Some over- lap of the conscious & unconscious What is proprioception?
Law of specific nerve energies (specific modalities): One type of receptor responds to one type of sensory modality Give an example. Receptors have various structures Ex: Pain receptor, Pacinian Auditory corpuscle (touch) receptor (hair cell) Dorsal root neuron = first order sensory neuron 1 Where does sensory information enter the CNS? Where are the synaptic relays? What is the final destination? (brainstem ~ pons/medulla) -Dorsal spinal cord medulla thalamus cortex -or directly to thalamus cortex (vision) -or directly to cortex (smell) What happens if the information never reaches the sensory cortex? Ex. Only to Thalamus = relay center We will learn the major sensory pathways in later slides. See some summaries below. How does convergence of neurons reduce the ability to locate a stimulus? Ex: reduces ability to distinguish two points (two-point discrimination) How does convergence confuse the brain about stimulus location? the major sensory tracts are the dorsal columns and the anterolateral tracts (spinothalamic tracts) Dorsal columns Dorsal columns Anterolateral cross in medulla cross in cord pressure, vibration pain, temperature proprioception, crude touch fine touch large, myelinated small fibers, fibers unmyelinated transmitter = transmitter = glutamate substance P Anterolateral tract Learn the above information and use that information along with the next slide to determine the effects of spinal lesions (injuries): Lesions of the dorsal columns on one side of the cord causes loss of specific senses (ex. pressure) on the same (ipsilateral) side below the lesion; however, dorsal columns cross above the cord in the medulla & lesions above the cross cause loss of sensation on the contralateral (opposite) side of the body. Because the anterolateral (spinothalamic) tracts cross in the cord immediately after receiving information, lesions of the anterolateral tracts in the cord cause loss of specific senses (ex. temperature) on the opposite (contralateral) side. See next slide and your text. Somatic sensory pathways (tracts) in spinal cord Anterolateral tracts cross in the cord and ascend on contralateral side See red tract Each side of the cortex receives sensations from the contralateral (opposite) side. Remember that both the dorsal columns and the anterolateral tracts eventually cross to the opposite side; therefore, lesions in the primary sensory cortex cause loss of sensation on the contralateral side of the body. Medulla Dorsal columns ipsilateral until medulla Cross in medulla black tract The cartoon-like image displayed on top of the brain is called a somatosensory homunculus. Note the upside-down arrangement of the sensory cortex: The top of the cortex receives input that originated in lower body areas as the foot. Only a small area of the sensory cortex is devoted to the trunk, and a large area receives sensation from the mouth and face. Receptive Fields The receptive field of a neuron is the area in the periphery (area of receptors) that receives signals and sends them to the neuron.
To define the receptive field of a neuron: Record from the neuron while stimulating various peripheral areas. When a signal is recorded, we can draw a map of the peripheral area (the receptive field) that received & sent signals. The figure below shows receptive fields of both the primary dorsal root neurons and the secondary neuron in the CNS. DRAW A LINE AROUND THE RECEPTIVE FIELD OF THE SECONDARY NEURON. Stimulate Record Record Annular Dorsal root receptive field neurons on skin Dorsal - column neuron - - inhibitory + - (-) interons - - (-)
-
An annular receptive field (off-center and on-surround) may result from the following connecting neurons
+ - + - (-) +
Complicated receptive fields - Ex. Annular Receptive field of one dorsal column neuron cell body located in the medulla oblongata. This neuron (pink below) generates spontaneous low-frequency tonic signals even when not stimulated. Pink neuron turned off or stimulated by dorsal root neurons. Recording = \ \ \ \ \ \ impulses if no stimulation to the skin (tonic signals). Recording = no impulses if press on the skin in () areas of skin. Recording = \\\\\\\\\\\\\\\\ if press on the skin in the (+) areas. Why is the annular receptive field designated On-center/Off-Surround? Recording here Stimulating here Important terminology: Generator potentials = Changes in receptor membrane potential Sensory stimulus graded changes in receptor voltage (generator potentials) Summation to threshold
Action potential in sensory nerve How are EPSPs and generator potentials similar & different?
