potentials recorded from a preganglionic neuron (T3 and T4 level) activated by one (A) and two (B and C) antidromic stimuli. Arrow in A indicates the onset of the slow phase of repolarization (see text). Records B and C indicate that, when the interval between stimuli is sufficiently short, the antidromic action potential conducted up the axon may fail to depolarize the soma to a level at which an action potential is initiated. Time marker, 5 ms; amplitude; 50 mV.

From De Molina Fernandez et al. 86


Figure 25.

Intracellular potentials recorded from tonically active neurons (1, 2, and 3) of the rabbit's superior cervical ganglion. 1: upper trace, respiration with upward movement indicating inspiration; middle trace, blood pressure; lower trace, intracellular potentials. Lower record shows intracellular potentials of the same neuron recorded with a faster movement of a film. 2: upper trace, respiration; lower trace, intracellular potentials. 3: intracellular potentials. Calibration, 50 mV; time mark on each record, 0.5 s.

From Skok 260


Figure 26.

Intracellular recording from a cell in the guinea pig inferior mesenteric ganglion before, during, and after cutting the lumbar colonic nerves. Moment of cutting the nerve is indicated by a dot.

From Crowcroft et al. 79


Figure 27.

Effect of periarterial nerve stimulation on transmission in the myenteric plexus of guinea pig small intestine. Upper trace in each illustration records the membrane potential of a myenteric neuron at slow scan speed (horizontal calibration, 200 ms); lower traces show ESP's evoked by single transmural stimulus applied across the plexus flap close to the recording site (horizontal calibration, 50 ms). In A and C, recorded 15 s before and 15 s after B, respectively, it can be seen that such stimuli each evoked 2 ESP's. However, when the periarterial nerves leading to the plexus flap were stimulated (solid bar, 10 Hz for 1 s) just before a transmural stimulus (B) only a single ESP was evoked. Vertical calibration bar (10 mV) applies to each trace.

From Hirst & McKirdy 139


Figure 28.

Intracellular recordings from 3 different myenteric neurons (A‐C). Each neuron was situated in a flap of myenteric plexus attached to the aboral end of an intestinal segment [see 138]. Calibration bars, 40 mV and 200 ms.

From Hirst & McKirdy 138


Figure 29.

Intracellular recording from circular layer of smooth muscle showing the sequence of changes in membrane potential evoked by a brief distension (1‐s duration; distending volume, 0.3 ml; recording site, 3 cm aboral to balloon). Upper trace was made using drug‐free solution. An IJP followed by an EJP was recorded. Lower trace was made 8 min after changing to atropine solution (2 × 10−7 g/ml); it can be seen that the EJP has been abolished.

From Hirst, Holman, and McKirdy 136
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Article Title: Junctional Transmission in Smooth Muscle and the Autonomic Nervous System
 

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Mollie E. Holman, G. D. S. Hirst. Junctional Transmission in Smooth Muscle and the Autonomic Nervous System. Compr Physiol 2011, Supplement 1: Handbook of Physiology, The Nervous System, Cellular Biology of Neurons: 417-461. First published in print 1977. doi: 10.1002/cphy.cp010112