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Physical Factors of Regulation

Paul M. Vanhoutte

10.1002/cphy.cp020216

Source: Supplement 7: Handbook of Physiology, The Cardiovascular System, Vascular Smooth Muscle

Originally published: 1980

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Temperature
1.1 Effect of Temperature on Isolated Blood Vessel Wall
1.2 Analysis of Effects of Temperature on Different Functional Components of Blood Vessel Wall
1.3 Role in the Intact Organism
2 Light
2.1 Photoactivated Contraction
2.2 Photorelaxation
2.3 Summary
3 Vibration
Figure 1. Figure 1.

Effect of temperature on isolated segment of saphenous vein perfused at constant flow with Krebs‐Ringer bicarbonate solution. A: under resting conditions, changes in perfusate temperature had no effect. B: with venoconstriction induced and sustained by continuous electrical stimulation (3 Hz), warming perfusate decreased and cooling augmented venoconstriction. C: with distending pressure increased by increasing flow rate through segment, changes in perfusate temperature had no effect.

From Vanhoutte and Shepherd 291
Figure 2. Figure 2.

Local cooling depresses and local warming augments the spontaneous activity of the mesenteric vein of the dog.

From Vanhoutte and Lorenz 287
Figure 3. Figure 3.

Comparison of thermal sensitivity of isolated strips from saphenous, mesenteric, and femoral veins in control conditions and during sustained electrical stimulation. In saphenous and mesenteric veins, but not in femoral veins, the response to nerve stimulation is augmented by cooling. In all three veins, however, cooling causes slight relaxation when nerve stimulation is absent.

From Vanhoutte and Lorenz 287
Figure 4. Figure 4.

Effect of cooling and rewarming on the response of the ear (•) and femoral (○) arteries to epinephrine. Response at each temperature is expressed as a percentage of the response at 37°C. Optimal responsiveness is obtained at much lower temperatures in the cutaneous artery.

From Glover et al. 112
Figure 5. Figure 5.

Potentiation of β‐receptor mechanism by cooling in cutaneous veins. Effect of increasing doses of isoproterenol on the reaction of saphenous vein strips to electrical stimulation. Upper curve, isoproterenol at 37°C; lower curve, isoproterenol at 29°C. Stimulation parameter selected (8 V, 2 ms, 5 Hz for 15 s) caused contractile responses of the same magnitude at both temperatures.

From Vanhoutte and Shepherd 290
Figure 6. Figure 6.

In dogs' saphenous veins the contractile response to increasing concentrations of acetylcholine is augmented by cooling from 37°C (○—○) to 24°C (•—•), despite the presence of an inhibitor of acetylcholinesterase (physostigmine) and of an α‐adrenergic blocking agent (phentolamine). All values shown at 25°C are significantly different from those of control (37°C).

Data from Janssens and Vanhoutte 145
Figure 7. Figure 7.

In isolated cutaneous veins of the dog cooling from 37°C to 24°C shifts to the right the dose‐response curve to norepinephrine in presence of cocaine (left) but not that to K+ in presence of the α‐adrenolytic drug phentolamine (right). Solid symbols indicate that the difference from control (37°C) is statistically significant.

Data from Janssens and Vanhoutte 145
Figure 8. Figure 8.

Opposite effect of altering perfusate temperature during sustained constriction of segments of dog's saphenous veins caused by nerve stimulation (left) and by barium chloride infusion (right). Segments were perfused at constant flow.

From Vanhoutte and Shepherd 291
Figure 9. Figure 9.

Effect that cooling from 37°C to 28°C has on contraction and 3H‐efflux in a helical strip of dog's saphenous vein previously incubated with [3H]norepinephrine. Cooling augments the contraction but depresses the output of tritiated compounds; the latter is used as a marker of the amount of norepinephrine overflowing in the extracellular space, and thus as a qualitative measure of the amounts of transmitter released.

From Shepherd and Vanhoutte 249
Figure 10. Figure 10.

