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Sensory Systems Picture: Rene Descartes (1596-1650)
Dr W Kolbinger, Sensory Systems (2009) 1
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Lecture Outline Common Plan of Sensory Systems
Four Sensory Receptor Classes Three Basic Processes in a Sensory Receptor Encoding of Four Stimulus Attributes Convergence, Divergence and Lateral Inhibition 2
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Common Plan of Sensory Systems
Pathway Perception Behavior Receptor Sensory systems have a common plan. They are designed to detect a physical stimulus (physical energy), such as light, sound, etc. This stimulus is “translated” into cell activity in sensory receptor cells or sensory receptor neurons. The scientific term for this “translation” process is signal transduction, another term for sensory receptors is therefore transducers. These receptors (when we talk about sensory systems, the term “receptors” usually refers to receptor cells or receptor neurons, rather than to receptor molecules) transmit the signal on to second order neurons, which are part of a sensory pathway. Most sensory pathways include the thalamus as a relay station. Finally, the encoded sensory information reaches specialized regions of the cerebral cortex. In general, most sensory cortex areas are located in the parietal, occipital, temporal, or insular lobes. The figure on the right, which has been originally drawn by Rene Descartes, a 17th century French philosopher, includes already all the essential elements of the common plan of sensory systems. In the cortex, sensory information reaches consciousness. Sensory information is interpreted there and may trigger behavior. The process of interpretation of sensory input is called perception. Some behavioral responses, such as reflexes, can be triggered sub-cortically, at the level of the spinal cord, for example. Stimulus Picture: Rene Descartes (1596–1650) 3
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Fourth-order sensory afferent neurons Third-order sensory afferent neurons Second-order sensory afferent neurons First-order sensory afferent neurons Sensory receptors
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Sensory Systems and their Receptors
Sensory Receptor Class Somatosensory System (touch, vibration, proprioception, pain and temperature) Nociceptors Visual System Vestibular System Auditory System Olfactory System Gustatory System Mechanoreceptors Thermoreceptors Chemoreceptors Photoreceptors Mechanoreceptors During the lecture series we will cover the six sensory systems, the somatosensory system, the visual system, the vestibular system, the auditory system, the olfactory system (smell) and the gustatory system (taste). The olfactory system and the gustatory system together are also called the chemical senses. The six sensory systems and their Receptors are shown in the table . You realize that all sensory systems together function with a total of four basic sensory receptor classes, mechanoreceptors, which are sensitive to mechanical stimuli, thermoreceptors, which are sensitive to temperature, chemoreceptors, which are sensitive to chemical ions or molecules and photoreceptors, which are sensitive to light. Mechanoreceptors Chemoreceptors Chemoreceptors 5
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Examples of Sensory Receptors
Sensory receptor neuron(somatosensory and olfactory systems Sensory receptor cell(visual, taste, and auditory systems 6
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1. Sensory receptors A: Free nerve endings (pain, temperature)
B C D A: Free nerve endings (pain, temperature) B: Pacinian corpuscle (pressure) C: Meissner’s corpuscle (touch) D: Muscle spindle (stretch) Mechanoreceptors are activated by pressure or changes in pressure. Mechanoreceptors include, but are not limited to, the pacinian corpuscles in subcutaneous tissue, Meissner's corpuscles in nonhairy skin (touch), baroreceptors in the carotid sinus (blood pressure), and hair cells on the organ of Corti (audition) and in the semicircular canals (vestibular system). Photoreceptors are activated by light and are involved in vision. Chemoreceptors are activated by chemicals and are involved in olfaction, taste, and detection of oxygen and carbon dioxide in the control of breathing. Thermoreceptors are activated by temperature or changes in temperature. Nociceptors are activated by extremes of pressure, temperature, or noxious chemicals.
