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1/23/2016 Receptors The Special Senses Introduction Vision General Properties of Receptors • Specialized structures designed to respond to specific stimuli • Variable complexity General Properties of Receptors • Transducers – Stimulus energy is converted to an AP • 2 types of graded potentials generated by receptors – Receptor potential • Occurs in a separate receptor cell (not the neuron) • Example: Rods and cones use phototransduction – Generator potential Stimulus causing receptor potentials ⇓ Generator potential in afferent neuron ⇓ Nerve impulse • Occurs when either: – The neuron is the receptor » Example: neurons in olfactory epithelium – The receptor releases NT and stimulates the sensory neuron Sensation and Perception Stimulatory input General properties of receptors • Information conveyed by receptors – Modality • Type of stimulus or sensation – hearing, vision, taste, etc. – Location • Brain refers pain to its point of origin – Intensity • Determined by impulse transmission rate, variations in sensory thresholds in a group of sensory neurons Conscious level = perception – Duration • Encoded by changes in the pattern of impulse transmission over time Detection of stimulus = sensation 1 1/23/2016 Sensory Adaptation Classification of Receptors • Stimulus Modality Chemoreceptors Thermoreceptors Nociceptors Mechanoreceptors Photoreceptors • Reduction in rate of impulse transmission when stimulus is prolonged • Occurs in all receptors except those for pain • Let’s try it… Classification of Receptors • Stimulus Modality Chemoreceptors Respond to chemical concentrations Odors, tastes, dissolved chemicals, etc Thermoreceptors Nociceptors Mechanoreceptors Photoreceptors Classification of Receptors • Stimulus Modality Chemoreceptors Thermoreceptors Nociceptors Free nerve endings Respond to tissue damage Give the perception of pain Mechanoreceptors Photoreceptors Classification of Receptors • Stimulus Modality Chemoreceptors Thermoreceptors Respond to heat and cold Nociceptors Mechanoreceptors Photoreceptors Classification of Receptors • Stimulus Modality Chemoreceptors Thermoreceptors Nociceptors Mechanoreceptors Respond to physical deformity Touch pressure, stretch, vibration, tension Photoreceptors 2 1/23/2016 Classification of Receptors • Stimulus Modality Classification of Receptors • Origin of stimuli Chemoreceptors Thermoreceptors Nociceptors Mechanoreceptors Photoreceptors Exteroceptors Interoceptors Proprioceptors Respond to light Rods, cones, certain retinal ganglion cells Classification of Receptors • Origin of stimuli Classification of Receptors • Origin of stimuli Exteroceptors Exteroceptors Interoceptors Respond to stimuli that originate outside the body • Examples: light, smell, sound Respond to stimuli that originate inside the body • Examples: pressure, hunger, thirst Interoceptors Proprioceptors Proprioceptors Classification of Receptors • Origin of stimuli Exteroceptors Interoceptors Proprioceptors Respond to stimuli that originate due to the positions and tensions of joints and muscles Special senses • • • • Vision Hearing Olfaction Gustation 3 1/23/2016 Vision Introduction Vision Introduction • 70% of all sensory receptors are in the eye • Nearly half of the cerebral cortex is involved in processing visual information • Optic nerve is one of body’s largest nerve tracts • The eye is a photoreceptor organ • Refraction – Light bends when it passes from one medium to another • Conversion (transduction) of light into AP’s • Information is interpreted in cerebral cortex Eyebrow Orbicularis oculi muscle Eyebrow Eyelid Eyelashes Site where conjunctiva merges with cornea Palpebral fissure Palpebral conjunctiva (covers inside of eyelids) Cornea Lateral commissure Eyelashes Bulbar conjunctiva (covers outer surface of eye) Eyelid Conjunctival sac Medial commissure Orbicularis oculi muscle (a) Surface anatomy of the right eye (b) Lateral view; some structures shown in sagittal section Figure 15.1a Figure 15.1b Accessory Structures Lacrimal sac • Lacrimal apparatus – Series of structures that produce tears, direct their flow, and drain them • Lacrimal gland and ducts – Connect to nasal cavity • Lacrimal secretion (tears) – Dilute saline solution » Mucus, antibodies, and lysozyme – Blinking spreads the tears toward medial commissure – Drain into the nasolacrimal duct Lacrimal gland Excretory ducts of lacrimal glands Lacrimal punctum Lacrimal canaliculus Nasolacrimal duct Inferior meatus of nasal cavity Nostril Figure 15.2 4 1/23/2016 Accessory Structures Superior oblique muscle Superior oblique tendon Superior rectus muscle • Six extrinsic eye muscles