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Physiologic and Pharmacologic Considerations
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The function of the autonomic nervous system in its regulation of the visceral organs is to a high degree independent When the autonomic nerves are interrupted, these organs continue to function (the organism survives), but they are no longer as effective in maintaining homeostasis and adapting to the demands of changing internal conditions and external stresses It was learned long ago that most viscera have a double nerve supply, sympathetic and parasympathetic, and that in general these two parts of the autonomic nervous system exert opposite effects For example, the effects of the sympathetic nervous system on the heart are excitatory and those of the parasympathetic inhibitory However, some structures sweat glands, cutaneous blood vessels, and hair follicles receive only sympathetic postganglionic bers, and the adrenal gland, as indicated above, has only a preganglionic sympathetic innervation Also, some parasympathetic neurons have been identi ed in sympathetic ganglia Neurohumoral Transmission All autonomic functions are mediated through the release of chemical transmitters The modern concept of neurohumoral transmission had its beginnings in the early decades of the twentieth century In 1921, Loewi discovered that stimulation of the vagus nerve released a chemical substance ( Vagustoff ) that slowed the heart Later this substance was shown by Dale to be acetylcholine (ACh) Also in 1921, Cannon reported that stimulation of the sympathetic trunk released an epinephrinelike substance, which increased the heart rate and blood pressure He named this substance sympathin, subsequently shown to be noradrenaline, or NE Dale found that ACh had pharmacologic effects similar to those obtained by stimulation of parasympathetic nerves; he designated these effects as parasympathomimetic These observations placed neurochemical transmission on solid ground and laid the basis for the distinction between cholinergic and adrenergic transmission in the autonomic nervous system The most important of the autonomic neurotransmitters are ACh and NE ACh is synthesized at the terminals of axons and stored in presynaptic vesicles until it is released by the arrival of nerve impulses ACh is released at the terminals of all preganglionic bers (in both the sympathetic and parasympathetic ganglia) as well as at the terminals of all postganglionic parasympathetic and a few special postganglionic sympathetic bers, mainly those subserving sweat glands (Of course, ACh is also the chemical transmitter of nerve impulses to the skeletal muscle bers) Parasympathetic postganglionic function is mediated by two distinct
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types of ACh receptors nicotinic and muscarinic, so named by Dale because the choline-induced responses were similar either to those of nicotine or to those of the alkaloid muscarine The postganglionic parasympathetic receptors are located within the innervated organ and are muscarinic; ie, they are antagonized by atropinic drugs The receptors in ganglia, like those of skeletal muscle, are nicotinic; they are not blocked by atropine but by other agents (eg, tubocurarine) It is likely that more than ACh is involved in nerve transmission at a ganglionic level Many peptides substance P, enkephalins, somatostatin, vasoactive intestinal peptide, adenosine triphosphate (ATP), and most recently nitric oxide have been identi ed in the autonomic ganglia, localizing in some cases to the same cell as ACh (This negates Dale s law, which stipulates that one neuron elaborates only one neurotransmitter) Particular neuronal ring rates appear to cause the preferential release of one or another of these substances Most of the neuropeptides exert their postsynaptic effects through the G-protein transduction system which utilizes adenylcyclase or phospholipase C as intermediaries The neuropeptides act as modulators of neural transmission, although their exact function in many cases remains to be determined With two exceptions, postganglionic sympathetic bers release only NE at their terminals The sweat glands and some blood vessels in muscle are innervated by postganglionic sympathetic bers, but their terminals, as mentioned, release ACh The NE that is discharged into the synaptic space activates speci c adrenergic receptors on the postsynaptic membrane of target cells Adrenergic receptors are of two types, classi ed originally by Ahlquist as alpha and beta In general, the alpha receptors mediate vasoconstriction, relaxation of the gut, and dilatation of the pupil; beta receptors mediate vasodilatation, especially in muscles; relaxation of the bronchi; and an increased rate and contractility of the heart Each of these receptors is subdivided further into two types Alpha1 receptors are postsynaptic; alpha2 receptors are situated on the