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Three Basic Processes in Different Components of a Sensory Receptor
Cell body transduction site synaptic Terminal “graded potentials” 2) Action potentials The components of a sensory receptor in general are the cell body (soma, perikaryon), the transduction site, where the signal transduction takes place, an axon, and a synaptic terminal where the signal is transmitted, usually to the second order neuron of the sensory pathway. As an example, the diagram below shows a typical sensory receptor of the somatosensory system, a Pacinian corpuscle, which is, like all somatosensory receptors, a pseudounipolar neuron. Three basic processes occur in different parts of the sensory receptor: Receptor Potential Receptor potentials are generated in the transduction site of a sensory receptor. These receptor potentials are “graded potentials”, which means they do not follow the all-or-nothing rule. They are more like “analog signals”, with variable durations and intensities. Action Potentials In case the receptor potential is above threshold, action potentials are generated in a defined area of the axon, which is called the trigger zone. Action potentials are then conducted, first along the distal part of the axon, followed by the proximal part of the axon until they reach the synaptic terminal. Please note that in this example of a sensory receptor neuron, the action potentials are not generated at the axon hillock, as it occurs for example in multipolar motor neurons. Transmitter Release Transmitter release takes place at the synaptic terminals of the sensory receptor. The transmitter released into the synaptic cleft will then bind to receptors (in this case receptor molecules) in the postsynaptic membrane of the postsynaptic cell, the second order neuron of the sensory pathway. Not all sensory receptors are receptor neurons, like the one shown above. Some sensory receptors, such as haircells, which are the mechanoreceptors of the vestibular and the auditory systems, are receptor cells. These receptor cells, instead of being of neuronal origin, are specialized epithelial cells. Their apical pole is still the transduction site, but they lack an axon, and they don’t produce action potentials. Instead, the graded receptor potential directly induces the release of a signaling substance (transmitter) at the base of this cell. 1) Receptor potential 3) Transmitter release 8
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Encoding of Four Stimulus Attributes
Modality labeled lines Intensity Amplitude of graded receptor potentials Frequency code of action potentials Duration Mechanisms depending on receptor adaptation Location Concept of receptive fields and other mechanisms Four attributes characterize a sensory stimulus: Modality refers to the physical type (or quality) of stimulus energy. This might be light (for the visual system), sound (for the auditory system, touch (for the somatosensory system), etc. Intensity refers to the amplitude (or quantity) of a stimulus. Duration is defined by the time between the start and the end point of the stimulus. Location refers to the place where the stimulus is located or originated. This might be some place in our three dimensional environment around us (as for light, or sound), or a place on the surface of our body (as for touch). 9
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Encoding of Stimulus Intensity
Sensory Transduction and Receptor Potentials 1. The environmental stimulus interacts with the sensory receptor and causes a change in its properties 2. receptor potential or generator potential. 3.receptor potentials are graded in amplitude
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Encoding of Stimulus Intensity
Encoding of Stimulus Intensity starts at the transduction site of a sensory receptor. Low stimulus intensity causes only minor changes in the polarity of the receptor, i.e. causes small graded receptor potentials. Sub-threshold stimuli may well induce small receptor potentials, without inducing any action potentials. Frequency Code Once receptor potentials are above threshold, action potentials are induced. Higher stimulus intensities induce higher receptor potentials, which in turn lead to a generation of higher numbers of action potentials per time unit. This primary encoding mechanism for stimulus intensity, which is demonstrated in the diagram above, is called the frequency code of individual axons. Population Code refers to the fact that high intensity stimuli can activate more individual axons than low intensity stimuli. Think about increasing the pressure on your skin. The more you press, the larger the area and the number of axons that are stimulated.