Lateral rectus muscle Inferior rectus Inferior oblique muscle muscle (a) Lateral view of the right eye Figure 15.3a the Eyeball Muscle Lateral rectus Medial rectus Superior rectus Inferior rectus Inferior oblique Superior oblique Action Moves eye laterally Moves eye medially Elevates eye and turns it medially Depresses eye and turns it medially Elevates eye and turns it laterally Depresses eye and turns it laterally Controlling cranial nerve VI (abducens) III (oculomotor) III (oculomotor) III (oculomotor) III (oculomotor) IV (trochlear) • Wall of eyeball contains three layers (tunics) –Fibrous –Vascular –Sensory (retinal) (c) Summary of muscle actions and innervating cranial nerves Figure 15.3c the eyeball • Three layers 1) Fibrous tunic Sclera Cornea • Sclera – “white” of the eye • Cornea – clear continuation over the iris and pupil Scleral venous sinus (AKA canal of Schlemm) Fibrous Tunic Figure 15.4a 5 1/23/2016 the eyeball • Cornea must be actively dehydrated to remain transparent • Sodium pumps remove sodium from the endothelium – water follows the eyeball • Three layers 2) Vascular tunic (uvea) – 3 components –Choroid –Ciliary body –Iris Corneal Edema the eyeball • Three layers the eyeball • Three layers 2) Vascular tunic (uvea) – 3 components 2) Vascular tunic (uvea) –Choroid –Choroid –Ciliary body » Highly vascular » Darkly pigmented - Absorbs stray light » Ciliary processes - Arranged in a ring around the lens » Ciliary muscle - Used to focus the eye –Ciliary body –Iris –Iris the eyeball • Three layers 2) Vascular tunic (uvea) – 3 components –Choroid –Ciliary body –Iris - Radial muscle fibers contract = pupil dilation - Circular muscle fibers contract = pupil constriction Ciliary body Ciliary zonule (suspensory ligament) Sclera Choroid Iris Pupil Cornea Lens Scleral venous sinus VascularTunic Figure 15.4a 6 1/23/2016 the eyeball the eyeball • Three layers • Rods and cones 3) Retina (innermost layer) – – – – • Photoreceptor neurons – Rods and Cones • Bipolar neurons • Ganglion neurons – Axons form the optic nerve – Exit eyeball at optic disc Ciliary body Ciliary zonule Specialized photoreceptors Rods = black and white vision Cones = daylight color vision Fovea centralis • Cones most concentrated here • Point of highest visual acuity • Other retinal layers are displaced (forms a depression) Ganglion cells Sclera Choroid Bipolar cells Photoreceptors • Rod • Cone Retina Cornea Iris Pupil Macula lutea Fovea centralis Optic nerve Lens Scleral venous sinus Central artery and vein of the retina Optic disc (blind spot) Amacrine cell Horizontal cell Pigmented Pathway of signal output layer of retina Pathway of light (b) Cells of the neural layer of the retina Retina Figure 15.4a Retinal Blood Supply Figure 15.6b Pathway of light • Outer 1/3 of retina Neural layer of retina – Choroid • Inner 2/3 of retina Pigmented layer of retina Choroid Sclera – Central artery and vein Optic disc Central artery and vein of retina Optic nerve (a) Posterior aspect of the eyeball Figure 15.6a 7 1/23/2016 Blood supply to the retina The Eyeball The eyeball • Photoreceptors • Photoreceptors – Rods – Cones • Red, blue, and green – Highest density in macula lutea • More numerous at peripheral region • Dim light » Concentrated in fovea centralis – Indistinct, fuzzy, non-color peripheral vision • Operate in bright light • High-acuity color vision • About 100 million per eye The eyeball Central artery and vein emerging from the optic disc • Lens – – – – Macula lutea Optic disc Biconvex, transparent, flexible Attached to ciliary body by suspensory ligaments Allows precise focusing of light on the retina Forms a partition • Creates an anterior and posterior cavity Retina Figure 15.7 8 1/23/2016 The eyeball Emmetropic eye (normal) • Two Cavities – Anterior cavity Focal plane • Filled with aqueous humor – Modified plasma – Constantly produced, in constant circulation – Accumulation may lead to glaucoma Focal point is on retina. Figure 15.14 (1 of 3) Iris Lens epithelium Lens Cornea Corneal epithelium Corneal endothelium Aqueous humor Anterior Anterior segment chamber Posterior (contains chamber aqueous humor) Scleral venous sinus 1 Aqueous humor is Cornealformed by filtration from the capillaries in scleral junction the ciliary processes. 2 Aqueous humor flows from the posterior chamber through the pupil into the anterior chamber. Some also flows through the vitreous humor (not shown). 