Sensory Coding = mechanisms used to determine the following properties of a stimulus: Stimulus modality How does the brain know the type of stimulus (light, temperature, etc.)? Stimulus location How does the brain know the location of a stimulus? Stimulus intensity - How know stimulus strength? Stimulus duration How know the time a stimulus lasts? Coding for stimulus modality: If pain and temperature come from the same peripheral area, how does the CNS distinguish the type of stimulus? by label line coding - one receptor and its nerve fiber transmit only one type of sensation. Ex: if one touches a photoreceptor or the nerve from a photoreceptor, the patient detects light, not touch.
Coding for stimulus location (1) by the law of projection - if a sensory nerve is stimulate at any point along the nerve fiber, the CNS interprets the location of the stimulus at the receptor (sensations are projected to the site of the receptor). The brain area that receives the signal believes the stimulus came from the receptor. The law of projection and label-line coding help to explain the phantom limb. Coding for stimulus location (2) by timing of stimuli arriving at the brain (see Figure on next slide to understand how timing of signals enables us to determine the location of sounds).
Coding for stimulus location (3) by lateral inhibition See Figure at right to see how neurons in the direct path from the stimulus suppress signals from lateral neurons, which helps to locate the stimulus precisely. Ex. of location of stimulus by timing of signal arrival Coding for stimulus intensity If action potentials in sensory nerves are all-or none, how does the CNS know the difference between a strong stimulus and weak stimulus? by two mechanisms: frequency coding and population coding
Frequency coding - A stronger stimulus causes action potentials to be generated at a higher frequency (increased number of action potentials per unit time) weak stimulus | | | | strong stimulus |||||||||
Population coding - A stronger stimulus affects a larger population of receptors, a larger number of receptors
Sensory Adaptation- Failure to detect continuous, repetitive stimuli after some period of time. Touch is a rapidly adapting sensation, and pain is a slowly adapting sensation.
Tonic and phasic receptors Tonic receptors continuously generate action potentials (turned on continuously) and change the frequency of action potentials when there is a change in stimuli. Tonic receptors usually are slowly adapting. Phasic receptors respond (generate action potentials) only when when there is a change in intensity; if there is no change in intensity of a stimulus, phasic receptors rapidly adapt. In other words, phasic receptors turn off when there is no change in stimulus intensity. (Ex. smell) Examples of touch/pressure receptors and pain receptors (nociceptors) Although most pain fibers are unmyelinated, the small, fast pain fibers are myelinated. Note the picture below of 5 types of touch receptors & the free nerve endings that are pain receptors. The change of shape of touch receptors may cause voltage changes (generator potentials) that sum to threshold potential. Stimulated & Adapted Pacinian corpuscles Unstimulated Pacinian corpuscle What is the explanation for referred pain? Explain the diagram above. Is this divergence or convergence of sensory information? Remember: The brain refers to the area from which it usually receives signals. Modulation of incoming sensory information (1) by other sensory neurons -see next slide (Ex. touching or rubbing an area located close to a painful stimulus may reduce the signal in the pain neuron because the touch neuron activates a small inhibitory neuron that influences the pain pathway (next slide). If the inhibitory neuron has a synapse on the receptor of the incoming sensory neuron, it will exert postsynaptic inhibition of the sensory neuron. See next slide. Usually inhibitory neurons release gaba, but some inhibitory neurons release the opioid - enkephalin, which inhibits release of the neurotransmitter substance P from pain fibers (presynaptic inhibition - See slide from lecture prior to Exam 1). (2) Also, a motor neuron in the brain may send a signal to a small inhibitory neuron in the spinal cord that inhibits release of neurotransmitter by the sensory neuron (presynaptic inhibition) See next slide red descending pathway.
