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NEUROPHYSIOLOGY THE SENSORY SYSTEMS Dr. Jonathan H. Jaggar Professor Department of Physiology STRUCTURE OF THE CENTRAL NERVOUS SYSTEM The CNS is comprised of the brain and spinal cord. 2 BRAIN 3 major subdivisions of the brain: 1) Forebrain 2) Brainstem 3) Cerebellum The forebrain is comprised of the cerebrum and the diencephalon. The brainstem consists of the midbrain, pons, and medulla oblongata. In addition there are 4 interconnected cavities called the cerebral ventricles that contain cerebrospinal fluid. Brainstem All fibers that pass between spinal cord, forebrain and cerebellum go though brainstem. A large portion of the brainstem is a structure called the reticular formation, which is absolutely essential for life. The reticular formation receives and integrates information from all regions of the CNS. 3 Cerebellum Almost exclusively involved in coordinating movements and controlling posture and balance. Receives information from muscles, joints, skin, eyes, ears, vicera and other parts of the brain involved in the control of movement. Forebrain The cerebrum consists of left and right hemispheres. Each cerebral hemisphere is divided into 4 lobes: The 2 cerebral cortex hemispheres are connected by a massive bundle of nerve fibers called the corpus callosum. Cerebral hemispheres consist of the cerebral cortex and a number of cell clusters, called the subcortical nuclei. 4 Cells in cerebral cortex are organized into 6 layers. Cortical neurons are of 2 basic types: pyramidal and non-pyramidal neurons. In the cerebral cortex, basic information is processed into meaningful images, and fine control of skeletal muscles occurs. Nerve fibers enter the cortex from: 1) the diencephalon, particularly the thalamus. 2) other regions of the cortex. 3) the reticular formation of the brainstem. The limbic system is an interconnected group of brain structures that are associated with learning, emotion and behaviour. 5 SPINAL CORD Butterfly shaped gray matter composed of: 1) Interneurons 2) Cell bodies and dendrites of efferent (i.e. away from brain) neurons. 3) Fibers of afferent (i.e. towards brain) neurons. 4) Glial cells – physically and metabolically support neurons. Gray matter surrounded by white matter, which consists of interneurons. Afferent fibers enter on the dorsal side of spinal cord via dorsal roots. Axons of efferent neurons exit the spinal cord on the ventral side, via the ventral roots. The dorsal root ganglia, are small bumps that contain the cell bodies of the dorsal roots. Dorsal and ventral roots combine to form a spinal nerve on each side of the spinal cord. There are 31 pairs of spinal nerves: •8 cervical: associated with the neck, shoulders, arms, hands. •12 thoracic: chest and abdominal walls. •5 lumbar: hip and legs. •5 sacral: genitals and lower digestive tract. 6 Afferent fibers enter on the dorsal side of spinal cord via dorsal roots. Axons of efferent neurons exit the spinal cord on the ventral side, via the ventral roots. The dorsal root ganglia, are small bumps that contain the cell bodies of the dorsal roots. Dorsal and ventral roots combine to form a spinal nerve on each side of the spinal cord. There are 31 pairs of spinal nerves: •8 cervical: associated with the neck, shoulders, arms, hands. •12 thoracic: chest and abdominal walls. •5 lumbar: hip and legs. •5 sacral: genitals and lower digestive tract. Dorsal view, spinal cord 7 PERIPHERAL NERVOUS SYSTEM 12 pairs of Cranial Nerves 31 pairs of Spinal Nerves. 1) Afferent Division: carries information towards CNS. i.e. sensory system 2) Efferent Division: carries information away from CNS. i) Somatic nervous system. Consists only of motor neurons. Innervates skeletal muscle. ii) Autonomic nervous system. Innervates smooth and cardiac muscle, glands, GI. Autonomic can be further subdivided into: a) Sympathetic b) Parasympathetic c) Enteric Major differences in Sympathetic/Parasympathetic 8 Autonomic Nervous System Sympathetic and Parasympathetic usually have opposing effects: e.g. in heart Sympathetic Parasympathetic Sinoatrial Node ↑ heart rate ↓ heart rate Atria ↑ contractility ↓ contractility Atrioventricular Node ↑ conduction velocity ↓ conduction velocity Ventricles ↑ contractility ↓ contractility 9 Cranial Nerves NAME FIBERS COMMENTS I. Olfactory Afferent Neuroepithelium II. Optic Afferent Eye receptors III. Oculomotor Efferent Eye muscles IV. Trochlear Efferent Afferent Eye muscles Eye muscles V. Trigeminal Efferent Afferent Chewing muscles Skin and skeletal muscles VI. Abducens Efferent Afferent Eye muscles Muscle VII. Facial Efferent Afferent Skeletal muscles/ salivary glands Taste buds VIII. Vestibulocochlear Afferent Ear receptors IX. Glossopharyngeal Efferent Afferent Swallowing/salivary Taste buds X. Vagus Efferent Afferent Thorax/abdomen muscle and glands Receptors in thorax/abdomen XI. Accessory Efferent Neck skeletal muscles XII. Hypoglossal Efferent Tongue skeletal muscles 10 Enteric Nervous System Localized to gastrointestinal tract. Consists of: 1) Myenteric plexus 2) Submucous plexus Many interactions between these 2 nerve networks. 