CENTRAL REGULATION OF AUTONOMIC FUNCTIONS.

 

 

Change of functional condition of organs in the case of stimulation of autonomic nerves

Symptoms

Sympathetic effecrs

Parasympathetic effecrs

Pupil of eye

Increase

Normal or decrease

Cardiovascular system:

heart beat

strength of cardiac contractility

 

Increase

Increase

 

Decrease

Decrease

Rate of breathing

Diameter of bronchs

Normal or increase

Increase

Decrease

Decrease

Digestive tract:

Salivation

Motility

Secretory function

Sphincters

 

Increase, viscous saliva

Decrease

Decrease

Contract

 

Increase, liguid saliva

Increase

Increase

Relex

Vessels of sceletal muscles

Vessels of skin

Sweet glands

Increase

Decrease

Secretion

 

CRANIAL NERVES THAT CARRY PARASYMPATHETIC FIBERS. If the oculomotor nerve is cut experimentally, the pupil dilates. The parasympathetic fibers within the oculomotor nerve carry nervous impulses that cause the pupil to constrict. Cutting the nerve destroys the balance between parasympathetic and sympathetic innervation. The sympathetic nervous impulses then cause the pupil to dilate. The "drops" placed in the eye for optical examination apparently act in much the same way by blocking the parasympathetic nerve endings.

It has been mentioned that there are four cranial nerves arising from the medulla that carry autonomic fibers and therefore are a part of the craniosacral system. These nerves are the facial glossopharyngeal, vagus, and accessory nerves. The facial nerve includes parasympathetic fibers that are secretary to the lacrimal gland and to the sublingual and submaxillary salivary glands. The lacrimal gland is supplied with postganglionic fibers from the sphenopalatine ganglion. The sublingual and submaxillary salivary glands receive postganglionic fibers arising in the submaxillary ganglion.

Preganglionic fibers in the glossopharyngeal nerve extend outward to the optic ganglion. Postganglionic fibers arise in the otic ganglion and supply the parotid salivary gland. These glands, including the lacrimal, have a double innervation. They derive their sympathetic innervation by way of the superior cervical sympathetic ganglion and carotid plexuses. The action of the two sets of nerves is not clear. Apparently they both contain secretory fibers, but the secretory action of the parasympathetic system seems to be dominant. The vagus nerve contains both motor and visceral afferent fibers. The motor fibers are long preganglionic fibers that extend out to the organ supplied. Very short postganglionic fibers are contained within the organ. Motor fibers are supplied to the larynx, trachea, bronchioles, heart, esophagus, stomach, small intestine, and some parts of the large intestine. Stimulation of the vagus acts as an inhibitor to the heart, causing its rate of beating to slow or to stop. To the muscles of the wall of the digestive tract, branches of the vagus act as accelerator nerves. Peristalsis is increased by parasympathetic stimulation. Parasympathetic fibers to the glands of the digestive tract have regulatory function on secretion, but food content of the stomach or intestine and hormones circulating in the blood can also stimulate secretion.

Parasympathetic fibers from both the right and left vagus nerves enter the great plexuses of the sympathetic system. There is, however, a definite parasympathetic nerve supply to such organs as the pancreas, liver, and kidneys. Nervous stimulation of these organs is, for the most part, merely regulatory. Hormones in the blood normally cause the pancreas and liver to secrete, but stimulation of the vagus increases the flow of pancreatic juice and bile. While sympathetic stimulation of the kidneys by way of the splanchnic nerves results in vasoconstriction and therefore reduced flow of urine, there are many other physiological factors that affect the function of the kidneys. A part of the accessory nerve contains visceral motor and cardiac inhibitory fibers. Certain types of allergy offer examples of overstimulation of the parasympathetic system. Epinephrine can be used to counteract these effects, since it is associated with the action of the sympathetic system.

THE SACRAL AUTONOMICS. The sacral portion of the craniosacral system is composed of preganglionic fibers incorporated in the second, third, and fourth sacral nerves. The fibers extend out to the pelvic plexuses, where they enter into close relationship with fibers of the sympathetic system. Parasympathetic fibers innervate the urogenital organs and the distal part of the colon. Postganglionic fibers are considered to be in the organs supplied or in small ganglia located close by. These parasympathetic fibers are motor to the muscles of the distal two-thirds of the colon, to the rectum, and to the urinary bladder. They carry vasodilator impulses to the penis and clitoris. Inhibitory impulses pass to the internal sphincter muscle of the bladder and to the internal sphincter of the anus.