In control solution (left) cooling (from 37°C to 24°C) significantly reduces the tissue uptake of [3H]norepinephrine by paired strips of dog's saphenous veins. In presence of cocaine, an inhibitor of neuronal uptake (right), the uptake of [3H]norepinephrine is not significantly altered by cooling.

Data from Janssens and Vanhoutte 145
Figure 11. Figure 11.

Tentative representation of the effects that moderate cooling has on adrenergic neuroeffector interaction in cutaneous vessels. At the lower temperatures the increased affinity of the α‐adrenergic receptors for norepinephrine is the main reason for the augmented contractile response to nerve stimulation. Moderate cooling depresses neuronal, and possibly extraneuronal, uptake of transmitter and may delay the diffusion of norepinephrine from the junctional cleft. These actions all would tend to increase the concentration of transmitter in the vicinity of the adrenergic receptors on the smooth muscle cells, and thus augment the response. Data available imply that the evoked release of norepinephrine is depressed by moderate cooling, which then could counteract the previously cited effects on the disposition of the transmitter and tend to depress the response to nerve stimulation; this assumption is made unlikely by observations that the response to sympathetic nerve stimulation is augmented by moderate cooling. No direct information on the effects of lowering the temperature on the enzymatic degradation of norepinephrine seems available; the biological data imply that, if there is such an effect, it is of little physiological importance. , cooling depresses the process; , cooling augments the process; , the effect of cooling on the process would tend to decrease the mechanical response to nerve stimulation; , the effect of cooling would tend to augment the response;?, data available but insufficient.
Figure 12. Figure 12.

Effect of temperature on rate and magnitude of response of glycerol‐extracted fibers of the hog carotid artery to 20 mM ATP.

From Bohr et al. 23
Figure 13. Figure 13.

Effect of temperature on the hydrolysis of ATP by actomyosin extracts of hog carotid arteries; the enzyme is activated by Mg2+ in solution of low‐ionic strength.

From Murphy 214
Figure 14. Figure 14.

Relationship between oxygen consumption (QO2) and temperature of the incubation medium in isolated human umbilical arteries. Relationship is neither linear nor exponential, but shows that the QO2 rises rapidly as the temperature is increased, particularly from 10°C to 30°C, which to the investigators suggested that the metabolic processes may be different at high and at low temperatures.

From Colthart and Roach 55. Reproduced by permission of the National Research Council of Canada from the Canadian Journal of Physiology and Pharmacology. 48: 377–381, 1970
Figure 15. Figure 15.

Lowering the temperature of the incubation medium significantly reduces 45C uptake by rat aorta microsomal membrane vesicles.

Data from Moore 211
Figure 16. Figure 16.

Potentiation by local cooling does not depend on external Ca2+ concentration. Reaction to electrical stimulation (2 Hz) at two different temperatures (•, 27°C; ○, 37°C) in a group of six saphenous veins incubated in solutions with increasing Ca2+ concentration. Broken line (▵) shows the difference in response between the two temperatures.

From Shepherd and Vanhoutte 249
Figure 17. Figure 17.

Temperature dependency of the K+‐induced relaxation of dog cerebral arteries. K+ (5 mM) is added during contractions caused by prostaglandin F2α (3 × 10−7 to 2 × 10−6 M). •, absolute values; ○, percentage changes.

From Toda 272
Figure 18. Figure 18.

Time course of the transient increase of contractile activity observed in a portal vein preparation upon return to standard Krebs solution (arrows) after previous equilibration in a hyperosmotic solution with 100 mM urea. Duration of the excitatory response is markedly increased with decreasing temperature.

From Jonsson et al. 151
Figure 19. Figure 19.