11 SENSORY SYSTEM - GENERAL PRINCIPLES The sensory system is part of the nervous system. It consists of sensory receptors that receive stimuli, the neural pathways that transmit that information to the brain, and the parts of the brain that process the information received. Definitions: Sensory information: conscious or unconscious. Sensation: conscious detection of sensory information. Perception: an understanding of sensory information that results from neural processing. Afferent Neuron: carries information towards CNS. Efferent Neuron: carries information away from CNS. 1) Receptors Respond to changes in environment. Two forms: i) On peripheral end of an afferent neuron. ii) Located on a separate cell that is adjacent to an afferent neuron. Receptors are activated by stimuli. Sensory information is transformed into an electrical response via a process known as signal transduction. Sensory receptors respond to specific stimuli, but can be activated by other stimuli if the stimulus is sufficiently high. The type of energy to which a receptor responds with most sensitivity is known as its adequate stimulus. The Receptor Potential Definition: A graded change in membrane potential that is induced by a stimulus that alters the activity of ion channels in a specialized receptor membrane. The localized steady depolarization induces subsequent action potential generation in the attached axon at the first node of Ranvier. When the receptor membrane is on a separate cell, activation of the receptor induces release of neurotransmitter that binds to specific sites on the afferent neuron and induces a graded membrane potential change. After initial stimulation, receptors may undergo adaptation, which is a decrease in the rate of firing of action potentials. The degree of adaptation varies widely between receptor types. The significance of adaptation will be discussed later. 13 2) Neural Pathways in Sensory Systems Sensory Unit: A single afferent neuron and all receptor endings. Receptive Field: Portion of the body that, when stimulated, activates an afferent neuron. Receptive fields of different afferent neurons overlap, so that stimulation activates several sensory units. Sensory unit and receptive field Ascending Pathways Afferent sensory neurons synapse on interneurons, termed “second-order” neurons, in the spinal cord or brain. In turn, these neurons synapse on “third-order” neurons, etc., etc., until the action potential reaches the cerebral cortex. Specific ascending pathways carry single types of stimuli (e.g. from thermoreceptors) to the brainstem and thalamus, before going to the cerebral cortex. Almost all specific pathways cross over to the opposite side of the central nervous system to that from which the stimulus came (i.e. contralateral). 14 Primary sensory areas cerebral cortex Examples of specific ascending pathways and primary receiving areas: •Somatic receptors => somatosensory cortex in parietal lobe of the brain •Eyes => visual cortex in occipital lobe •Ears => auditory cortex in temporal lobe •Taste buds => cortical area adjacent to somatosensory cortex •Olfactory => terminate in limbic system rather than going to thalamus Processing of this information does not end here but continues to the association areas of the cerebral cortex. Non-specific ascending pathways •Activated by sensory units of different types, i.e. signal general information •Neurons receiving input from multiple non-specific pathways are termed “polymodal neurons”. •Non-specific pathways terminate in areas of brainstem, thalamus, and cerebral cortex that are non discriminative, but control alertness. Specific/nonspecific sensory pathways 15 3) Association Cortex and Perceptual Processing The areas of the association cortex lie outside the primary cortical sensory or motor areas. These areas are not considered to be part of the sensory pathways, but their function is to process and analyze sensory information. Therefore, the