PARASYMPATHETIC PLEXUSES. Enteric Plexuses the digestive tube has its own intrinsic nerve supply, consisting of the myenteric plexus, located between the longitudinal and circular muscles and a submucous plexus, located under the mucous layer in the sub mucosa. This part of the nervous system extends the entire length of the digestive tube. It can be assumed that parasympathetic fibers entering the wall of the digestive tract are preganglionic fibers that make synaptic connections with neurons of the enteric system. Sympathetic fibers entering the muscular wall, however, are postganglionic fibers and terminate in the tissues that they supply without making synaptic connections.

The enteric plexuses function in maintaining rhythmic peristaltic movement along the digestive tract. Peristalsis is maintained if both sympathetic and parasympathetic nerve supply is cut. The nerves of the autonomic system, however, exert a regulatory effect.

 

SYMPATHETIC AND PARASYMPATHETIC RELATIONSHIPS. Autonomic effects are usually conditioned by other factors such as the presence of hormones in the bloodstream or by circulatory effects. The secretion of a gland can be depressed by the stimulation of an inhibitor nerve; secretion can also be depressed by vasoconstriction of blood vessels supplying the gland, thus limiting its blood supply. While the sympathetic system can be considered as an accelerator to the heart, the situation is reversed in the case of the action of the autonomic system upon the digestive tract. Here the action of sympathetic nerves depresses peristalsis and the secretion of digestive glands during emotional excitement, while the parasympathetic system, as an accelerator, effects a return to normal. When we speak of the sympathetic and parasympathetic nerves as being antagonistic, we mean this in the sense of antagonistic muscles. The nerves from the sympathetic and parasympathetic systems can produce opposite effects, but they provide a correlated adjustment to meet many physiological conditions. Autonomic effects are not always clearly antagonistic. The accommodation reflex of the eye whereby the lens and iris are adjusted to facilitate clear vision appears to be primarily a parasympathetic function so far as the ciliary’s muscle and the muscles of the iris are concerned. The two sets of muscles of the iris seem to have a synergistic relationship, which causes them to contract or dilate the pupil smoothly in a mild state of opposition to each other. The pupil can also dilate in response to an emotional state such as fear or pain. This is due to stimulation of the sympathetic system.

CHEMICAL TRANSMISSION AT AUTONOMIC FUNCTIONS

Transmission at the synaptic junctions between pre- and postganglionic neurons and between the postganglionic neurons and the autonomic effectors is chemically mediated. The principal transmitter agents involved are acetylcholine and norepinephrine, although dopamine is also secreted by interneurons in the sympathetic ganglia.

Chemical Divisions of the Autonomic Nervous System

On the basis of the chemical mediator released, the autonomic nervous system can be divided into cholinergic and noradrenergic divisions. The neurons that are cholinergic are (1) all preganglionic neurons; (2) the anatomically parasympathetic postganglionic neurons; (3) the anatomically sympathetic postganglionic neurons which innervate sweat glands; and (4) the anatomically sympathetic neurons which end on blood vessels in skeletal muscles and produce vasodilatation when stimulated. The remaining postganglionic sympathetic neurons are noradrenergic. The adrenal medulla is essentially a sympathetic ganglion in which the postganglionic cells have lost their axons and become specialized for secretion directly into the bloodstream. The cholinergic preganglionic neurons to these cells have consequently become the secret motor nerve supply of this gland.

RESPONSES OF EFFECTOR ORGANS TO AUTONOMIC NERVE IMPULSES

General Principles

On the basis of the chemical mediator released, the autonomic nervous system can be divided into cholinergic and noradrenergic divisions. The neurons that are cholinergic are (1) all preganglionic neurons; (2) the anatomically parasympathetic postganglionic neurons; (3) the anatomically sympathetic postganglionic neurons which innervate sweat glands; and (4) the anatomically sympathetic neurons which end on blood vessels in skeletal muscles and produce vasodilatation when stimulated. The remaining postganglionic sympathetic neurons are noradrenergic. The adrenal medulla is essentially a sympathetic ganglion in which the postganglionic cells have lost their axons and become specialized for secretion directly into the bloodstream. The cholinergic preganglionic neurons to these cells have consequently become the secret motor nerve supply of this gland.

