Sensory Receptors General Properties Reading: None www.premedhq.com The Study of the Senses All that we know about the world around us is obtained through our senses (“Esse est percipi”): “For there is no conception in a man’s mind, which hath not at first, totally, or in parts, been begotten upon the organs of the Sense” (Thomas Hobbes, 1558-1679). “What is real? How do you define real? If you’re talking about what you can feel, what you can smell, what you can taste and see, then real is simply electrical signals interpreted by your brain… this is the world that you know” – Morpheus to Neo in The Matrix, 1999. Thomas Hobbes Goals: • How does the nervous system encode and process sensory stimuli? • How does activity and circuitry in the nervous system account for sensory experience at the behavioral level? NSC201 Basic Neurobiology 2 Sensation and Perception Sensation: • Related to the physical interaction of a stimulus with a sensory receptor. • Law of Specific Nerve Energies (Johannes Müller, 1829): sensation depends on the type of receptor activated, not the form of activation. Perception: • Conscious awareness and interpretation of sensation. • Related to the particular sensory pathway in the nervous system that processes information from a receptor. • Influenced by “top down” processes: cognition, attention, experience. • Not directly related to the actual stimulus activating the receptor: it is the brain’s interpretation of sensation, based on its internal model of reality. NSC201 Basic Neurobiology 3 Sensation ≠ Perception Parallel! NSC201 Basic Neurobiology 4 Stimulation of Sensory Receptors Stimulation: Stimulus interacts with primary sensory receptor. Accessory structures: Shapes the input to the receptor. Examples: Cornea, iris, and lens of eye Outer/middle/inner ear structures Pacinian corpuscle touch receptor. NSC201 Basic Neurobiology 5 Transduction and Transmission Transduction: the conversion of stimulus energy into electrophysiological response. Transduction current: membrane currents that result from sensory transduction. Receptor Potential: Change in membrane potential produced by transduction currents. Transmission: Direct: transduction currents generate action potentials (typical of “long” neuronal receptors). Indirect: transduction currents cause transmitter release, activating primary afferent fibers ( “short” epithelial receptors). NSC201 Basic Neurobiology 6 Receptor Classification: Processes Neuronal (“long" receptors): Have dendritic and axonal processes. Somatosensory Auditory Visual Transmit information via action potentials. Examples: most somatosensory and olfactory receptors. Epithelial (“short" receptors): Interneuron Ganglion cell NO axonal process. Ganglion cell "Synaptomimetic": functionally and morphologically similar to presynaptic nerve endings. transduction currents modulate release of neurotransmitter, in some cases generating APs in receptor cell. Ganglion cell Examples: visual, auditory/vestibular, taste receptors. NSC201 Basic Neurobiology 7 Receptor Classification: Stimulus Energy Mechanoreceptors • • • • • Touch/pressure Pain Hearing Balance Joint/muscle proprioception Chemoreceptors • • • • Taste Smell Pain Itch Electromagnetic receptors • Vision • Electroreception • Magnetoreception Thermoreceptors • Warm • Cold NSC201 Basic Neurobiology 8 Receptor Classification: Stimulus Origin Exteroceptive: • Located at or near the surface of the body • Provide information about the external environment (senses) Interoceptive: • Found in blood vessels, connective tissues and organs • Provide information about the internal environment (organs, blo0d chemistry) Proprioceptive: • Located in muscles, tendons, joints, muscles and the internal ear (vestibular) • Provide information about body and limb positions, skeletal muscle movements, balance Nociceptive: • Located throughout the body • Provide information about pain NSC201 Basic Neurobiology 9 Types of Transduction: Direct Stimulus interacts directly with ion channels (analogous to ionotropic synaptic receptors). Hair Cells (Auditory/vestibular mechanoreceptors) Transduction channels (stretch activated, cation selective) Transduction current (carried by K+) Ca2+ influx (via Cav channels Synaptic transmission NSC201 Basic Neurobiology 10 Types of Transduction: Indirect Stimulus acts on G-protein coupled receptor E.g., photoreceptors have lightsensitive photopigments that regulate ion channels via second messenger (analogous to