Tentative representation of the effects of moderate cooling (e.g., from 37°C to 24°C) on vascular smooth muscle. Major difference between cutaneous and deep vessels appears to be the degree by which cooling augments the affinity of the membrane receptors for vasoconstrictor agonists, norepinephrine in particular; such an increase in affinity tends to potentiate the contractile response to exogenous norepinephrine and to sympathetic nerve stimulation. The depressing effect that cooling has on other cellular processes may, when considered individually, either facilitate or inhibit the contractile process. Among the facilitatory effects are the inhibition of the Na+‐K+ exchanges, and the decreased sequestration and extrusion of Ca2+. Among the inhibitory phenomena are the possibilities that cooling hyperpolarizes the cell membrane, decreases the influx of Ca2+ and its release from cellular stores, depresses the actomyosin ATPase, and decreases intrinsic velocity of the contractile proteins. It is also likely, but not proven, that moderate cooling depresses the metabolic regeneration of ATP. It appears that the other cellular effects of cooling are negative, so that in most deep vessels an actual depression of the contractile response to norepinephrine is observed; by contrast, in cutaneous vessels the increase in receptor affinity for norepinephrine overrules the other negative influences of moderate cooling, and the contractile response is enhanced. NE, norepinephrine; ATP, adenosine triphosphate; ADP, adenosine diphosphate; , cooling depresses the process; , cooling seemingly does not affect the process; , cooling facilitates the process; , effect of cooling on the process tends to decrease the mechanical response of the cell to norepinephrine; , effect of cooling tends to augment the response; ?, data available but insufficient.
Figure 20. Figure 20.

When arteries of cold‐acclimated rabbits (maintained for 3–6 wk at 5°C) are cooled, their optimal response to norepinephrine is obtained at temperatures markedly lower than that of the body core (•—‐•). This potentiating effect of moderate cooling is not observed in arteries from warm‐acclimated rabbits (○—‐○). Responses to norepinephrine are expressed as a percentage of the response at 37°C.

From McClelland et al. 204
Figure 21. Figure 21.

Local cooling of the skin (upper graph) does not affect basal venomotor tone, which is measured as changes in venous pressure in an occluded limb (middle graph), but greatly augments the reflex increase in venous tone caused, for example, by deep breath (DB in lower graph). T, temperature.

From Zitnik et al. 322
Figure 22. Figure 22.

In the dog, local cooling greatly reinforces the centrally induced increase in cutaneous venomotor tone during thermoregulatory adaptations. At the beginning of the experiment, core temperature is maintained at 37°C (rectal and esophageal temperature traces) by shivering (EMG trace) and by peripheral vasoconstriction; the latter is illustrated by the progressive drop in skin temperature (ear temperature trace) and by the relatively high resistance to flow in the lateral saphenous vein (with saphenous perfusion pressure at constant flow). When the blood perfusing the saphenous vein is cooled, a marked augmentation in perfusion pressure is noted. Injection of epinephrine (which is thought to mediate the thermoregulatory responses causing heat dissipation) into the third ventricle caused cessation of shivering, dilatation of the vein to basal levels, and an increase in ear temperature. EMG, electromyogram.

From Shepherd and Vanhoutte 249
Figure 23. Figure 23.

Relaxing effect of radiation on an aortic strip brought to different levels of contraction with phenylephrine. Upper curve, lights off; lower curve, lights on. Difference between points on two curves at given concentration of phenylephrine indicates extent of relaxation produced by 3‐min exposure to 350 nm radiation from xenon arc.

From Furchgott et al. 98
Figure 24. Figure 24.

Tension‐strain curves from two iliac arteries. Left graph is for a 68‐yr‐old man. ○, control; , after 6 h at 300 Hz; and ▾, after 13 h at 300 Hz. Progressive increase in distensibility is obvious. Right graph is for a 65‐yr‐old woman. ○, control; and ▾, after 14 h without vibration in an environment identical to that in the left graph.

From Roach and Melech 239. Reproduced by permission of the National Research Council of Canada from the Canadian Journal of Physiology and Pharmacology. 49: 288–291, 1971
Figure 25. Figure 25.