The smooth muscle in the walls of the hollow viscera is generally innervated by both noradrenergic and cholinergic fibers, and activity in one of these systems increases the intrinsic activity of the smooth muscle whereas activity in the other decreases it. However, there is no uniform rule about which system stimulates and which inhibits. In the case of sphincter muscles, both noradrenergic and cholinergic innervations are excitatory, but one supplies the constrictor component of the sphincter and the other the dilator.

There is usually no acetylcholine in the circulating blood, and the effects of localized cholinergic discharge are generally discrete and of short duration because of the high concentration of acetylcholinesterase at cholinergic nerve endings. Norepinephrine spreads farther and has a more prolonged action than acetylcholine. The epinephrine and some of the dopamine come from the adrenal medulla, but much of the norepinephrine diffuses into the bloodstream from norad-renergic nerve endings.

Cholinergic Discharge

In a general way, the functions promoted by activity in the cholinergic division of the autonomic nervous system are those concerned with the vegetative aspects of day-to-day living. For example, cholinergic action favors digestion and absorption of food by increasing the activity of the intestinal musculature, increasing gastric secretion, and relaxing the pyloric sphincter. For this reason, and to contrast it with the ''catabolic'' noradrenergic division, the cholinergic division is sometimes called the anabolic nervous system.

Noradrenergic Discharge

The noradrenergic division discharges as a unit in emergency situations. The effects of this discharge are of considerable value in preparing the individual to cope with the emergency, although it is important to avoid the teleologic fallacy involved in the statement that the system discharges in order to do this. For example, noradrenergic discharge relaxes accommodation and dilates the pupils (letting more light into the eyes), accelerates the heartbeat and raises the blood pressure (providing better perfusion of the vital organs and muscles), and constricts the blood vessels of the skin (which limits bleeding from wounds). Noradrenergic discharge also leads to lower thresholds in the reticular formation (reinforcing the alert, aroused state) and elevated blood glucose and free fatty acid levels (supplying more energy). On the basis of effects like these, Cannon called the emergency-induced discharge of the noradrenergic nervous system the ''preparation for flight or fight.''

The emphasis on mass discharge in stressful situations should not obscure the fact that the noradrenergic autonomic fibers also subserve other functions. For example, tonic noradrenergic discharge to the arterioles maintains arterial pressure, and variations in this tonic discharge are the mechanism by which the carotid sinus feedback regulation of blood pressure is effected. In addition, sympathetic discharge is decreased in fasting animals and increased when fasted animals are refed. These changes may explain the decrease in blood pressure and metabolic rate produced by fasting and the opposite changes produced by feeding.

Adrenergic Fibers The terminal filaments of most sympathetic postganglionic neurons produce an adrenalin-like substance and are classified as adrenergic. Sympathetic fibers to sweat glands, blood vessels of the skin, and to the arrestors pylorus muscles are exceptions. These postganglionic fibers enter spinal nerves through the gray rami and reach the skin incorporated in peripheral nerves.

The effects of norepinephrine, in conjunction with epinephrine, can be general and widespread. There is experimental evidence that the chemical substance resulting from excitation of sympathetic postganglionic fibers is carried by the bloodstream and can affect organs remote from the point of origin. It is interesting to note that the sympathetic ganglia and the modularly portion of the adrenal gland have the same embryonic origin. They both arise from neural crest cells. Cholinergic Fibers Parasympathetic fibers also produce a chemical mediating substance. In this case the substance is acetylcholine, which is promptly converted to choline and acetic acid by the action of an enzyme called cholinesterase. Since acetylcholine does not remain in its most active state for any great length of time, it is probable that its effects are entirely local. Unlike norepinephrine, it is probably not carried by the bloodstream.

All preganglionic fibers, whether sympathetic or parasympathetic, have been shown to liberate a cholinergic substance, probably identical with acetylcholine. This means that the transmission of the nervous impulse across the point of synapse between the preganglionic and postganglionic fiber is accomplished by the production of acetylcholine.

