BIOL 2305 Peripheral Nervous System - Afferent Division - Part I Peripheral Nervous System - Afferent Peripheral Nervous System – all neural structures outside the brain and spinal cord Includes sensory receptors, peripheral nerves, associated ganglia, and motor endings PNS provides links to and from the external environment Properties of Sensory Systems All afferent pathways have in common: Stimulus (physical energy – internal or external) Sensory receptor (transducer – converts stimulus into AP) Sensory neuron (carries AP along afferent pathway to CNS) Destination: CNS Integration Some travel to cerebral cortex for conscious perception Some reach only spinal cord or lower levels of cerebrum, cerebellum, & brain stem for unconscious perception and/or reflexive response 1 From Sensation to Perception Survival depends upon sensation and perception Sensation is the awareness of changes in the internal and external environment Perception is the conscious interpretation of those stimuli Sensory Receptors: Transducers Transduction – the process on converting stimulus energy into electrical impulses that can be sent to the CNS Sensory receptors – sensory nerve endings that responds to changes in the environment around them by transducing stimuli into electrical impulses Ion channels or second messengers initiate a change in membrane potential of receptor Local depolarizations (graded potentials) trigger electrical impulses (action potentials) that travel to the CNS (brain and spinal cord) The realization of these stimuli (sensation and perception) occur in the brain 2 Sensory Receptor Types Receptor Classification Mechanoreceptors – respond to touch, pressure, vibration, stretch, and itch e.g., Pacinian corpuscles in skin, baroreceptors in aorta Thermoreceptors – sensitive to changes in temperature Photoreceptors – respond to light energy e.g., rods and cones of the retina Chemoreceptors – respond to chemicals e.g., olfactory neurons (smell), taste buds, carotid and aortic bodies (changes in blood chemistry) Nociceptors – sensitive to pain-causing stimuli e.g., free nerve endings Osmoreceptors – detect changes in concentration of solutes, osmotic activity (primarily found in the hypothalamus) Receptor Characteristics The receptor must have specificity for the stimulus energy The receptor’s receptive field must be stimulated Stimulus energy must be converted into a graded potential A generator potential in the associated sensory neuron must reach threshold 3 Types of Graded Potentials Generator potentials Occur in specialized nerve endings of sensory neurons Stimulus opens ion channels in receptor causing local current flow Local current flow opens ion channels in afferent neuron AP-generating region If threshold reached, AP is generated which propagates to CNS Receptor potentials Occur in separate receptor cells at distal end of sensory neurons Stimulus opens ion channels in receptor causing graded membrane potential Receptor cell releases chemical messengers Chemical messenger opens ion channels in afferent neuron AP generating region If threshold reached, AP is generated which propagates to CNS Sensory Pathways Stimuli exist in a variety of energy forms or modalities – heat, light, sound, pressure, chemical, etc. Stimulus as physical energy makes contact with sensory receptor Receptor acts as a transducer Intracellular signal: usually change in membrane potential Stimulus generator potential threshold action potential to CNS Integration in CNS: cerebral cortex or acted on subconsciously in brain stem or spinal cord Transduction The process of converting energy forms into electrical signals via a generator potential which triggers an action potential if it is large enough to reach threshold. A generator potential is a type of graded potential similar to an EPSP (excitatory postsynaptic potential) Somatic Senses – Internal Stimuli Vision Hearing Taste Smell Equilibrium Somatic Senses 4 Somatic Senses – Internal Stimuli Somatic Pathways First-order neurons (1o) soma reside in dorsal root or cranial ganglia conduct impulses from the skin to the spinal cord or brain stem Second-order neurons (2o) soma reside in the dorsal horn of the spinal cord or medullary nuclei transmit impulses to the thalamus or cerebellum Third-order neurons (3o) located in the thalamus conduct impulses from thalamus to the somatosensory cortex of the cerebrum 5 Sensory Coding Sensory systems code 4 aspects of a stimulus: Modality – type of stimulus chemo-, thermo-, noci-, mechano-, osmo-, photoLocation Physical site of the stimulated receptor Acuity - precision of stimulus location Larger receptive field size and field overlap decreases acuity Lateral inhibition increases acuity Intensity Stronger stimuli result in higher frequency of receptor potentials leading to a higher frequency of action potentials Stronger stimuli also affect a larger area and leads to recruitment of a larger number of receptors and their corresponding sensory neurons Duration/Adaptation Tonic receptors – generally do not adapt Phasic receptors – adapt readily Receptive Fields of Sensory Neurons Receptive Field: Two-point