Effect of temperature on vibration‐induced inhibitory responses of K+ contracture in the rat portal vein. Vibrations at a frequency of 200 Hz and an amplitude of 3% and 6% of total tissue length causes prompt reductions in active force both at 25°C (A) and at 5°C (B). Note that the rate of recovery of force is considerably reduced at 5°C.

From Ljung and Sivertsson 185, with permission of S. Karger AG, Basel


Figure 1.

Effect of temperature on isolated segment of saphenous vein perfused at constant flow with Krebs‐Ringer bicarbonate solution. A: under resting conditions, changes in perfusate temperature had no effect. B: with venoconstriction induced and sustained by continuous electrical stimulation (3 Hz), warming perfusate decreased and cooling augmented venoconstriction. C: with distending pressure increased by increasing flow rate through segment, changes in perfusate temperature had no effect.

From Vanhoutte and Shepherd 291


Figure 2.

Local cooling depresses and local warming augments the spontaneous activity of the mesenteric vein of the dog.

From Vanhoutte and Lorenz 287


Figure 3.

Comparison of thermal sensitivity of isolated strips from saphenous, mesenteric, and femoral veins in control conditions and during sustained electrical stimulation. In saphenous and mesenteric veins, but not in femoral veins, the response to nerve stimulation is augmented by cooling. In all three veins, however, cooling causes slight relaxation when nerve stimulation is absent.

From Vanhoutte and Lorenz 287


Figure 4.

Effect of cooling and rewarming on the response of the ear (•) and femoral (○) arteries to epinephrine. Response at each temperature is expressed as a percentage of the response at 37°C. Optimal responsiveness is obtained at much lower temperatures in the cutaneous artery.

From Glover et al. 112


Figure 5.

Potentiation of β‐receptor mechanism by cooling in cutaneous veins. Effect of increasing doses of isoproterenol on the reaction of saphenous vein strips to electrical stimulation. Upper curve, isoproterenol at 37°C; lower curve, isoproterenol at 29°C. Stimulation parameter selected (8 V, 2 ms, 5 Hz for 15 s) caused contractile responses of the same magnitude at both temperatures.

From Vanhoutte and Shepherd 290


Figure 6.

In dogs' saphenous veins the contractile response to increasing concentrations of acetylcholine is augmented by cooling from 37°C (○—○) to 24°C (•—•), despite the presence of an inhibitor of acetylcholinesterase (physostigmine) and of an α‐adrenergic blocking agent (phentolamine). All values shown at 25°C are significantly different from those of control (37°C).

Data from Janssens and Vanhoutte 145


Figure 7.

In isolated cutaneous veins of the dog cooling from 37°C to 24°C shifts to the right the dose‐response curve to norepinephrine in presence of cocaine (left) but not that to K+ in presence of the α‐adrenolytic drug phentolamine (right). Solid symbols indicate that the difference from control (37°C) is statistically significant.

Data from Janssens and Vanhoutte 145


Figure 8.

Opposite effect of altering perfusate temperature during sustained constriction of segments of dog's saphenous veins caused by nerve stimulation (left) and by barium chloride infusion (right). Segments were perfused at constant flow.

From Vanhoutte and Shepherd 291


Figure 9.

Effect that cooling from 37°C to 28°C has on contraction and 3H‐efflux in a helical strip of dog's saphenous vein previously incubated with [3H]norepinephrine. Cooling augments the contraction but depresses the output of tritiated compounds; the latter is used as a marker of the amount of norepinephrine overflowing in the extracellular space, and thus as a qualitative measure of the amounts of transmitter released.

From Shepherd and Vanhoutte 249


Figure 10.

In control solution (left) cooling (from 37°C to 24°C) significantly reduces the tissue uptake of [3H]norepinephrine by paired strips of dog's saphenous veins. In presence of cocaine, an inhibitor of neuronal uptake (right), the uptake of [3H]norepinephrine is not significantly altered by cooling.

Data from Janssens and Vanhoutte 145


Figure 11.