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Encoding of Duration: Different Strategies
Slowly adapting receptors remain active for the duration of a stimulus Rapidly adapting receptors are active only during times of changes (on/off) Encoding of Stimulus Duration Encoding of stimulus duration is another feature of sensory systems. It has its origin again at the receptor level. We all know from every day experience that a while after the onset of a sensory stimulus, the sensation induced by the stimulus may disappear, until it is no longer recognized. This process takes already place at the receptor level and is called receptor adaptation. For example while you dress in the morning, sensory information caused by your clothes touching the skin help you in the process. During the day, you no longer feel the touch of your clothiers, although you are still dressed and the fabric touches your skin. Sensory receptors differ widely in their adaptation rate. We differentiate between slowly adapting receptors and rapidly adapting receptors. Ideally, slowly adapting receptors remain depolarized and produce increased rates of action potentials for the whole duration of the stimulus. The duration of the stimulus is indicated by the time the frequency of action potentials is elevated. Slowly adapting receptors are important for regulatory functions, when a physiological parameter has to be kept at a certain value. Rapidly adapting receptors, on the other hand, may only signal the onset of the stimulus, as shown in the lower portion of the diagram. The receptor potential quickly returns to baseline values and no action potentials are generated after this initial phase, although the stimulus is still on with the same intensity. In reality, it is rare that sensory receptors act as simple. Rapidly adapting receptors often have a basal discharge in the absence of a stimulus. Their firing rate may increase during the onset of the stimulus, it may return to the basal rate during the duration of the stimulus, and the offset of the stimulus might cause a short drop in firing rate. As a consequence, slowly adapting receptors are better in constantly monitoring levels of stimulation, whereas rapidly adapting receptors are most sensitive to changes, not to constant stimulation. slowly adapting receptors are better in constantly monitoring levels of stimulation, whereas rapidly adapting receptors are most sensitive to changes, not to constant stimulation. 12
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Encoding of Location: Receptive Field of a Sensory Receptor
Stimulation Recordings 1 1 2 2 Encoding of Stimulus Location The way a stimulus is localized may vary in different sensory systems and we will focus on mechanisms for the somatosensory system, for the visual system and for the auditory system later in the lecture series. One basic principle I like to introduce at this point is the concept of a receptive field of a sensory receptor. The diagram on the right shows a schematic drawing of a somatosensory touch receptor neuron (pseudounipolar neuron) innervating a certain area of the skin. The square on the left of the diagram depicts a part of the body’s surface. A recording electrode measures the action potentials at one point along the axon and the recordings are shown on the right hand side. When the skin is touched at the tip of arrow # 1, the recording shows baseline activities. When the skin is touched at # 2, the number of action potentials increases. You would repeat the experiment until you could outline the whole area where a stimulus would induce a response in this neuron. The area is then called the receptive field of this neuron. A receptive field defines an area of the body that when stimulated results in a change in firing rate of a sensory neuron 13
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Convergence and Divergence
convergence answers the question “Where does the information come from?” divergence answers the question “Where does the information go to?” 14
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lateral inhibition Receptive fields can be excitatory or inhibitory.
The areas of inhibition contribute to a phenomenon called lateral inhibition, and aid in the precise localization of the stimulus by defining its boundaries and providing a contrasting border
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Somatosensory System Lecture Outline
Five Modalities and their Receptors Different Fibers for Different Receptors Segmental Spinal Nerves and Dermatomes Pathways for Different Modalities Clinical Correlations 16
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Modalities of the Somatosensory System
Touch (discriminative touch) Vibration Proprioception Pain (nociception) Temperature most of the somatic sensory modalities refer to sensations of the skin, proprioception refers to sensory receptors originating in the skeleto-muscular system. The somatic sensory system comprises of five different modalities: touch, vibration, proprioception, pain and temperature. Whereas most of the somatic sensory modalities refer to sensations of the skin, proprioception refers to sensory receptors originating in the skeleto-muscular system. All receptors of the somatic sensory system are pseudo-unipolar neurons. Their sensory endings can be found in skin, or in (or close to) the muscle. Their fibers run in peripheral nerves and their cell bodies are located in ganglia (dorsal root ganglia in case the fibers run in spinal nerves or in cranial nerve ganglia). All receptors of the somatic sensory system are pseudo-unipolar neurons. Their sensory endings can be found in skin, or in (or close to) the muscle. Their fibers run in peripheral nerves and their cell bodies are located in ganglia (dorsal root ganglia in case the fibers run in spinal nerves or in cranial nerve ganglia). 17
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Touch is transduced by Merkel’s disks (discriminative touch) and Ruffini’s endings (skin stretch). Vibration is transduced by Meissner’s corpuscles (for lower frequencies of about 50 Hz) and Pacinian corpuscles (for higher frequencies of about 300 Hz). Pain (pricking pain by rapidly adapting mechano-sensitive or thermo-sensitive receptors, burning pain by slowly adapting polymodal receptors) and Temperature (cold receptors and warm receptors) are transduced by free nerve endings.