3 Aqueous humor is reabsorbed into the venous blood by the scleral venous sinus. The eyeball Posterior segment (contains vitreous humor) 2 • Two Cavities Ciliary zonule (suspensory ligament) Ciliary body 3 Bulbar conjunctiva Sclera 1 Ciliary processes • Filled with vitreous humor – Gelatinous mass – Avascular – Very few cells » A few phagocytes that remove debris from the field of vision – Responsible for eye holding its spherical shape Ciliary muscle Cornea – Posterior cavity Lens Figure 15.8 The Physiology of Vision • Visual acuity – ability to discern detail – Ratio of distances • 20/20 • 20/40 – Less visual acuity • 20/15 – More visual acuity The Physiology of vision • 5 processes produce a visual image 1. Refraction 2. Accommodation 3. Pupil constriction 4. Convergence 5. Photoreception 9 1/23/2016 Refraction Refraction • The bending of light rays • Most of the light we see is reflected off of objects Refraction Refraction • This light begins as a single point, but spreads as distance from the object increases • The eye must collect and re-bend the light to a single point in order to construct an image • Since focal length is fixed, the eye changes the shape of the lens to accomplish this Focal length is fixed in the eye Refraction Emmetropic eye (normal) Focal plane • Light passing through a convex lens (as in the eye) is bent so that the rays converge at a focal point • The image formed at the focal point is upside-down and reversed right to left Focal point is on retina. Figure 15.14 (1 of 3) 10 1/23/2016 Refraction Sympathetic activation Nearly parallel rays from distant object • Lens Refracting media • Light is refracted by – Cornea – Aqueous humor – Lens – Vitreous humor Ciliary zonule Inverted image Ciliary muscle • Change in lens curvature – Allows for fine focusing of an image (a) Lens is flattened for distant vision. Sympathetic input relaxes the ciliary muscle, tightening the ciliary zonule, and flattening the lens. Figure 15.13a Accommodation Sympathetic activation Nearly parallel rays from distant object • Eye is adapted for distance vision Lens • Ciliary muscle is relaxed = lens is flat As light moves closer greater curvature required • Ciliary muscle contracts = lens is curved Ciliary zonule Inverted image Ciliary muscle (a) Lens is flattened for distant vision. Sympathetic input relaxes the ciliary muscle, tightening the ciliary zonule, and flattening the lens. Figure 15.13a Parasympathetic activation Divergent rays from close object Accommodation Inverted image • Near vision – Light rays from near objects require extensive refraction – Accommodation • Ciliary muscles contract • Lens shape changes • Eye strain (b) Lens bulges for close vision. Parasympathetic input contracts the ciliary muscle, loosening the ciliary zonule, allowing the lens to bulge. Figure 15.13b 11 1/23/2016 Accommodation • Near point – Closest point at which an object can be brought into focus • Typically 25cm – Maximum bulge – Presbyopia - Aging Close Vision • Close vision requires –Accommodation –Pupil constriction • Prevents most divergent light rays from entering the eye –Convergence • Medial rotation of the eyeballs toward the object being viewed (binocular vision) Pupil Manipulation • Contraction of the circular muscles = constriction • Contraction of the radial muscles = dilation Photoreception • Sensory transduction – Light energy is converted into nerve impulses • Takes place in retina – Photoreceptors 12 1/23/2016 Process of bipolar cell Photoreception • Rods and cones –Outer segment of each contains visual pigments Synaptic terminals Rod cell body Rod cell body Cone cell body Nuclei Outer fiber Mitochondria Pigmented layer Outer segment Inner segment • Molecules that change shape as they absorb light Inner fibers The outer segments of rods and cones are embedded in the pigmented layer of the retina. Connecting cilia Apical microvillus Discs containing visual pigments Discs being phagocytized Pigment cell nucleus Melanin granules Basal lamina (border with choroid) Figure 15.15a In the dark Na+ Ca2+ Rod discs In the dark, cGMP opens sodium channels and depolarizes photoreceptors Photoreceptor cell (rod) Ca2+ Inhibitory neurotransmitter is released (glutamic acid) Note that they are connected to each other here Bipolar cells are constantly inhibited in the dark Visual pigment consists of • Retinal • Opsin Bipolar cell Ganglion cell (b) Rhodopsin, the visual pigment in rods, is embedded in the membrane that forms discs in the outer segment. Figure 15.15b Breakdown of rhodopsin causes closing of sodium channels 11-cis-retinal 2H+ Oxidation Vitamin A 11-cis-retinal Rhodopsin Reduction 2H+ 2 Regeneration of the pigment: Enzymes slowly convert all-trans retinal to its 11-cis form in the pigmented epithelium; requires ATP. Figure 15.18 (1 of 2) Dark Light In the light 1 cGMP-gated channels are closed, so Na+ influx stops; the photoreceptor hyperpolarizes. Light 1 Bleaching of the pigment: Light absorption by rhodopsin triggers a rapid series of steps in which retinal changes shape (11-cis to all-trans) and eventually releases from opsin. Photoreceptor cell (rod) 2 Voltage-gated Ca2+ channels close in synaptic terminals. 