BIOL 2305 Peripheral Nervous System – Efferent Division Somatic & Autonomic Efferent division provides communication link between CNS and the activities of muscle and glands – the “effector organs” Two Branches of the Peripheral Efferent Division: Autonomic (ANS) – involuntary cardiac, smooth muscle, exocrine glands, and some endocrine glands Somatic – voluntary movement and secretions Autonomic Pathways consist of a 2 neuron chain: The first motor neuron, called the preganglionic neuron, has its cell body in the CNS and its myelinated axon called the preganglionic fiber extends to autonomic ganglion. The second motor neuron called the postganglionic neuron has its soma in that autonomic ganglion and its unmyelinated axon called the postganglionic fiber extends directly to the effector cell or organ 1 Efferent Pathways Somatic: Single, heavily myelinated axon of the somatic motor neuron extends from the CNS to the effector (lacks ganglia) ANS (Parasympathetic & Sympathetic): Pathways in the ANS are a two-neuron chain 1. The preganglionic (first) neuron has a lightly myelinated axon 2. The postganglionic (second) unmyelinated neuron extends to an effector organ via the postganglionic axon Comparison of Somatic and Autonomic Systems 2 Autonomic Nervous System The ANS operates without conscious control; it is primarily regulated by the hypothalamus and the medulla oblongata with input from the limbic system and other regions of the cerebrum. The afferent component of the ANS consists of general visceral sensory neurons: Interoreceptors such as chemoreceptors (CO2 levels) and mechanoreceptors (degree of stretch of organs and vessels) Afferent signals are not consciously recognized unless intense enough to cause pain or nausea from damaged viscera, fullness of bladder, angina pectoris (inadequate blood flow to heart) The efferent component consists of the autonomic motor neurons that excite or inhibit visceral activities of effector tissues: cardiac muscle, smooth muscle, glands Efferent responses include automatic activities beyond conscious control - dilation/constriction of pupils, accommodation of lens, dilation of blood vessels, heartbeat, GI/GU movement, glandular secretions. Efferent Reflex Pathways Efferent motor neuron synapses at NMJ with skeletal muscle Preganglionic neuron synapses at sympathetic chain ganglion Efferent motor neuron releases NT (ACh) at NMJ Postganglionic neuron releases NT (NE/ACh) at varicosities onto viscera/glands Voluntary Involuntary 3 Synapses in Autonomic Nerves Varicosities Form strings of synapses (called boutons) NT synthesized in varicosities AP from axon triggers voltage-gated Ca2+ channels Influx of Ca2+ stimulates exocytosis of NT NT released into general ISF in synaptic cleft NT diffuses slowly through interstitial fluid to receptors No motor end plate (meaning receptors on the muscle are not clustered) Impact: Large area Slow acting Long duration NT removed by reuptake, degradation, or diffusion 4 Efferent Pathways: Motor & Autonomic Autonomic Nervous System: Sympathetic & Parasympathetic Regulation of the internal environment generally “autonomous” (outside of our conscious control) ANS innervates smooth muscle, cardiac muscle, and glands “Visceromotor” – movement of viscera; effector targets are visceral organs and blood vessels Involve 2 neurons that synapse in a peripheral ganglion: Preganglionic neuron is myelinated Postganglionic neuron is unmyelinated Autonomic nerves release NT’s that may be stimulatory (NE) or inhibitory (ACh) ANS Autonomic nerve pathway Extends from CNS to an innervated organ Two-neuron chain Preganglionic fiber (synapses with cell body of second neuron) Postganglionic fiber (innervates effector organ) 5 Two Divisions of the ANS (Sympathetic and Parasympathetic) Both have preganglionic neurons that originate in CNS. Both have postganglionic neurons that originate outside of the CNS in ganglia. Differences: Sympathetic vs Parasympathetic Parasympathetic Nervous System (PNS) Fibers originate from cranial and sacral areas of CNS CN 3, 7, 9, 10 Sacral 2, 3, 4 Preganglionic fibers are longer Very short postganglionic fibers Preganglionic fibers release acetylcholine (ACh) Postganglionic fibers release acetylcholine (ACh) Sympathetic Nervous System (SNS) SNS Fibers originate in thoracic and lumbar regions of spinal cord T1 – L2 Most preganglionic fibers are