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Golgi tendon organs are
Muscle spindles are embedded in extrafusal fibers of the working musculature of the muscle. The primary receptor of a muscle spindle is a rapidly adapting receptor with a Ia ("one a") afferent fiber. It carries sensory information of muscle stretch. This receptor forms the afferent limb of the myotatic reflex (deep tendon reflex), The secondary receptor of a muscle spindle is a slowly adapting receptor carrying sensory information of muscle length. It uses a class II ("two") afferent fiber. Golgi tendon organs are positioned close to the border between muscle and tendon. Their Ib afferent fibers form the afferent limb of the reverse myotatic reflex (inverse myotatic reflex).
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Afferent Fiber Classification
12-20 μm 6-12 μm 1-6 μm 0.2-1 μm A alpha A beta A delta C I II III IV m/sec 36-72 m/sec 4-36 m/sec 0.4-2 m/sec Three types of primary afferent fibers (with cell bodies in the dorsal root ganglia) mediate cutaneous sensation: (1) large myelinated A and A fibers that transmit impulses generated by mechanical stimuli; (2) small myelinated A fibers, some of which transmit impulses from cold receptors and nociceptors that mediate pain and some of which transmit impulses from mechanoreceptors; and (3) small unmyelinated C fibers that are concerned primarily with pain and temperature. However, a few C fibers also transmit impulses from mechanoreceptors. 20
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Afferent Fiber Classification and Somatosensory Modalities
From skin: From muscle: 12-20 μm 6-12 μm 1-6 μm 0.2-1 μm Touch, Vibration Proprioception Pain, Temperature 21
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Segmental Organization of Spinal Nerves and Dermatomes- segmental organization of the sensory innervation of the skin
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Pathway for Touch, Vibration, Proprioception: Dorsal Column / Medial Lemniscus System
Touch, vibration and proprioception are carried in a pathway called the dorsalcolumn / medial lemniscus system 23
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Topographical Organization of the Spinal Cord
Dorsal column From arm From trunk From leg Midline 24
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Spatial Orientation of Signals from Different Parts of the Body
in Somatosensory Area Somatosensory area has a high degree of localization of the different parts of the body
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Sensory Neurons: Two-Point Discrimination
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Pathway for Pain and Temperature: Anterolateral System
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Topographical Organization of the Spinal Cord
ALS From arm From trunk From leg Midline 28
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Topographical Organization of the Spinal Cord
Touch, vibration, proprioception from left side of body L T A Right Left L T A Pain, temperature from right side of body 29
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Lissauer’s Tract and the Anterolateral System
Right Left 30
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Chief Complaint: Left Leg Paralysis, Right Leg Numbness
History: A 75 year old retired pastry maker was in good health until about one year ago, when he started to develop gait difficulties and numbness of his right leg. Recently, he also experienced urinary urgency with occasional incontinence. Though he started taking laxatives, he experienced problems with bowel movements. His wife reported that he also had stiffness in the legs bilaterally. At times, the left leg has unexpectedly not been able to support his body weight for a brief period, causing him to stumble to maintain balance. He also reported that in addition to the numbness in the right leg, he also has a constant tingling feeling in the same limb, which he describes as “intolerable.” General Examination: Normal vital signs. Patient has no significant cardiovascular history. Abdomen was soft and non-tender. No palpable abdominal masses. Digital rectal examination showed significantly reduced muscle tone in the external sphincter and weakness of voluntary contraction. Prostate was felt to be enlarged with a highly nodular, irregular surface. Neurological examination: Patient was fully alert and oriented x 3. Cranial nerve exam was unremarkable. Upon motor examination, the upper extremities had normal strength, bulk, and tone, and the reflexes were 2+ throughout the C5 to C8 spinal level. In the lower extremities, however, the muscle tone was increased in left leg and the left iliopsoas muscle was weaker than the right (4/5). The muscle bulk in both legs was normal. Reflexes in the right leg were 2+, knee jerk on the left leg was 3+, and ankle jerk was 4+. Plantar response was extensor the left and flexor on the right. Finger to nose and heel to shin testing was normal. Pinprick testing and temperature sensation showed decreased sensitivity on the right side of the trunk below the umbilicus. Vibration and joint position sense was significantly reduced in the left leg and foot.