association cortex is important for determining perception. These factors can affect perception •Sensory receptor adaptation •Personal experience, emotion, personality, social background •Afferent sensory processing and signal discrimination. Occurs at the level of receptor and higher centers. •No feasible sensation – lack receptor for certain energy forms. •Faulty perception from damaged neural networks. e.g. phantom limb. •Pharmacological modulation. e.g. drug induced hallucinations Areas of association cortex 16 4) Primary Sensory Coding 4 aspects of a stimulus are coded: stimulus type, intensity, location, and duration. a) Stimulus Type Modalities are broken down into Submodalities. e.g Taste/sweet. All receptors of a single afferent neuron respond preferentially to the same stimulus type. Overlap of different sensory neuron receptive fields allow a single stimulus to provide multiple sensations because multiple neurons that respond to different modalities will be stimulated. b) Stimulus Intensity Stimulus intensity is translated in 3 primary ways: •Action potential firing frequency. Action potentials do not vary in amplitude, they are all-or-none. Thus, increased stimulus intensity at a single receptor is translated into sensory information by increasing the frequency of evoked action potentials. •Multiple receptor stimulation. An increased stimulus will most likely activate receptors on other branches of the same afferent neuron. Action potentials evoked by these receptors will add to the train of action potentials in the main fiber. •Recruitment. A stronger stimulus will usually affect a larger tissue area and stimulate receptors on other afferent neurons. Action potentials/afferent fiber 17 c) Stimulus Location Touch: location on body. Smell, vision, hearing: origin of stimulus. Stimulus location is coded by the site of the stimulated receptor. Several factors affect the acuity (or precision) of stimulus location: i) The amount of convergence in the ascending pathway. > convergence = < acuity. ii) Size of the receptive field covered by the afferent neuron. > receptive field = < acuity. Stimulus location/two neurons (a) Small receptive field (b) Large receptive field iii) Receptor density. Neurons produce more action potentials if a stimulus occurs in center of receptive field, due to increased receptor density. However, this is not a precise mechanism because an increase in the number of action potentials could also mean a more intense stimulus was applied. Two stimulus points 18 iv) Receptive field overlap. Stimuli will usually activate receptors in the receptive field of more than 1 neuron. Thus, differential firing frequency of multiple neurons will allow coding of stimulus location. A stimulus point 19 v) Lateral Inhibition. This is the most important mechanism to determine stimulus localization. Focuses sensory processing on important messages, allowing signal discrimination by inhibiting information from afferent neurons whose receptors are at the edge of a stimulus. Lateral inhibition can occur at all levels in the sensory pathway, but it usually occurs at the early stages. It occurs most commonly in pathways that require accurate localization, e.g. skin. Afferent pathways/lateral inhibition Effect on actionpotential frequency 20 d) Stimulus Duration The action potential frequency that occurs at the beginning of a stimulus indicates intensity. After the initial response the action potential frequency depends on the receptor type: Rapidly adapting receptors: after initial burst, fires slowly or stops firing. Important for signaling rapid changes in stimuli. Some of these receptors only fire 1 action potential at the initial stimulus onset (“on-response”). Some receptors respond at the beginning and at the end of the stimulus. Slowly adapting receptors: Maintain firing near to initial frequency throughout stimulus duration. Signal slow changes in stimuli, or prolonged events. Rapidly/slowly adapting receptors 21 Central Control of Afferent Information In addition to information filtering that occurs in the ascending pathways, information is also controlled from the higher centers in the brain by descending pathways. In particular, the reticular formation and the cerebral cortex control afferent information via descending pathways. Inhibition from descending pathways can occur: 1) directly, by synapses onto the axon terminals of the primary afferent neuron (termed “presynaptic inhibition”). 2) indirectly, via interneurons. Descending pathways control sensory information 22