short Long postganglionic fibers Preganglionic fibers release acetylcholine (ACh) Most postganglionic fibers release norepinephrine (noradrenaline, NE) 6 Functional Differences Sympathetic - “fight or flight” Catabolic (expend energy) Release of norepinephrine from postganglionic fibers and epinephrine from adrenal medulla Mass activation prepares for intense activity ↑Heart rate ↑ Bronchodilation ↑ Blood [glucose] Parasympathetic - “feed & breed”, “rest & digest” Maintain homeostasis Normally not activated as a whole, stimulation of separate parasympathetic nerves. Release ACh as NT onto effector organs Relaxing effects: ↓ Heart Rate ↑ Vasodilation (to visceral organs) ↑ digestive activity (motility/secretions) Dual innervation of many organs — having a brake and an accelerator provides more control Neurochemistry of the ANS All preganglionic fibers release acetylcholine (cholinergic) Postganglionic PARASYMPATHETIC fibers release acetylcholine (cholinergic) Postganglionic SYMPATHETIC fibers release norepinephrine (adrenergic) Some exceptions: Adrenal medullary chromaffin cells secrete epinephrine (80%) and norepinephrine (20%) into bloodstream Sympathetic nerves innervating sweat glands secrete acetylcholine Sympathetic nerves innervating blood vessels in skeletal muscle secrete acetylcholine Sympathetic nerves innervating renal blood vessels secrete dopamine Autonomic Pathways 7 Synaptic Organization Adrenal Glands Adrenal medulla secretes epinephrine (Epi) and norepinephrine (NE) when stimulated by the sympathetic nervous system. Modified sympathetic ganglion, derived from same embryonic tissue that forms postganglionic sympathetic neurons. Sympathoadrenal system: mass activation of the sympathetic nervous system. Innervated by preganglionic sympatheti c fibers. Secretion of hormones into blood About 20% of hormone release is norepinephrine About 80% of hormone released is epinephrine 8 Adrenergic and Cholinergic NTs Axons of postganglionic neurons have numerous varicosities along the axon that contain NT ACh is NT for all preganglionic fibers of both sympathetic and parasympathetic nervous systems. Transmission at these synapses is termed cholinergic: ACh is NT released by most postganglionic parasympathetic fibers at synapse with effector. NT released by most postganglionic sympathetic nerve fibers is NE Transmission at these synapses is called adrenergic Note: Epi, released by the adrenal medulla, is synthesized from the same precursor as NE Epi and NE collectively called catecholamines Responses to Cholinergic Stimulation All somatic motor neurons, all preganglionic and most postganglionic parasympathetic neurons are cholinergic Release ACh as NT Somatic motor neurons and all preganglionic autonomic neurons are excitatory Postganglionic axons, may be excitatory or inhibitory Muscarinic receptors: ACh binds to receptor Requires the mediation of G-proteins Nicotinic receptors (ligand-gated): ACh binds to nicotinic receptor binding sites Opens a Na+/K+ channel Always excitatory 9 Responses to Cholinergic Stimulation Nicotinic receptors (ligand-gated) open a Na+ & K+ channels; excitatory only Muscarinic receptors involve G-proteins; excitatory or inhibitory M1 receptor – common in exocrine glands and in the CNS; IP3 pathway M2 receptor – receptors are located in the heart, cause a decrease in cAMP in the cell, inhibition of voltage-gated Ca2+ channels, and efflux of K+ M3 receptor – smooth muscle in bronchioles. In blood vessels it causes increased synthesis of nitric oxide by endothelial cells causing vasodilation, stimulates secretion in many glands; IP3 pathway M4 receptor – found in the CNS; decrease cAMP in the cell and, thus, produce generally inhibitory locomotor effects M5 receptor – location is not well known Cholinergic Stimulation (ACh) 10 Responses to Adrenergic Stimulation Excitatory or inhibitory effects, depending on receptor on target cell All act through G-proteins Alpha adrenergic responses: α1 receptors - vasoconstriction in GI tract, skin, kidney – IP3, DAG, Ca2+ α2 receptors - sphincter constriction in GI tract, inhibit insulin release – reduced cAMP Beta adrenergic responses: β1 receptors - increases HR and force of contraction, increase renin secretion – cAMP β2 receptors - bronchiole dilation, vasodilation skeketal muscle , decrease motility GI tract– cAMP β3 receptors - stimulates lipolysis