When would inhibition of pain fibers be beneficial to an animal? Descending motor neuron may modulate sensory neurons A nearby sensory neuron may modulate a sensory pathway The pain fiber releases what neurotransmitter? What spinal inhibitory neurotransmitter is released by inter-neurons that inhibit release of the pain neurons neurotransmitter? What other inhibitory neurotransmitter is released by inter-neurons? Explain pre- and post-synaptic inhibition? see slides before exam 1 Try to answer this example question:
1. If the dorsal column on the left side of the spinal cord is damaged, the patient will lose
(a) vibration on the left side below the injury
(b) vibration on the right side below the injury
(c) temperature sensation on the right side below the injury
(d) a and c
(e) b and c Special Senses
Chemoreceptive Senses: Smell and Taste
Olfactory bulb area of the brain receiving signals from the primary olfactory neurons see next slide; this area is proportionally larger in dogs and rats than in human beings. Olfactory receptors in the olfactory epithelium in the nose = metabatropic receptors linked to G proteins, which stimulate second messengers, which open ion channels. Secondary and higher order neurons project to the olfactory cortex, hippocampus, amygdala Unlike other neurons, olfactory receptors can divide to replace worn-out receptor cells. Because the axons of the newly formed cells must find their way to the olfactory bulb, the receptor cells also are a model for determining how developing neurons find the correct target. Taste
Many different signal transduction mechanisms enable the taste receptors to discriminate 5 taste modalities: sweet, salty, bitter, sour, and umami (responds to taste-enhancing stimuli).
Taste receptors are clustered in taste buds on the tongue. Receptors for salty and sour are ion channels (which ions? See next slide ) Receptors for the other tastes are linked to G-proteins.
Note in figure on next slide - All of the signal transduction mechanisms of the taste buds eventually increase intracellular calcium
Signal transduction mechanisms of the taste buds increase intracellular calcium release of neurotransmitter action potentials in primary gustatory neurons medulla secondary neurons in medulla thalamus tertiary neurons in thalamus gustatory cortex.
HEARING We characterize sound waves by pitch and loudness. Pitch is determined by the frequency of the sound waves measured in hertz (Hz, waves/sec). High frequency waves high-pitched sounds Low frequency waves low-pitched sounds.
Human ears sometimes detect frequencies 20 20,000 Hz. Most acute hearing in humans 1000 - 3000 Hz. Mice, dogs hear ultra-high frequency sounds and others, Elephants hear ultra-low frequency sounds not detected by humans.
Loudness (intensity) of sounds depends on amplitude of sounds waves - measured in bels or decibels (dB).
normal conversation ~ 60 dB 80 dB often damage sound receptors How does the frequency of sound waves in the right and left panels differ? Does the amplitude of the waves differ in the two panels?
What does this tell you about the pitch and loudness of the sounds graphed in the two panels? Graphs of sound waves illustrate the frequency & amplitude of the waves The inner ear contains (1) sound receptors in the cochlea, and (2) Receptors for equilibrium in the vestibular apparatus (the semicircular canals, and two otolith organs called the utricle and saccule).
The middle ear is the amplification area of the ear (tympanic membrane = eardrum and the ear bones).
outer ear = pinna - directs sound waves and aids with localizing sounds Cochlea and auditory (cochlear) nerve Cochlear Nerve
Medulla oblongata
Thalamus Auditory cortex Organ of Corti in the cochlea contains the hair cells shown on the next slide. These Hair cells = Sound Receptors. Hair cells in the cochlea These hair cells are sound receptors, just as rods and cones are photo- receptors. Sound waves cause vibrations of the tympanic membrane, ear bones, and fluids of the ear, which promotes action potentials in hair cells in the cochlea. Hair cells within the cochlea are the sound receptors. When stimulated, hair cells release neurotransmitter that binds to receptors on the auditory nerve endings. Auditory nerve action potentials activate medullary nuclei medial geniculate nucleus in the thalamus temporal cortex (auditory cortex). Note Path Hair cells bend backwards to close ion channels and turn off signals. Coding pitch = coding the frequency of sound waves: frequency coding: High frequency sounds create maximal displacement of the basilar membrane near the oval window and do not travel far into the cochlea. Low frequency sounds travel along the basilar membrane and reach peak amplitude at the apex of the cochlea base apex Coding the loudness of sound = coding the amplitude of the sound wave, which is coded by the number of action potentials generated. (High amplitude sounds (louder sounds) stimulate more action potentials per unit time). CAUTION: Do not confuse frequency of action potentials indicating loudness with frequency of sound waves = pitch.
One Unusual feature: Projection of auditory signals to the cortex differs from projection of most somatosensory information Auditory nerves from each ear project to both sides of the brain.
