2.4 Autonomic Nervous System

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1 2.4 Autonomic Nervous System The ANS regulates visceral activities normally outside the realm of consciousness and voluntary control: Circulation. Digestion. Sweating. Pupillary size. The ANS consists of: Sympathetic nervous system: the accelerator of the body. Parasympathetic nervous system: the decelerator of the body. Structures In the ANS: Preganglionic fibre: comes from a neuron in the CNS to a ganglion. Autonomic ganglion: region where many synapses occur, allowing signals to diverge from the CNS to a greater number of effectors. Postganglionic fibre: comes from a ganglion to an effector organ. Acetylcholine is the primary neurotransmitter involved in ganglion neurotransmission. However, at the end of postganglionic fibre: Sympathetic neurons release noradrenaline which acts on α1, α2, β1 and β2 receptors. Parasympathetic neurons release acetylcholine which acts on muscarinic acetylcholine receptors. At the effector, receptors for neurotransmitters are GPCRs (no electrical signalling involved) which induce Ca2+ influx and muscle contraction. The location of ganglia differ: Sympathetic ganglia occur close to the spinal cord, in the sympathetic chain. Parasympathetic ganglia occur close to the effector or in the effector (e.g. heart). The origin of nerves of the sympathetic and parasympathetic nervous systems differ: Sympathetic neurons arise from the thoracic and lumbar regions. Parasympathetic neurons arise from the cervical and sacral regions. Sympathetic nervous system

2 Effects of sympathetic stimulation include: Increased cardiac output (heart rate x stroke volume). Generalised vasoconstriction. Vasodilation of blood vessels supplying skeletal muscle. Increased blood pressure. Airway dilation. Pupil dilation (farther and more vision). Glycogenolysis (breakdown of glycogen). Increased sweating. Inhibited urinary and digestive activities. Sympathetic varicosities are small, numerous enlargements of postganglionic axons at neuromuscular junctions. Inter-varicosity axons are 5 μm long. Can be visualised with DiOC2 labelling and microscopy. Contain the presynaptic components for vesicular release of neurotransmitter. Noradrenaline Noradrenaline is synthesised through: Tyrosine transported into neurons is converted to DOPA by tyrosine hydroxylase. DOPA is converted to dopamine by DOPA decarboxylase. Dopamine is converted to noradrenaline by dopamine β-hydroxylase. Noradrenaline is stored in vesicles until release: Exocytosis occurs to release noradrenaline. Noradrenaline binds receptors on postsynaptic membranes. Noradrenaline is removed from the synapse through: Neuronal re-uptake into the presynaptic or postsynaptic neuron. Diffusion into surrounding extracellular space. Metabolism by enzymes. Noradrenaline acts on receptors on the presynaptic membrane to inhibit release (negative feedback). Parasympathetic nervous system Effects of parasympathetic stimulation include: Increased urinary activity. Increased digestive activity (increased insulin secretion, increased motility of

3 digestive tract). Stimulation of mucus production in airways. Sex? Acetylcholine Acetylcholine is synthesised through: Choline and acetyl CoA is combined to form CAT. CAT is converted to acetylcholine. Acetylcholine is stored in vesicles until release (similarly to noradrenaline). Acetylcholine can be broken down to acetate and choline. Acetylcholine can also be converted to acetylcholinesterase. Muscarinic acetylcholine receptors Muscarinic acetylcholine receptors are GPCRs. They produce parasympathetic effects in different body parts: Heart: receptor opens K+ channels, causing hyperpolarisation and slower heart rate. Digestive tract smooth muscle: receptor inactivates K+ channels, causing depolarisation and contraction. Adrenal medulla The adrenal medulla is a modified sympathetic ganglion that, on stimulation by the preganglionic fibre, secretes hormones into the blood. 2.5 ANS and Enteric Nervous System Autonomic nervous system In peripheral sympathetic nerve varicosities on smooth muscle, cotransmission occurs: ATP acts as a neurotransmitter (binding P2X), leading to the release of Ca2+ and fast muscle contraction. Noradrenaline also causes Ca2+ release (via GqPCR and IP3), but slow muscle contraction. Neuropeptide Y (NPY) enhances the response to ATP and noradrenaline. Visceral smooth muscle cells are linked by gap junctions:

4 Allow for Ca2+ and electrical signalling to pass quickly from one cell to the other. Only the surface cells are directly innervated by neurons. All muscle cells contract as a group, and are called unitary smooth muscle. Enteric nervous system The enteric division of the ANS: A different system of afferent neurons, interneurons and motor neurons that form plexuses. Surrounds the gastrointestinal tract. Can function as a separate and independent nervous system. Normally controlled by the CNS through sympathetic and parasympathetic fibres. Movement of food throughout the GI tract occurs in one direction. In the duodenum: Muscularis mucosae doesn t promote muscle contraction. Muscularis externa (containing outer longitudinal muscle layer, neuronal cell layer and inner circular muscle layer) provides lateral contraction and causes gut to close off. Myenteric (Auerbach s) and submucosal (Meissner s) plexi comprise the ENS: Contain ganglia, sensory and motor neurons, and inputs and outputs. In the mucosa, sensory neuron system may promote motor responses.

5 Intestine motility Types of motility in the intestine are: Gastric mixing. Segmentation. Peristalsis. Migrating motor complex. Gastric mixing involves: Breakdown and neutralisation of food through waves of contraction in musculature of stomach. Forming waves of chyme in stomach through opening and closing of sphincters. Segmentation is the most frequent type of motility: Closely spaced contraction of the circular muscle layer. Divides the small intestine into small neighbouring segments. In rhythmic segmentation the sites of circular contractions alternate, leading to mixing. Frequency of segmentations decreases further away from the mouth (11-12/min in duodenum, 8-9/min in ileum) leading to slow forward transport of food content. Peristalsis is the unidirectional downward flow of food. Smooth muscles contract above and relax below the bolus of food. Migrating motor complex occurs in the gut at rest, in the jejunum: General contraction during periods of no feeding. Maintains downward movement of food. Timed like peristalsis. Regulation of gastrointestinal function GI function is regulated by: Smooth muscle: promotes waves of contraction, controlled by interstitial cells. Neurons: sympathetic nervous system, enteric nervous system senses acidity, calcium ions and proteins. Hormones: influence motility of gut, released from one part of the gut but targets another part. In intestinal smooth muscle, contraction requires an increase of cytosolic calcium. Histamine and acetylcholine stimulation cause slow waves of calcium.

6 Slow waves cause spike potentials which lead to increased internal calcium concentration. Calcium causes calcium-induced calcium release. Calcium, along with hormones from PIP2 pathway (calmodulin), activates MLCK. MLCK phosphorylates myosin, causing muscle contraction. Distension of the bowels by food bolus initiate reflexes which cause movement of food: Ascending excitatory reflex: nicotinic cholinergic transmission from sensory neurons, interneurons, excitatory motor neurons cause acetylcholine-stimulated muscle contraction, pushing food away from the oral cavity. Descending inhibitory reflex: nicotinic cholinergic transmission from sensory neurons, interneurons, inhibitory motor neurons cause vasoactive intestinal polypeptide (VIP) or nitric oxide (NO)-induced muscle relaxation, allowing food to move towards the anus. Defecation reflexes involves sympathetic and parasympathetic control arising from the hypothalamus: Sympathetic: hypothalamic autonomic centre sends signal to mesencephalon and pons, which exits the spinal cord at L1-L3 directly as hypogastric nerves to the inferior mesenteric ganglion (bypasses sympathetic chain). Postganglionic neurons innervate the internal anal sphincter. Parasympathetic: hypothalamic autonomic centre sends signal to dorsal motor vagal nucleus and down the spinal cord. Vagus nerve, and pelvic nerve from S2-S4 increases GI motility. The pudendal nerve from S2-S4 innervates the internal and external anal sphincters. 2.6 Stress * see bottom Stress has been defined as a state of threatened homeostasis. Beneficial in small amounts but detrimental in large amounts. A stressor is a set of conditions that cause adaptive physiological and behavioural changes in animals exposed to conditions such as: Unfamiliar procedures. Restraint. Overstimulation of senses. Transportation. Isolation and overcrowding. Barren environments. Handling and injections.

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