metabotropic synaptic receptors). Photoreceptors Transduction channels: (cGMP gated Na channel). “Dark Current”: Transduction channels open at rest, net inward Na+ current, depolarizes photoreceptor. When photons captured by pigment, then breakdown of cGMP is catalyzed, dark current decreases and cell hyperpolarizes. NSC201 Basic Neurobiology 11 Information Coded by Sensory Systems Modality: What form does the stimulus take? Intensity: How strong is the stimulus? Timing: When did the stimulus occur? Location: Where is the stimulus in the world, and what is it’s spatial relation to other stimuli? NSC201 Basic Neurobiology 12 Determinants of Sensory Modality Modality Hearing, Vision balance Taste Smell Proprioception touch, pain, temp Receptor Modality determined by Form of energy in the stimulus. Type of sensory receptor specialized to respond best to the form of energy in stimulus (“labelled line”). NSC201 Basic Neurobiology 13 Submodality Cones Each modality subdivides sensory “space” to extract different attributes of a stimulus. Examples: Taste: Taste receptors specialized for sweet, sour, salty, bitter, umami Vision: Photoreceptors express different photopigments selective for particular wavelengths of light. Hearing: Hair cells in cochlea are tuned for sound frequency by location and mechanical properties of the sensory epithelium. NSC201 Basic Neurobiology Rods Hair cells 14 Sensory Coding: Intensity SA Touch Receptor : (tonically active, slowly adapting) Electrophysiological response (i.e. change in receptor potential) is proportional to stimulus amplitude. Most basic code for amplitude is firing rate of sensory neurons or primary sensory afferents (but code may change centrally). Adaptation: Response often declines under constant stimulation (often interpreted as evidence that change is more important than steady-state). (Note slowing of firing rate over time in illustrated touch receptor) Stimulus Intensity (Skin Deflection) Intensity proportional to magnitude of energy in stimulus. - Response - Stimulus Time Firing rate encodes intensity NSC201 Basic Neurobiology 15 Sensory Coding: Time • Temporal properties of a stimulus are encoded by changes in the discharge pattern of peripheral sensory neurons. • Latency: time from stimulus onset to response (basis for reaction times). • Onsets and offsets: transient spiking. • Duration. • Stimulus “shape”. Pacinian Corpuscle • Pattern codes for mimicking stimulus features may be converted to a “rate/place” code centrally. NSC201 Basic Neurobiology stimulus •Phasic on and off response. •rapidly adapting 16 Sensory Coding: Spatial Location Receptive field: the region of “sensory space” over which a receptor integrates energy. Retinal circuitry underlying an “opponent” visual receptive field Skin innervation creating an opponent tactile receptive field NSC201 Basic Neurobiology 17 Explicit Space Mapping Physical space directly (explicitly) “mapped” by spatial pattern (topography) of activated receptors. Vision Optics of the eye project visual scene onto a 2D sensory epithelium (retina). Visual stimulus directly encoded by locus of stimulated photoreceptors. Somatosensation Stimulus location is encoded by location of sensory receptors on the body surface. NSC201 Basic Neurobiology 18 Implicit Space Mapping Hearing Sensory epithelium represents sound frequency, not space. Space perception derived from neural analysis of spatial cues (neural computation) Spatial cue disparities at the two ears generated by head (horizontal plane). Directionally-dependent spectral filtering of incoming sounds by the ears (vertical plane). Smell Sensory epithelium sensitive to odorants, not space. Space computed centrally based upon reconstruction of odor “trails”. Porter et al, Nat. Neuro. (2007) NSC201 Basic Neurobiology 19 Sensory Discrimination “Minimum Audible Angle” (MAA) Acuity: a form of “two-point discrimination” (Ernst Weber, 1795-1878): What is the minimal perceptible difference (just-noticeable difference or JND, difference limen) between two stimuli? NSC201 Basic Neurobiology 20 Acuity Acuity is related to Size of the receptive field i.e., diameter of the photoreceptor, relative to the projected image on the retina: Smaller is better. Spatial density (receptors/mm2) of receptors in the retina: higher is better. 400 receptors 3,600 receptors 14,400 receptors 160,000 receptors Examples: density of pixels on a TV screen or image sensor of a digital camera. NSC201 Basic Neurobiology 21