As we have indicated, postganglionic sympathetic fibers to the sweat glands and to smooth muscles of the skin are cholinergic. These fibers are carried by peripheral nerves. Voluntary motor nerves to skeletal muscles are also cholinergic. On the basis of chemical transmitter substances it appears that the division of the autonomic system into sympathetic and parasympathetic is somewhat artificial.

MEDULLA OBLONGATA

Control of Respiration, Heart Rate, & Blood Pressure

The medullary centers for the autonomic reflex control of the circulation, heart, and lungs are called the vital centers because damage to them is usually fatal. The afferent fibers to these centers originate in a number of instances in highly specialized visceral receptors. The specialized receptors include not only those of the carotid and aortic sinuses and bodies but also receptor cells that are apparently located in the medulla itself. The motor responses are graded and delicately adjusted and include somatic as well as visceral components.

Other Medullary Autonomic Reflexes

Swallowing, coughing, sneezing, gagging, and vomiting are also reflex responses integrated in the medulla oblongata. Coughing is initiated by irritation of the lining of the respiratory passages. The glottis closes and strong contraction of the respiratory muscles builds up intrapulmonary pressure, whereupon the glottis suddenly opens, causing an explosive discharge of air. Sneezing is a somewhat similar response to irritation of the nasal epithelium. It is initiated by stimulation of pain fibers in the trigeminal nerves.

RELATION OF HYPOTHALAMUS TO AUTONOMIC FUNCTION

Many years ago, Sherrington called the hypothalamus "the head ganglion of the autonomic system." Stimulation of the hypothalamus produces autonomic responses, but there is little evidence that the hypothalamus is concerned with the regulation of visceral function per se. Rather, the autonomic responses triggered in the hypothalamus are part of more complex phenomena such as rage and other emotions.

"Parasympathetic Center"

Stimulation of the superior anterior hypothalamus occasionally causes contraction of the urinary bladder, a parasympathetic response. Largely on this basis, the statement is often made that there is a "parasympathetic center " in the anterior hypothalamus. However, bladder contraction can also be elicited by stimulation of other parts of the hypothalamus, and hypothalamic stimulation causes very few other parasympathetic responses. Thus, there is very little evidence that a localized "parasympathetic center" exists. Stimulation of the hypothalamus can cause cardiac arrhythmias, and there is reason to believe that these are due to simultaneous activation of vagal and sympathetic nerves to the heart.

Sympathetic Responses

Stimulation of various parts of the hypothalamus, especially the lateral areas, produces a rise in blood pressure, pupillary dilatation, piloerection, and other signs of diffuse noradrenergic discharge. The stimuli that trigger this pattern of responses in the intact animal are not regulatory impulses from the viscera but emotional stimuli, especially rage and fear. Noradrenergic responses are also triggered as part of the reactions that conserve heat.

Low-voltage electrical stimulation of the middorsal portion of the hypothalamus causes vasodilatation in muscle. Associated vasoconstriction in the skin and elsewhere maintains blood pressure at a fairly constant level. This observation and other evidence support the conclusion that the hypothalamus is a way station on the so-called cholinergic sympathetic vasodilator system, which originates in the cerebral cortex. It may be this system, which is responsible for the dilatation of muscle blood vessels at the start of exercise.

Stimulation of the dorsomedial nuclei and posterior hypothalamic areas produces increased secretion of epinephrine and norepinephrine from the adrenal medulla. Increased adrenal medullary secretion is one of the physical changes associated with rage and fears and may occur when the cholinergic sympathetic vasodilator system is activated. It has been claimed that there are separate hypothalamic centers for the control of epinephrine and norepinephrine secretion. Differential secretion of one or the other of these adrenal medullary catecholamines does occur in certain situations, but the selective increases are small.

RELATION TO SLEEP Lesions of the posterior hypothalamus cause prolonged sleep, and stimulation of the dorsal hypothalamus in conscious animals causes them to go to sleep. These observations have led to consideralable speculation about the existence of 'sleep centers" a ' 'wakefulness centers '' in the hypothalamus.