Tentative representation of the effects that moderate cooling has on adrenergic neuroeffector interaction in cutaneous vessels. At the lower temperatures the increased affinity of the α‐adrenergic receptors for norepinephrine is the main reason for the augmented contractile response to nerve stimulation. Moderate cooling depresses neuronal, and possibly extraneuronal, uptake of transmitter and may delay the diffusion of norepinephrine from the junctional cleft. These actions all would tend to increase the concentration of transmitter in the vicinity of the adrenergic receptors on the smooth muscle cells, and thus augment the response. Data available imply that the evoked release of norepinephrine is depressed by moderate cooling, which then could counteract the previously cited effects on the disposition of the transmitter and tend to depress the response to nerve stimulation; this assumption is made unlikely by observations that the response to sympathetic nerve stimulation is augmented by moderate cooling. No direct information on the effects of lowering the temperature on the enzymatic degradation of norepinephrine seems available; the biological data imply that, if there is such an effect, it is of little physiological importance. , cooling depresses the process; , cooling augments the process; , the effect of cooling on the process would tend to decrease the mechanical response to nerve stimulation; , the effect of cooling would tend to augment the response;?, data available but insufficient.



Figure 12.

Effect of temperature on rate and magnitude of response of glycerol‐extracted fibers of the hog carotid artery to 20 mM ATP.

From Bohr et al. 23


Figure 13.

Effect of temperature on the hydrolysis of ATP by actomyosin extracts of hog carotid arteries; the enzyme is activated by Mg2+ in solution of low‐ionic strength.

From Murphy 214


Figure 14.

Relationship between oxygen consumption (QO2) and temperature of the incubation medium in isolated human umbilical arteries. Relationship is neither linear nor exponential, but shows that the QO2 rises rapidly as the temperature is increased, particularly from 10°C to 30°C, which to the investigators suggested that the metabolic processes may be different at high and at low temperatures.

From Colthart and Roach 55. Reproduced by permission of the National Research Council of Canada from the Canadian Journal of Physiology and Pharmacology. 48: 377–381, 1970


Figure 15.

Lowering the temperature of the incubation medium significantly reduces 45C uptake by rat aorta microsomal membrane vesicles.

Data from Moore 211


Figure 16.

Potentiation by local cooling does not depend on external Ca2+ concentration. Reaction to electrical stimulation (2 Hz) at two different temperatures (•, 27°C; ○, 37°C) in a group of six saphenous veins incubated in solutions with increasing Ca2+ concentration. Broken line (▵) shows the difference in response between the two temperatures.

From Shepherd and Vanhoutte 249


Figure 17.

Temperature dependency of the K+‐induced relaxation of dog cerebral arteries. K+ (5 mM) is added during contractions caused by prostaglandin F2α (3 × 10−7 to 2 × 10−6 M). •, absolute values; ○, percentage changes.

From Toda 272


Figure 18.

Time course of the transient increase of contractile activity observed in a portal vein preparation upon return to standard Krebs solution (arrows) after previous equilibration in a hyperosmotic solution with 100 mM urea. Duration of the excitatory response is markedly increased with decreasing temperature.

From Jonsson et al. 151


Figure 19.

Tentative representation of the effects of moderate cooling (e.g., from 37°C to 24°C) on vascular smooth muscle. Major difference between cutaneous and deep vessels appears to be the degree by which cooling augments the affinity of the membrane receptors for vasoconstrictor agonists, norepinephrine in particular; such an increase in affinity tends to potentiate the contractile response to exogenous norepinephrine and to sympathetic nerve stimulation. The depressing effect that cooling has on other cellular processes may, when considered individually, either facilitate or inhibit the contractile process. Among the facilitatory effects are the inhibition of the Na+‐K+ exchanges, and the decreased sequestration and extrusion of Ca2+. Among the inhibitory phenomena are the possibilities that cooling hyperpolarizes the cell membrane, decreases the influx of Ca2+ and its release from cellular stores, depresses the actomyosin ATPase, and decreases intrinsic velocity of the contractile proteins. It is also likely, but not proven, that moderate cooling depresses the metabolic regeneration of ATP. It appears that the other cellular effects of cooling are negative, so that in most deep vessels an actual depression of the contractile response to norepinephrine is observed; by contrast, in cutaneous vessels the increase in receptor affinity for norepinephrine overrules the other negative influences of moderate cooling, and the contractile response is enhanced. NE, norepinephrine; ATP, adenosine triphosphate; ADP, adenosine diphosphate; , cooling depresses the process; , cooling seemingly does not affect the process; , cooling facilitates the process; , effect of cooling on the process tends to decrease the mechanical response of the cell to norepinephrine; , effect of cooling tends to augment the response; ?, data available but insufficient.