discrimination Receptive field – area within which a receptor can detect a stimulus 6 Lateral Inhibition Lateral Inhibition - a process by which information from neurons at the edge of a stimulus is inhibited; a means of increasing acuity To facilitate localization and sharpen contrast, the most strongly activated pathway at the center inhibits the less excited pathways from the fringe areas Sensory Coding: Stimulus Intensity & Duration Intensity - coded by number of receptors activated and frequency of action potentials Duration - coded by duration of action potentials;, although some receptors can adapt or cease to respond 7 Adaptation Adaptation occurs when sensory receptors are subjected to an unchanging stimulus Receptor membranes become less responsive Receptor potentials decline in frequency or stop Adaptation occurs in the receptor, not the CNS Tonic receptors – do not adapt or adapt very slowly Important when maintaining information about a stimulus is valuable – stretch, pain receptors Phasic receptors – readily adapt Useful in situations where it is important to signal a change in stimulus – tactile (touch) receptors Sensory Adaptation Tonic receptors (pain): Do not adapt or adapt very slowly Produce constant rate of firing as long as stimulus is applied Phasic receptors: Adapt very quickly Burst of activity but quickly reduce firing rate (adapt) if stimulus maintained. Sensory adaptation: cease to pay attention to constant stimuli. Tonic receptors Phasic receptors Receptor Types and Adaptation Receptors responding to pressure, touch, and smell adapt quickly Receptors responding slowly include Merkel’s discs, Ruffini’s corpuscles Pain receptors and proprioceptors do not exhibit adaptation Adaptation mechanisms Mechanical – specialized receptor ending consists of concentric layers of connective tissue. Sustained pressure causes layers to slip, dissipating stimulus intensity Chemical – Na+ channels that initial opened are slowly inactivated Touch (pressure) Mechanoreceptors: Free nerve endings Enclosed nerve endings Lamellated (Pacinian) corpuscles – rapidly adapting skin receptor that detects pressure and vibration. Corpuscle of touch (Meissner‘s) – receptor for discriminative touch Type I cutaneous (Merkel) receptors for discriminative touch Type II cutaneous (Ruffini) receptor for continuous touch sensation Baroreceptors – receptors to detect pressure changes 8 Proprioceptors Muscle spindles In muscles Detect stretch (change in the length of the muscle) Golgi tendon organs In tendons At the insertion of skeletal muscle fibers into their respective tendons Sense force and stretch Joint receptors In the synovial junctions between bones Sense position & pressure Muscle Spindle Structure Consist of collections of specialized muscle fibers known as intrafusal fibers Lie within spindle-shaped connective tissue capsules parallel to extrafusal fibers Each spindle has its own private efferent and afferent nerve supply Play key role in stretch reflex Muscle spindles monitor changes in length of muscle by responding to the rate and degree of change 9 Tendon Organs (Golgi tendon organs) Consists of sensory fiber penetrating a thin capsule of connective tissue and entwining around a few collagen fibers Found at the junctions of a tendon with a muscle Help protect tendons and associated muscles from damage due to excessive tension or stretching Golgi Tendon Organ Stretch Reflex Primary purpose is to resist tendency for passive stretch of extensor muscles by gravitational forces when person is standing upright Classic example is patellar tendon, or knee-jerk reflex 10 Pain Nociceptors Reflexive path Fast pain (Aδ) Slow pain (C) Pain Receptors Three types of receptors (nociceptors): mechanical, thermal, polymodal All are naked nerve endings and do not adapt All can be sensitized by prostaglandins (increase pain) Prostaglandins derived from lipid bilayer of membrane released from damaged tissues Mechanical (crushing, cutting, pinching) and thermal (extreme temperatures) are transmitted over small myelinated A-delta fibers – 30m/sec fast pain pathway Polymodal respond to all kinds of damaging stimuli and is carried by small unmyelinated C-fibers 12m/sec slow pain pathway Nociceptive Transmission Pathway & Fibers A-Delta Fibers (Aδ) “Fast pain” Small, thinly myelinated 10 % sensory pain fibers Conduct at 5-30 m/sec Mechanical and thermal stimuli Sensations of sharp, pricking pain C Fibers “Slow pain” Small, unmyelinatd fibers. 90% of afferent sensory fibers. Conduct at 0.5-2.0 m/sec. Mechanical, thermal, chemical. Long lasting, burning pain. Aδ and C Nociceptors Mediate Pain 11 Neurotransmitters in Spinal Cord Substance P: Key nociceptor transmitters Released from first order sensory neurons Activates ascending pathways that transmit nociceptor impulses Glutamate: Make up majority of excitatory synapses in brain/spinal cord Modifiable Synapses (can increase or decrease excitability) Binds to AMPA receptors, increases permeability, increasing likelihood of AP Binds to NMDA receptors increases excitability of dorsal horn neurons. Spinal Cord: Excitatory Transmitters 12