3 No neurotransmitter is released. In the presence of light bipolar cells are not inhibited 4 Lack of IPSPs in bipolar cell results in depolarization. Bipolar cell Ca2+ Ganglion cell Opsin and All-trans-retinal 5 Depolarization opens voltage-gated Ca2+ channels; neurotransmitter is released. 6 EPSPs occur in ganglion cell. 7 Action potentials propagate along the optic nerve. All-trans-retinal Figure 15.16 Figure 15.18 (2 of 2) 13 1/23/2016 Photoreception • Excitation of cones – Method of excitation is similar to that of rods – There are three types of cones • Named for the colors of light absorbed: blue, green, & red • Light absorbed depends upon opsins present Photoreception • Light adaptation – Occurs when moving from darkness into bright light • Large amounts of pigments are broken down instantaneously – Produces glare • Pupils constrict • Dramatic changes in retinal sensitivity – Rod function ceases • Cones and neurons rapidly adapt – Visual acuity improves over 5–10 minutes Photoreception Photoreception • Dark adaptation – Occurs when moving from bright light into darkness • • • • The reverse of light adaptation Cones stop functioning in low-intensity light Pupils dilate Rhodopsin accumulates in the dark – Retinal sensitivity increases within 20–30 minutes • Duplicity theory – Why do we have 2 kinds of photoreceptors? • Neural circuits for night are not suited for day – Rods:Bipolar cell • 600:1 • Spatial summation of large field = poor detail – Cones:Bipolar cell • 1:1 • Best for detail in small receptive fields Visual Pathway to the brain Fixation point • Axons from medial portion of each retina cross at optic chiasma – Continue as optic tracts • Optic radiation fibers connect to the primary visual cortex in the occipital lobes – Visual input from both eyes is interpreted by brain • Distance • Depth perception Optic nerve Suprachiasmatic nucleus Pretectal nucleus Lateral geniculate nucleus of Thalamus – primary connection of optic nerve to occipital lobe Superior colliculus Optic chiasma Optic tract Uncrossed (ipsilateral) fiber Crossed (contralateral) fiber Optic radiation Occipital lobe (primary visual cortex) The visual fields of the two eyes overlap considerably. Note that fibers from the lateral portion of each retinal field do not cross at the optic chiasma. Figure 15.19a 14 1/23/2016 Abnormalities of Vision • Myopia (nearsightedness) Abnormalities of Vision • Hyperopia (farsightedness) – Close objects seen clearly – Image is focused in front of the retina – Correction = concave lens – Distant objects seen clearly – Image is focused behind the retina – Correction = convex lens Abnormalities of vision • • • • • • • • • Abnormalities of vision • Astigmatism – Abnormal curvature of eye – Light rays don’t meet at a common focus – Causes visual distortion Astigmatism Detached retina Conjunctivitis Glaucoma Cataract Diplopia Night blindness Color blindness Macular Degeneration Abnormalities of vision • Detached retina – Retina becomes separated from underlying choroid – Causes loss of vision in affected area 15 1/23/2016 Abnormalities of vision • Conjunctivitis – Inflammation of the conjunctiva – Causes by bacteria, viruses, irritants, allergies – “Pink eye” Abnormalities of vision • Glaucoma – – – – Increased pressure in the eyeball Leads to optic nerve damage Causes gradual loss of sight Damage is irreversible, but may be slowed or stopped Abnormalities of vision • Cataract – Lens becomes progressively opaque – Results in blurred vision – Can be surgically removed and replaced with an artificial lens Abnormalities of vision • Nightblindness (nyctalopia) – Loss of low light vision – Most common cause: retinitis pigmentosa • Rods lose ability to respond to light Abnormalities of vision • Diplopia – Double vision – At least 20 different causes Cranial nerve III palsy Aneurysm High blood pressure Diabetes Inflammatory conditions Guillain-Barre syndrome Problems with the lid, pupil, or eye muscles • Etc. • • • • • • • Abnormalities of vision • Colorblindness – Defect in perception of colors – Defect in cones – Can be acquired; usually genetic 16 1/23/2016 Abnormalities of vision • Macular degeneration – Damage to macula and retina – Risk factors • Genetic • Lifestyle Abnormalities of vision • Bonus: amblyopia – – – – “Lazy eye” Decreased vision in eye that appears normal Developmental problem of the brain Brain doesn’t recognize image from one eye 17