RELATION TO CYCLIC PHENOMENA Lesions of the suprachiasmatic nuclei disrupt the circadian rhythm in the secretion of ACTH and melatonin. In addition, these lesions interrupt estrous cycles and activity patterns in laboratory animals. The suprachiasmatic nuclei receive an important input from the eyes via the retinohypothalamic fibers, and it appears that they normally function to entrain various body rhythms to the 24-hour light-dark cycle. There is a prominent serotonergic input from the raphe nuclei to the supra-chiasmatic nuclei, but the exact relation of this input to their function is not known. Feeding & Satiety Centers Hypothalamic regulation of the appetite for food depends primarily upon the interaction of 2 areas: a lateral "feeding center" in the bed nucleus of the medial forebrain bundle at its junction with the pallid hypothalamic fibers, and a medial "satiety center" in the ventromedial nucleus. Stimulation of the feeding center evokes eating behavior in conscious animals, and its destruction causes severe, fatal anorexia in otherwise healthy animals. Stimulation of the ventromedial nucleus causes cessation of eating, whereas lesions in this region cause hyperphagia and, if the food supply is abundant, the syndrome of hypothalamic. Destruction of the feeding center in rats with lesions of the satiety center causes anorexia, which indicates that the satiety center-functions by inhibiting the feeding center.

ANATOMIC CONSIDERATIONS

The term limbic lobe or limbic system is applied to the part of the brain that consists of a rim of cortical tissue around the hilus of the cerebral hemisphere and a group of associated deep structures – the amygdala, the hippocampus, and the septal nuclei. The region was formerly called the rhinencephalon because of its relation to olfaction, but only a small part of it is actually concerned with smell.

LIMBIC FUNCTIONS

Stimulation and ablation experiments indicate that in addition to its role in olfaction, the limbic system is concerned with feeding behavior. Along with the hypothalamus, it is also concerned with sexual behavior, the emotions of rage and fear, and motivation.

Autonomic Responses & Feeding Behavior

Limbic stimulation produces autonomic effects, particularly changes in blood pressure and respiration. These responses are elicited from many limbic structures, and there is little evidence of localization of autonomic responses. This suggests that the autonomic effects are part of more complex phenomena, particularly emotional and behavioral responses. Stimulation of the amygdaloid nuclei causes movements such as chewing and licking and other activities related to feeding. Lesions in the amygdala cause moderate hyperphagia, with indiscriminate ingestion of all kinds of food.

Influence of parasympathetic nervous system on the heart activity

To narcotize a rat and fix it on the preparative table. To make the middle cut on the neck. Find and to separate nervus vagus. Registrate the ECG before and after the electric stimulation of the nerve. Compare the frequency of the heart contraction before and after stimulation of the nerve.

            Pilomotor reflex

To make a thermal (ice) or mechanical stimulus of skin in area of trapezoidal muscle. Pay attention on development of anserine skin on the part of the body. Rise of intensive anserine skin on the whole body testifies of increased of irritation of the sympathetic nervous system (slight anserine skin testifies of normal reaction). It is known, that pileous muscles of head and neck are connected with I-III thoracic segments, pileous muscles of hands are connected with IV-VII thoracic segments, pileous muscles of trunk are connected with VIII-IX thoracic segments.

Functional significance of posterior hypothalamus (stereotaxic research)

To determine stereotaxic coordinates of posterior hypothalamus. To narcotize a rat and fix it on a table. Put the identeferentive electrode into the cervic muscles of a rat. The active electrode into the electrodo-holder and lead it into the posterior hypothalamus.

To count a quantity of respiratorical movements during one minute. To make the stimulation and then count a quantity of respiratorical movements once more.

AUTONOMIC CIRCULATORY EFFECTS. Vasoconstriction is a function of the sympathetic system. Although vasodilation may be a function of the parasympathetic system, experimental results are not conclusive. It appears that sympathetic nerves also can include vasodilator fibers. Other factors can influence the blood vessels, such as hormones circulating in the blood stream, the CO2 content of the blood, and temperature.