Responses to Adrenergic Stimulation 11 Responses to Adrenergic Stimulation Dual Innervation Organs With Dual Innervation: Innervated by both sympathetic and parasympathetic fibers) Antagonistic – Sympathetic and parasympathetic fibers innervate the same cells. Actions counteract each other (e.g.: heart rate) Complementary – sympathetic and parasympathetic stimulation produces similar effects ex. salivary gland secretion Cooperative – Sympathetic and parasympathetic stimulation produce different effects that work together to produce desired effect. ex: Parasympathetic fibers penile erection Sympathetic fibers ejaculation Organs Without Dual Innervation: Regulation achieved by increasing or decreasing firing rate Examples: Adrenal medulla, arrector pili muscle, sweat glands, and most blood vessels receive only sympathetic innervation 12 Sympathetic vs Parasympathetic Levels of ANS Control The hypothalamus is the main integration center of ANS activity Subconscious cerebral input via limbic lobe connections influences hypothalamic function Other controls come from the cerebral cortex, the reticular formation, and the spinal cord 13 Regulation of the ANS by CNS Prefontal Association Cortex and Limbic System: Responsible for visceral responses that are characteristic of emotional states and behavior Hypothalamus: Primal sympathetic response to anger or fear is brought on by hypothalamus through medulla Medulla: Most directly controls activity of ANS Location of centers for control of cardiovascular, pulmonary, urinary, reproductive, and digestive systems. Spinal cord: Some autonomic reflexes integrated at spinal cord (urination, erection) Somatic Motor Controls Skeletal Muscles Body movement Appendages Locomotion Single neuron CNS origin Heavily myelinated Terminus Branches (motor unit) Neuromuscular junction (motor end plate) Somatic Efferent Consists of the axons of motor neurons which originate in the spinal cord and terminate on skeletal muscle Acetylcholine released from a motor neuron stimulates muscle contraction Motor neurons are the final common pathway by which various regions of the CNS exert control over skeletal muscle activity The areas of the CNS that influence skeletal muscle activity by acting through the motor neurons are the spinal cord, motor regions of the cortex, basal nuclei, cerebellum, and brain stem Nerve Stimulus Skeletal muscles are stimulated by motor neurons of the somatic nervous system Axons of these neurons travel in nerves to muscle cells Axons of motor neurons branch repeatedly as they enter muscles Each axonal ending forms a neuromuscular junction (NMJ) with a muscle fiber 14 Neuromuscular Junction (NMJ) When a nerve impulse reaches the neuromuscular junction: Voltage-regulated calcium channels in the axon membrane open and allow Ca2+ to enter the axon Ca2+ inside the axon terminal causes some of the synaptic vesicles to fuse with the axon membrane and release ACh into the synaptic cleft (exocytosis) ACh diffuses across the synaptic cleft and attaches to ACh receptors on the sarcolemma Binding of ACh to receptors on the sarcolemma initiates an action potential in the muscle ACh is quickly degraded by the enzyme acetylcholinesterase (AChE) 15 Motor Unit: Neuromuscular Functional Unit Motor Unit – a motor neuron and all the muscle fibers it supplies is called a Motor Unit Each muscle has at least one motor nerve that may contain hundreds of motor neuron axons. Axons branch into terminals, each forming a neuromuscular junction with a single muscle fiber 16 Motor Unit The number of muscle fibers per motor unit can vary from a few to several hundred Muscles that control fine movements (fingers, eyes) have small motor units Large weight-bearing muscles (thighs, hips) have large motor units Muscle fibers in a single motor unit are spread throughout the muscle. As a result, stimulation of a single motor unit causes weak contraction of the entire muscle Summary Autonomic branches: sympathetic and parasympathetic Regulate glands, smooth & cardiac muscles Team with endocrine to regulate homeostasis Are regulated by hypothalamus, pons & medulla Have pathways with 2 neurons and a ganglion Use varicosities to release NTs Have diverse receptors: tonic & antagonistic regulation Efferent motor neurons control skeletal muscles Single long myelinated neuron from CNS Neuromuscular junction structure & mechanism 17