Coding location of sounds: The brain compares differences in the timing and intensity of sounds to locate a particular sound (e.g., sounds reach the two ears at slightly different times - see next slide) Coding location of a sound by timing of signal arrival Hearing Disorders: Nerve deafness: Examples include sensorineural hearing loss = damage to hair cells from loud noises and central hearing loss due to damage to the auditory nerve or the auditory cortex. These disorders are not improved significantly with a hearing aid. Conduction deafness (amplification problem) Ex: otosclerosis inflammation of the ear bones. Improved with a hearing aid. Weber-Rennie test can be used as a crude screen for the above types of deafness. All deaf patients cannot hear a tuning fork, but patients with conduction deafness notice improved hearing if the fork is placed on a jaw bone or on the forehead Why?. Auditory aphasia (lesion in the auditory association cortex preventing interpretation of sounds). Ex. Patient may hear voices as sounds but not realize that the sounds are voices. Equilibrium - vestibular system in the inner ear. semicircular canals - receptors for rotational acceleration within cristae otolith organs = utricle and saccule - receptors for linear acceleration and head position within maculae Vestibular apparatus is NOT pointing to the cochlea. It consists of utricle, saccule, and semi- circular canals. Cochlea has receptors for? All vestibular receptor cells are hair cells. Receptor areas of the semicircular canals = cristae consisting of [cupula = gelatinous masses + hair cells], which bend when the head is rotated (rotational acceleration). Receptor areas of the utricle and saccule = maculae which contain hair cells + otoliths (crystals embedded in gelatinous material), The otoliths move in response to gravity (linear acceleration) bending hair cells. Hair cells of the vestibular apparatus stimulate the vestibular nerve, which projects either directly to the cerebellum or to the vestibular nuclei of the medulla, which then send axons to the cerebellum, which is the major final destination of most vestibular information. In other words, the cerebellum integrates signals for equilibrium. Note path
Visual and proprioceptive information and vestibular signals are used by the cerebellum to coordinate equilibrium. not a good image of fovea see subsequent slides Retina interrupted by the optic nerve Blind spot Fig. 10-2-e. Know the cells of the retina & path of signals: Light Rods & Cones Signal Bipolar cells Ganglion cells Optic Nerve [amacrine and horizontal cells = modulating cells] Note that light must travel through other cells to reach the rods and cones. Then neuronal signals go in opposite direction through cells This figure has a better illustration of the fovea. The fovea is an area of high acuity vision (sharp vision). What anatomical feature allows sharp focus? See above Substitute this image for the ones in your book. This figure clarifies that the left field is what is seen by the left eye, and right field is what is seen by the right eye. Know that lateral outer fibers carry nasal field information and do not cross. Inner fibers carry temporal information and do cross. Know these lesions: # 2 Cutting one optic nerve loss of all vision from same eye. # 5 Cutting optic tract Loss of nasal field from same eye and temporal field from the other eye.
When cutting optic tract on one side, explain why you lose the temporal field from the opposite side. Clue involves fibers. Accomodation = adjusting the shape of the lens to keep objects in focus. Near response = adjustments to view objects near the eyes. Near responses include: accomodation, pupillary constriction and convergence of the visual axes. Different groups of ocular muscles are responsible for each component of the response. Accomodation is only one part of the near response Photoreceptors (rods and cones). Cones - most important for high-acuity (sharp) day vision and color vision. Rods - very sensitive to dim light, especially important for night vision. Both needed for dark adaptation. Signal transduction in photoreceptors Light change in Rhodopsin Activates Transducin cGMP GMP CLOSE Na+ channels Hyperpolarization of photoreceptors decreased release of Neurotransmitter
Darkness Depolarization of photoreceptors Caution: It is unusual for a stimulus to hyper- polarize Decreases the tonic inhibition of Bipolar cells Common Visual disorders
Look at the visual defects in the figure below. Recall that light should be focused on the retina. Myopia light is focused in front of the retina (near-sighted corrected with concave lens) Hyperopia light is focused behind the retina. (far-sighted corrected with convex lens) Retina Astigmatism = blurred image due to abnormal curvature of the cornea.
Presbyopia = poor accomodation due to aging. The lens hardens with aging, which limits changes in shape.
Abnormalities of color vision are usually inherited; can be detected with Ishihara plates Example: Do you see a number on the plate below? People with red/green color blindness may not see a number All dots may appear to be the same color or black or gray.