Sympathetic fibers are conveyed to the blood vessels of the arms and legs by way of the spinal nerves of the central nervous system supplying these regions. Vasoconstriction can be localized or general. In an emergency calling tor quick action, general vasoconstriction causes a rise in blood pressure. At the same time Vasoconstriction may reduce the flow of blood to the digestive tract in a localized area. Muscular exercise requires an increased flow of blood to the skeletal muscles and, therefore, vasodilation of the blood vessels supplying them. Coronary arteries supplying the heart muscle are dilated also. The action of the sympathetic system is supported by epinephrine in the bloodstream. Central Regulation of Visceral Function

The levels of autonomic integration within the central nervous system are arranged, like their somatic counterparts, in a hierarchy. Simple reflexes such as contraction of the full bladder are integrated in the spinal cord. More complex reflexes that regulate respiration and blood pressure are integrated in the medulla oblongata. Those that control papillary responses to light and accommodation are integrated in the midbrain. The complex autonomic mechanisms that maintain the chemical constancy and temperature of the internal environment are integrated in the hypothalamus. The hypothalamus also functions with the limbic system as a until that regulates emotional and instinctual behavior.

Autonomic Reflexes of Spinal cord

Reflex contractions of the full bladder and rectum occur in spinal animals and humans, although the bladder is rarely emptied completely. Hyperactive bladder reflexes can keep the bladder in a shrunken state long enough for hypertrophy and fibrosis of its wall to occur. Blood pressure is generally normal at rest, but the precise feedback regulation normally supplied by the baroreceptor reflexes is absent and wide swings in pressure are common. Bouts of sweating and blanching of the skin also occur.

Sexual Reflexes of Spinal cord other reflex responses are present in the spinal animal, but in general they are only fragments ol patterns that are integrated in the normal animal into purposeful sequences. The sexual reflexes are example. Coordinated sexual activity depends upon a series of reflexes integrated at many neural levels arc is absent after cord transection. However, genital manipulation in male spinal animals and humans produces erection and even ejaculation. In female spina dogs, vaginal stimulation causes tail deviation and movement of the pelvis into the copulatory position.

Neurosecretion. The hormones of the posterior pituitary gland are synthesized in the cell bodies of neurons in the supraoptic and paraventricular nuclei and transported down the axons of these neurons to the posterior lobe. Some of the neurons make oxytocin and others make vasopressin, and oxytocin-containing and vasopressin-containing cells are found in both nuclei. The neurons also conduct action potentials, and action potentials reaching the endings of the axons trigger release of the hormones by Ca:+-dependent exocytosis. Oxytocin and vasopressin are neural hormones, ie, hormones secreted into the circulation by nerve cells. The term neurosecretion was originally coined to describe the secretion of hormones by neurons.

Effects of vasopressin because its principal physiologic effect is the retention of water by the kidney, vasopressin is often called the antidiuretic hormone (ADH). It increases the permeability of the collecting ducts of the kidney, so that water enters the hypertonic interstitium of the renal pyramids. The urine becomes concentrated and its volume decreases. The overall effect is therefore retention of water in excess of solute; consequently, the effective osmotic pressure of the body fluids is decreased.

Effect of Oxytocin on the Breast in mammals, an important physiological effect of oxytocin is on the myoepithelial cells, smooth muscle-like cells that line the ducts of the breast. The hormone makes these cells contract, squeezing the milk out of the alveoli of the lactating breast into the large ducts (sinuses) and thence out the nipple. Oxytocin causes contraction of the smooth muscle of the uterus. Feature of Hypothalamic Control Anterior pituitary secretion is controlled by chemical agents carried in the portal hypophyseal vessels from the hypothalamus to the pituitary. These substances have generally been referred to as releasing and inhibiting factors, but they are now commonly called hypophysiotropic hormones. The latter term seems appropriate, since they are secreted into the bloodstream and act at a distance from their site of origin. There are 7 relatively well established hypothalamic releasing and inhibiting hormones: corticotropin-releasing hormone (CRH); thyrotropin-releasing hormone (TRH); growth hormone-releasing hormone (GRH); growth hormone-inhibiting hormone (GIH; also called somatostatin); luteinizing hormone-releasing hormone (LHRH); prolactin-releasing hormone (PRH); and prolactin-inhibiting hormone (PIH) Hypothalamus and temperature regulation Anterior hypothalamus response to heat. Posterior hypothalamus response to cold. Afferents go from cutaneous cold receptors, temperature-sensitive cells in hypothalamus.

References:

1. Review of Medical Physiology // W.F. Ganong. – Twentieth edition, 2001. – P. 217-223, 226-229, 232, 233, 242.

2. Textbook of Medical Physiology // A.C. Guyton, J.E. Hall. – Tenth edition, 2002. – P. 364, 632, 681-684, 697-707, 736.