Sensory Physiology Receptor transduction Receptive fields and perception Phasic and tonic receptors Different somatosensory modalities Five special senses Somatic (= general) senses 1. Touch 2. Temperature 3. Nociception 4. Itch 5. Proprioception Special senses 1. Vision 2. Hearing 3. Taste 4. Smell 5. Equilibrium Touch Receptors Free or encapsulated dendritic endings In skin and deep organs, e.g.: Pacinian corpuscles • concentric layers of c.t. ⇒ large receptive field detect vibration opens mechanically gated ion channel adaptation ⇒ receptor type? rapid Temperature Receptors • AKA thermoceptors or thermorecetors • Free dendritic endings in hypodermis • Function in thermoregulation • Cold receptors (< body temp.) • Warm receptors (> body temp.) • Test if more cold or warm receptors in lab • Adaptation only between 20 and 40°C • Nociceptors activated if T > 45°C General Principles The process of stimulus transduction involves the opening or closing of ion channels on a specialised plasma membrane. The size of the receptor potential, just like the synaptic potential is graded and is determined by: • • • • The magnitude of the stimulus The rate of change of the stimulus strength The temporal summation of other receptor potentials The degree of ‘adaptation’ Sensory Physiology: General Principles Sensory information exists in a variety of forms: Light Sound Temperature Vibration Pressure Chemical Osmolarity Tension Sensory Physiology: General Principles Sensory systems convert one form of energy into an electrical signal This conversion of one energy form (eg. light) into an electrical signal (receptor potential) is known as stimulus transduction Sensory Physiology: General Principles Different sensory receptors for different sensory modalities Photoreceptors are responsive to light. (eyes) Mechanoreceptors are responsive to mechanical energy. (skin, ears) Thermoreceptors are response to either heat or cold. (skin) Osmoreceptors are responsive to the concentration of solutes in the extracellular fluid. (tongue, hypothalamus) Chemoreceptors are responsive to specific chemicals (tongue, hypothalamus) Nocireceptors are sensitive to tissue damage (pH, histamine, extreme heat/cold, extreme mechanical pressure). (skin) Types of Sensory Receptors 1. Chemo- (specific ligands) and Osmo(conc. of solutes) 2. Mechano- (touch, pressure, vibration, stretch) 3. Thermo- (temp. change) Cold receptors lower than body temp. Warm receptors (37 - 45oC) > 45oC ? 1. Photo- (light) Somatosensory System MECHANORECEPTORS (touch, pressure vibration) Skin: Meissners corpuscle, Merkles corpuscle / discs, Pancinian corpuscle THERMORECEPTORS (free nerve endings activated by a rising temperature OR a falling temperature) NOCIRECEPTORS (free nerve endings activated by mechanical deformation chemicals and temperature) In the dermis of the skin and are responsible for superficial somatic pain In muscles, tendons and joints and are responsible for deep somatic pain In the visceral organs and are responsible for visceral pain PROPRIOCEPTORS (muscle spindles in muscle; tendon organs in tendon) are responsible for the awareness of muscle length and body position in space Somatosensory System Mechanoreceptors Thermoreceptors Nocireceptors Proprioceptors General Principles Receptor potentials have the same properties as synaptic potentials A receptor may be either a specialised nerve ending of an afferent neuron or a separate cell that is intimately associated with the peripheral endings of the neuron. Figure 46-3 Excitation of a sensory nerve fiber by a receptor potential produced in a pacinian corpuscle. (Modified from Lo&#235;wenstein WR: Excitation and inactivation in a receptor membrane. Ann N Y Acad Sci 94:510, 1961.) Receptor Potentials Typical relation between receptor potential and action potentials when the receptor potential rises above threshold level. Relation of amplitude of receptor potential to strength of a mechanical stimulus applied to a pacinian corpuscle. (Data from Lo&#235;wenstein WR: Excitation and inactivation in a receptor membrane. Ann N Y Acad Sci 94:510, 1961.) Adaptation of different types of receptors, showing rapid adaptation of some receptors and slow adaptation of others. Figure 46-8 Translation of signal strength into a frequency-modulated series of nerve impulses, showing the strength of signal (above) and the separate nerve impulses (below). This is an example of temporal summation. Figure 46-17 The rhythmical output of summated nerve impulses from the respiratory center, showing that progressively increasing stimulation of the carotid body increases both the intensity and the frequency of the phrenic nerve signal to the diaphragm to increase respiration. Figure 46-18 Successive flexor reflexes showing fatigue of conduction through the reflex pathway. Sensory Pathway Stimulus Sensory receptor (= transducer) Afferent sensory neurons CNS Integration, perception Somatosensory Pathways Pathways are composed of a first, second and third order neuron First Order neuron is the 10 afferent. It conveys information to the spinal cord (via the dorsal roots) or the brainstem. Second