presynaptic membrane and, when stimulated, diminish the release of the transmitter Beta1 receptors are, for all practical purposes, limited to the heart; their activation increases the heart rate and contractility Beta2 receptors, when stimulated, relax the smooth muscle of the bronchi and of most other sites, including the blood vessels of skeletal muscle A comprehensive account of neurohumoral transmission and receptor function can be found in the review by Hoffman and colleagues and in the monograph by Cooper and colleagues Discussed in the following pages are the ways in which the two divisions of the autonomic nervous system, acting in conjunction with the endocrine glands, maintain the homeostasis of the organism As stated above, the integration of these two systems is achieved primarily in the hypothalamus In addition, the endocrine glands are in uenced by circulating catecholamines, and some of them are innervated by adrenergic bers, which terminate not only on blood vessels but also, in some cases, directly on secretory cells These autonomic-endocrine relations are elaborated in Chap 27
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heart Together, these actions serve to maintain normal blood pressure and allow re ex maintenance of blood pressure with changes in body position Two types of baroreceptors function as the afferent component of this re ex arc by sensing pressure gradients across the walls of large blood vessels Those in the carotid sinus and aortic arch are sensitive to reductions in pulse pressure (the difference between systolic and diastolic blood pressure), while those in the right heart chambers and pulmonary vessels respond more to alterations in blood volume The carotid sinus baroreceptors are rapidly responsive and capable of detecting beat-to-beat changes, in contrast to the aortic arch nerves, which have a longer response time and discriminate only the larger and more prolonged alterations in pressure The nerves arising from these receptors are small-caliber, thinly myelinated bers that course in cranial nerves IX and X and terminate in the NTS In response to increased stimulation of these receptors, vagal efferent activity is reduced, resulting in re ex cardioacceleration This is accomplished through polysynaptic connections between the NTS and the dorsal motor nucleus of the vagus; it is from this structure that vagal neurons project to the sinoatrial node Increased systemic vascular resistance is effected concurrently through parallel connections between the NTS and the medullary pressor areas that project, in turn, to the intermedolateral cells of the midthoracic cord The main sympathetic out ow from these segments is via the greater splanchnic nerve to the celiac ganglion, the postganglionic nerves of which project to the capacitance vessels of the gut The splanchnic capacitance veins act as a reservoir for as much as 20 percent of the total blood volume, and interruption of the splanchnic nerves results in severe postural hypotension After a high-carbohydrate meal there is a marked hyperemia of the gut and compensatory peripheral vasoconstriction in the muscles and skin It has also been noted that the mesenteric vascular bed is responsive to the orthostatic redistribution of blood volume but not to mental stress The opposite response to the one described above, namely bradycardia and hypotension, results when vagal tone is enhanced and sympathetic tone reduced This response can be triggered by baroreceptors, or it may arise from cerebral stimuli such as fear or sight of blood in susceptible individuals Two slower-acting humoral mechanisms regulate blood volume and complement the control of systemic vascular resistance Pressure-sensitive renal juxtaglomerular cells release renin, which stimulates production of angiotensin and in uences aldosterone production, both of which effect an increase of blood volume Of lesser in uence in the control of blood pressure is antidiuretic hormone, discussed in the next chapter; but the effects of this peptide become more important when autonomic failure forces a dependence on secondary mechanisms for the maintenance of blood pressure More recently, in addition to its presence in autonomic ganglia, nitric oxide has been found to have an important local role in maintaining vascular tone, mainly by way of attenuating the response to sympathetic stimulation The extent to which this latter function is under neural control is not clear Emergency and Alarm Reactions Inasmuch as the autonomic nervous system and the adrenal glands were accepted for many years as the neural and humoral basis of all instinctive and emotional behavior, it is remarkable how little sound information has been acquired about their roles in disease In states of chronic anxiety and acute panic reactions, depressive psychosis, mania, and schizophrenia, all of which are characterized by an altered emo-
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