Lecture 13
Fundamental Response Properties of Receptor Cells
Overview of Sensory Receptors
Sensory receptors respond to stimuli with specific qualities that define their receptive fields.
The receptive fields of neurons, further downstream in sensory pathways, are often characterized as:
More complex than initial sensory receptors.
The neurons fire under a more restricted set of conditions.
Definitions
Receptive Field: The specific area or conditions under which a sensory neuron or receptor responds to stimuli.
Neurons in Sensory Pathways
Neurons that are further downstream in the sensory pathways also have their own receptive fields.
Example of Receptive Fields:
Photoreceptor X: Fires when detecting light from a specific region of visual space.
Complex cell Y: Fires only to an edge at a specific orientation moving in a specific direction.
Peripheral Receptive Field Sizes and Sensory Resolution
Peripheral receptive field sizes ultimately define sensory resolution.
Better sensory discrimination requires more neurons at all steps in the circuit to preserve this resolution, leading to an energy and space trade-off.
Example of Sensory Discrimination
Stimulus Effect on Skin:
Lips: Two distinct points are felt due to the density of sensory receptors.
Back: Only one point is felt due to lower receptor density.
Coverage of Sensory Space
Receptive fields vary across cells, ensuring that sensory space is well-covered by the receptor array.
Spatial Receptive Fields of V1 Neurons: The representation of sensory information is structured, ensuring a comprehensive coverage of the sensory input.
Topographic Mapping of Sensory Information
Higher-level sensory systems represent sensory information in an orderly (topographic) map.
This involves:
Ordered Spatial Representation: Reflects the organization of the sensory/motor world in the Central Nervous System (CNS).
Dimensions of the map can include real spatial arrangements in the environment (e.g., visual field) or other dimensions like sound frequency.
Somatotopic maps located in primary somatosensory and motor cortex demonstrate this orderly representation.
Brain Area Allocation for Sensory Processing
Proportional size of brain areas is dedicated to processing somatosensory information from different body regions.
High sensory resolution is linked to expanded representation in the CNS.
Percentage of Somatosensory Cortex Devoted to Body Parts:
Forepaw: 10.2%
Trunk/Tail: 13.5%
Chin: 7.5%
Hindpaw: 7.8%
Incisors: 30.8%
Buccal Cavity: 11.5%
Head Vibrissae: 18.8%
Determinants of Touch Resolution
Factors contributing to touch resolution:
Densely packed receptors with small receptive fields have higher resolution.
Sparsely packed receptors with large receptive fields may converge projections significantly.
Specifics of Somatosensation
Definition: Somatic (body) sensation, referred to as somatosensation, involves the sensation of the body's physical interactions with itself and the environment.
Components of Somatosensation:
Mechanoreception (detection of mechanical pressure or distortion):
Cutaneous Mechanoreceptors: Involved in the sense of touch.
Proprioceptors: Monitor position and load of body parts.
Baroreception: Pertains to blood pressure detection.
Thermoreception: Involves temperature detection.
Nociception: Involves pain detection (some nociceptors are mechanoreceptors, but not all).
Mechanotransduction in Somatosensation
Mechanotransduction refers to the process whereby mechanical forces cause opening of ion channels in the cell membrane, leading to receptor potentials.
Key Mechanisms:
Mechanoreceptor proteins typically open cation channels when the cell membrane is deformed, resulting in depolarizing receptor potentials.
Receptor Potential and Stimulus Strength
The magnitude of receptor potential depends on the forces applied to the mechanoreceptor's cell membrane:
Ion Channel Dynamics:
Closed Ion Channels: At rest.
Weak Stimulus: Membrane is stretched; some ion channels open, leading to a receptor potential.
Moderate Stimulus: Increased stretching causing greater receptor potential.
Strong Stimulus: Significant stretching, resulting in clearly defined receptor potentials that may lead to action potentials if threshold is exceeded.
Types of Cutaneous Mechanoreceptors
Various types of mechanoreceptors exist in the skin, each with distinct properties and responses to physical stimuli:
Meissner's Corpuscle: Rapidly adapting mechanoreceptor for touch and pressure.
Merkel's Corpuscle: Slowly adapting mechanoreceptor for touch and pressure.
Free Nerve Ending: Some are nociceptors and thermoreceptors; they are slowly adapting.
Pacinian Corpuscle: Rapidly adapting mechanoreceptor for vibration and deep pressure.
Ruffini Corpuscle: Slowly adapting mechanoreceptor for skin stretch.
Tonic vs. Phasic Receptors
Tonic and phasic receptors differ in how they respond to stimuli:
Tonic Receptors (e.g., Merkel disc, Ruffini ending): Produce action potentials during sustained stimuli.
Phasic Receptors (e.g., Meissner and Pacinian Corpuscles): Begin firing during stimulus onset, with adaptation that results in fewer action potentials upon continued stimulation.
Pain Detection and Pathways
Specialized nociceptive and non-nociceptive somatosensory pathways exist, typically with higher firing thresholds than other somatosensory cells.
Labeled lines encoding indicates that no stimulation of certain cells will elicit the sensation of pain; pathways are distinguished for different modalities.
Proprioception Overview
Proprioception refers to the sensation of position and load on different body parts.
Involves sensory neurons located in muscles and connective tissues, with ascending afferent and descending efferent projections contributing to the understanding of body position.
Structure of Skeletal Muscles
Skeletal muscles attach to and cause movements of our bones and are primarily composed of large, parallel cells called extrafusal muscle fibers.
Muscle spindles, intrafusal fibers along with mechanosensory neurons, encode the degree of muscle stretch and contraction (rate coding).
Functions of Proprioceptors
Proprioceptors provide significant sensory feedback for:
Informing voluntary circuits about body part positions.
Triggering involuntary skeletal muscle stretch reflexes.
Involving components that compare expected and actual motor outputs with feedback mechanisms localized likely to the cerebellum, aiding in compensatory movements.