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Brown Sequard Syndrome
X X X X 32
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Visual System Lecture Outline
Structures of the Eye Refraction and Image Formation Visual Acuity Autonomic Control of Pupil Diameter Clinical Correlations Dr W Kolbinger, Visual System (2009) 33 33
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Anatomic Considerations
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The Ocular Fundus Optic Fovea disc Macula
The diagram on the right superimposes the passage of light originating from the point of fixation and from the blind spot region of the visual field on the underlying anatomical structures. Light originating from the point of fixation is projected onto the fovea. Light originating from the blind spot region of the visual field ends up in the optic disk region of the retina. Since the optic disk region does not contain photoreceptor, the visual information of this region is lost. Most of the time we are not aware of the loss of visual information, since our brain “completes” the picture with the “most likely” visual information. The optic disc region itself only contains axons of retinal ganglion cells, the output elements of the retina, but it lacks photoreceptors. As a consequence, the optic disc is responsible for the blind spot, a region inside the boundaries of the visual field, where we don’t receive visual information. 35
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Optics of the Eye Cornea refractive power: 42 D
Rounded lens refractive power: 26 D Flat lens refractive power: 13 D Refractive Power of an Optical System When light rays pass from air into another medium, such as water (or human tissue, which also has a high water component), or a glass lens for example, they change their direction. This interaction between light and its environment is called refraction. When incoming parallel light rays pass through a convex lens, all light rays after the passage will focus at a defined distance from the midline of the lens. This distance is called focal distance. The focal distance depends on the material and on the curvature of the lens. Refractive power is the reciprocal value of the focal distance in meters. Its unit of measurement is called diopter, abbreviated as “D” (plural: diopters). In the human eye, the cornea is responsible for the main refractive power of the eye of about 42 D, whereas the lens is responsible for the modulation of refractive power (refractive plasticity) during accommodation. Plasticity: 13 D 36
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Accommodation Far Vision
Ciliary muscle relaxed Flat lens refractive power: 13 D Suspensory ligaments tightened During far vision, light rays originating from a distant object can be considered (almost) parallel. The focus of light rays during far vision largely depends on the high refractive power of the cornea (42 D) and only a small refractive power of the flat lens (13 D). During far vision, the ciliary muscle, which is a circular muscle around the lens, is relaxed. The diameter of this circular structure is large, causing tightening of the suspensory ligaments (zonule fibers). As a consequence, these ligaments pull on the lenses’ equator, thereby flattening it and minimizing its refractive power. Focus on the Retina Accommodation Adjusts the Refractive Power of the Eye 37
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Accommodation Near Vision Ciliary muscle constricted
Rounded lens refractive power: 26 D Suspensory ligaments floppy During near vision incoming light rays can no longer be considered parallel. Therefore, a stronger refractive power is needed to focus the light rays on the retina. This is achieved by constriction of the circular ciliary muscle, which reduces its’ diameter and relaxes the suspensory ligaments. The lens follows its own elasticity and gets a more rounded (more convex) shape, which increases its refractive power to about 26 D in a young individual. The ciliary muscle is activated by parasympathetic fibers of the autonomic nervous system (ANS). Cell bodies of the first order efferent parasympathetic neurons are located in the upper midbrain, in the Edinger-Westphal nucleus of cranial nerve III. Together with the somatic efferent fibers innervating the extraocular muscles, the preganglionic fibers originating in the midbrain form the oculomotor nerve (CN III). The preganglionic parasympathetic fibers synapse on second order neurons in the ciliary ganglion. Postganglionic fibers originating in this ganglion form the short ciliary nerves which innervate the ciliary muscle. Focus on the Retina 38
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Presbyopia Near Vision Flat lens Blurred picture on the Retina
The variability of the refractive power of the lens between far vision (13 D) and near vision (26 D) is called refractive plasticity. Unfortunately, the lens looses its elasticity during aging, thereby reducing the ability to focus on near objects, a condition called presbyopia. Flat lens Blurred picture on the Retina 39
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