Order neuron conduct impulses from either the spinal cord or brainstem to the thalamus. Third Order neuron conducts impulses from the thalamus to the somatosensory cortex. There are three main spinal pathways that carry somatosensory information The Anterolateral Pathway (also called Spinothalamic) The Dorsal Column Pathway The Spinocerebellar Pathway (important for proprioception) Sensory Nerves, Spinal Cord & Spinal Nerves Central ‘butterfly’ shaped grey matter contains the cell bodies of motor neurons and interneurons Surrounding white matter contains the axons of ascending and descending neurons (more on this later) Dorsal roots (sensory) join with ventral roots (motor) to form a spinal nerve. Spinal nerve contains sensory and motor information. Spinal Cord & Spinal Nerves Central ‘butterfly’ shaped grey matter contains the cell bodies of motor neurons and interneurons Surrounding white matter contains the axons of ascending and descending neurons (more on this later) Dorsal roots (sensory) join with ventral roots (motor) to form a spinal nerve. Spinal nerve contains sensory and motor information. Dermatomes The area of skin that that is innervated by the sensory nerves of a single spinal nerves is called a dermatome Somatosensory Pathways 3O neuron 2O neuron 1O neuron Somatic Senses • Primary sensory neurons from receptor to spinal cord or medulla • Secondary sensory neurons always cross over (in spinal cord or medulla) → thalamus • Tertiary sensory neurons → somatosensory cortex (post central gyrus) Somatosensory Pathways There are specific pathways which covey sensory (ascending) information up the spinal cord to the brain Somatosensory Pathways Crossover at spinal cord Pain and Temperature Tickle and Itch Poorly localised touch Crossover in medulla Discriminative touch Shape, size texture, weight Vibration Proprioception DORSAL COLUMN PATHWAY • CARRIES FINE TOUCH, POSITION, PRESSURE, VIBRATION, TWO POINT DESRIMINATION stereognosis • AFFERENT SENSORY FIBERS Aβ TYPE. • VERY FAST VELOCITY 30 – 70 m/s • 3 NEURON SYSTEM ANTEROLATERAL PATHWAY • CARRIES PAIN & TEMPRATURE (lat. Sp.Th) • CRUDE TOUCH & PRESSURE ( VENT, Sp. Th) • AFFERENT SENSORY FIBERS Aδ (MYELINATED) FAST PAIN • C FIBERS( UNMYELINATED) SLOW PAIN • RELATIVELY SLOW VELOCITY Aδ – 6 – 30 m/s. C – 0.5 – 2 m/s. • 3 NEURON SYSTEM CNS Distinguishes 4 Stimulus Properties • Modality (nature) of stimulus – Type of receptor • Location – lateral inhibition – population coding • Intensity • Duration Somatosensory Cortex Area on somatosensory cortex related to degree of innervation Receptive Fields • Each 1° sensory neuron picks up information from a receptive field • Often convergence onto 2° sensory neuron ⇒ summation of multiple stimuli • Size of receptive field determines sensitivity to stimulus → Two point discrimination test Pattern of stimulation of pain fibers in a nerve leading from an area of skin pricked by a pin. This is an example of spatial summation. Two-points discrimination varies Lateral inhibition Lateral pathways are strongly inhibited causing most accurate localization. e.g. movement of skin hairs can be well located, temperature and pain are poorly located. Intensity & Duration of Stimulus • Intensity is coded by # of receptors activated and frequency of AP coming from receptor • Duration is coded by duration of APs in sensory neurons • Sustained stimulation leads to adaptation – Tonic receptors – Phasic receptors Tonic Receptors Phasic Receptors • Slow or no adaptation • Rapid adaptation • Continuous signal transmission for duration of stimulus • Cease firing if strength of a continuous stimulus remains constant • Monitoring of parameters that must be continually evaluated, e.g.: baroreceptors • Allow body to ignore constant unimportant information, e.g.: – Smell Tonic and Phasic Receptors • Two classes of receptors that encode stimulus duration – Phasic – produce APs only at the beginning or end of the stimulus encode changes in stimulus, but not stimulus duration – Tonic – produce APs as long as the stimulus continues • Receptor adaptation – AP frequency decreases if stimulus intensity is maintained at the same level Tonic and Phasic Receptors Nociceptors • Free dendritic endings • Activation by strong, noxious stimuli - Function? • 3 categories: – Mechanical – Thermal (menthol and cold / capsaicin and hot) – Chemical (includes chemicals from injured tissues) • Inflammatory Pain • May activate 2 different pathways: – Reflexive protective – integrated in spinal cord – Ascending to cortex (pain or pruritis) Epicritic-electric, lancinating, shock-like Protopathic-constant, burning Pain • Aβ, and AΔ fibers mediate pain • C fibers mediate pruritis • Fast (Aδ fibers) pain is sharp • Slow pain (C) is throbbing • Ascend to limbic system and hypothalamusEmotional Distress – Modulation • Gate Control Theory: We can inhibit the pain response (fig 1012c) • Pain control – NSAIDs (inhibit COX) – Opiates (inhibit NT release) Referred Pain Pain in organs is poorly localized May be displaced if Multiple 1° sensory neurons converge on single ascending tract Referred Pain Referred Pain: Heart