Figure 20.

When arteries of cold‐acclimated rabbits (maintained for 3–6 wk at 5°C) are cooled, their optimal response to norepinephrine is obtained at temperatures markedly lower than that of the body core (•—‐•). This potentiating effect of moderate cooling is not observed in arteries from warm‐acclimated rabbits (○—‐○). Responses to norepinephrine are expressed as a percentage of the response at 37°C.

From McClelland et al. 204


Figure 21.

Local cooling of the skin (upper graph) does not affect basal venomotor tone, which is measured as changes in venous pressure in an occluded limb (middle graph), but greatly augments the reflex increase in venous tone caused, for example, by deep breath (DB in lower graph). T, temperature.

From Zitnik et al. 322


Figure 22.

In the dog, local cooling greatly reinforces the centrally induced increase in cutaneous venomotor tone during thermoregulatory adaptations. At the beginning of the experiment, core temperature is maintained at 37°C (rectal and esophageal temperature traces) by shivering (EMG trace) and by peripheral vasoconstriction; the latter is illustrated by the progressive drop in skin temperature (ear temperature trace) and by the relatively high resistance to flow in the lateral saphenous vein (with saphenous perfusion pressure at constant flow). When the blood perfusing the saphenous vein is cooled, a marked augmentation in perfusion pressure is noted. Injection of epinephrine (which is thought to mediate the thermoregulatory responses causing heat dissipation) into the third ventricle caused cessation of shivering, dilatation of the vein to basal levels, and an increase in ear temperature. EMG, electromyogram.

From Shepherd and Vanhoutte 249


Figure 23.

Relaxing effect of radiation on an aortic strip brought to different levels of contraction with phenylephrine. Upper curve, lights off; lower curve, lights on. Difference between points on two curves at given concentration of phenylephrine indicates extent of relaxation produced by 3‐min exposure to 350 nm radiation from xenon arc.

From Furchgott et al. 98


Figure 24.

Tension‐strain curves from two iliac arteries. Left graph is for a 68‐yr‐old man. ○, control; , after 6 h at 300 Hz; and ▾, after 13 h at 300 Hz. Progressive increase in distensibility is obvious. Right graph is for a 65‐yr‐old woman. ○, control; and ▾, after 14 h without vibration in an environment identical to that in the left graph.

From Roach and Melech 239. Reproduced by permission of the National Research Council of Canada from the Canadian Journal of Physiology and Pharmacology. 49: 288–291, 1971


Figure 25.

Effect of temperature on vibration‐induced inhibitory responses of K+ contracture in the rat portal vein. Vibrations at a frequency of 200 Hz and an amplitude of 3% and 6% of total tissue length causes prompt reductions in active force both at 25°C (A) and at 5°C (B). Note that the rate of recovery of force is considerably reduced at 5°C.

From Ljung and Sivertsson 185, with permission of S. Karger AG, Basel
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Article Title: Physical Factors of Regulation
 

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Paul M. Vanhoutte. Physical Factors of Regulation. Compr Physiol 2011, Supplement 7: Handbook of Physiology, The Cardiovascular System, Vascular Smooth Muscle: 443-474. First published in print 1980. doi: 10.1002/cphy.cp020216