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CHAPTER 9 THE SENSORY ORGANS 王俊 副教授 博雅楼 402室 [email protected] Section 1 General Principles of Sensory Organs 一、Definitions and Classifications Receptors: structures that monitor both external and internal environmental conditions and conduct information about those stimuli to the central nervous system. 环层小体 触觉小体 肌梭 Pacinian copuscle Messner’s corpuscles Muscle spindle Receptor Classification by Location Exteroreceptors Respond to stimuli arising outside the body; Include touch, pressure, pain, and temperature receptors in the skin and most receptors of the special senses (vision, hearing equilibrium, smell, taste) Interoceptors Visceroceptors: Found in internal viscera and blood vessels Proprioceptors: Found in skeletal muscles, tendons, joints, ligaments Receptor Classification by Stimulus Type Mechanoreceptors – respond to touch, pressure, vibration, stretch, and itch Thermoreceptors – sensitive to changes in temperature Photoreceptors – respond to light energy (e.g., retina) Chemoreceptors – respond to chemicals (e.g., smell, taste, changes in blood chemistry) Nociceptors – sensitive to pain-causing stimuli 二、General Characteristics of Sensory Receptors 1. Adequate Stimulus 2. Transduction 3. Coding 4. Adaption Adequate stimulus Specific sensory R are more sensitive to certain types of stimuli (photo R in eye). The types of stimulus to which the R is most sensitive (has the lowest threshold) is called adequate stimulus. Transduction Receptor detects a stimulus and convert the energy (light, pressure) of stimulus into electrical signals (action potentials) in afferent nerve. During the transduction, receptor respond to a stimulus by generating a series of graded potentials in their associated afferent fiber, not action potentials directly. The response of the pacinian corpuscle to an adequate stimulus is called a generator potential. It is able to "generate" action potentials in the region of the receptor cell that has voltage-sensitive channels. However, not all receptors have voltage-sensitive channels. In receptor cells incapable of generating action potentials (typically specialized receptors such as the cones and rods of the retina), the change in membrane potential in response to an adequate stimulus is referred to as a receptor potential. The membrane of the receptor region is, however, electrically inexcitable; it contains no voltage-gated ionic channels and does not generate spikes. If the receptor region generated action potentials, the graded nature of the generator potential would be destroyed because as soon as the generator potential exceeded the critical firing level an action potential would be initiated, reversing the membrane polarization no matter how large or small the stimulus, i.e., the membrane potential would no longer encode the stimulus intensity. Coding The speed of conduction and other characteristics of sensory nerve fibers vary, but action potentials are similar in all nerves. This raises the question of why stimulation of a touch receptor causes a sensation of touch and not of warmth. It also raises the question of how it is possible to tell whether the touch is light or heavy. Adaption When a maintained stimulus of constant strength is applied to a receptor, the frequency of the action potentials in its sensory nerve declines over time. This phenomenon is known as adaption. 快适应感受器 慢适应感受器 举例 皮肤触觉感受器 肌梭、颈动脉窦、关节囊感 受器 特点 仅在刺激开始后的短时间内有 传入冲动发放,以后虽然刺激 仍在作用,但其传入冲动的频 率却很快降低到零 刺激持续作用时,一般仅在 刺激开始后不久出现冲动频 率的轻微降低,以后可以较 长时间维持在这一水平。 意义 对刺激变化十分灵敏,适于传 递快速变化的信息,有利于机 体探索新异的物体或障碍物, 有利于感受器和中枢再接受新 的刺激 有利于机体对某些功能状态 进行长时间持续的检测,并 根据其变化随时调整机体的 功能。 Section 2 Visual Function of the Eye 760 一、The light-Refracting System of the Eye (一)Optical Property of the Light-Refracting System of the Eye (二)Refraction of Light in the Eye and Reduced Eye AB Bn = ab nb (三)Accommodation Far point of vision : farthest point to the eye at which an object can be brought into clear focus without accomodation Near point of vision :nearest point to the eye at which an object can be brought into clear focus by accomodation 1. Accommodation of the Lenses 2. Accommodation of the Pupils (1.5~8mm) Pupillary accommodation reflex/Near reflex of pupil: is the reduction of pupil size in response to an object coming close to the eye. 强光--视网膜--中脑顶盖 前区--动眼神经缩瞳核-- 动眼神经副交感神经--瞳孔 括约肌收缩--瞳孔缩小 Pupillary light reflex: is the reduction of pupil size in response to light. consensual light reflex: each pupil constricts with light shone into one eye. 3. Convergence Convergence reflex: is the simultaneous inward movement of both eyes toward each other, usually in an effort to maintain single binocular vision when viewing an object far to near. (四)Defects in Accommodation and Light Reflex emmetropia ametropia myopia hyperopia astigmatism myopia hyperopia astigmatism (五)Aqueous humor Aqueous humor circulation non-pigmented epithelium of the ciliary body—posterior chamber— anterior chamber— trabecular meshwork—Schlemm's canal—vein Function Maintains the intraocular pressure and inflates the globe of the eye. Provides nutrition for the avascular ocular tissues; posterior cornea, trabecular meshwork, lens, and anterior vitreous. 二、Functional Anatomy of the Retina and Two Photoreceptor System (一)Functional Anatomy of the Retina Pigment epithelia cells Photoreceptor cell layer bipolar cells ganglion cells 1 色素上皮层 Pigment epithelia cells 2 光感受器细胞层 Photoreceptor cell layer Rod cell 视杆细胞 Cone cell 视锥细胞 Synaptic terminal 3 双极细胞和神经节细胞 bipolar cells and ganglion cells Physiological blind spot is the place in the visual field that corresponds to the lack of light-detecting photoreceptor cells on the optic disc of the retina where the optic nerve passes through it. Since there are no cells to detect light on the optic disc, a part of the field of vision is not perceived. (二)Retinal Photoreceptor System Rods system, Scotopic vision system Scotopic vision not color sensitive, more sensitive to light and mainly peripheral (rods) Cones system, Photopic vision system Photopic vision color sensitive, less sensitive to light and mainly central (cones) Rods Cones 1.2x108 6x106 Vision in shades of gray Color vision High sensitivity Low sensitivity Low acuity High acuity Night vision Day vision Much convergence in retinal pathways Little convergence in retinal pathways More numerous in periphery Concentrated in fovea (三)Mechanism of Photoreception of the Rod 1、Photochemistry of Rhodopsin Rhodopsin(500nm) opsin retinal VitA Rhodopsin light opsin + retinal 2、Generation of Receptor Potential Dark current is the depolarizing current, carried by Na+ ions, that flows into a photoreceptor cell when unstimulated. GP:-30~-40mv 超极化-感受器电位 GC Ca2+ Ca2+ (四)The Cone System and Color Vision Rhodopin opsin retinal 视锥细胞的特点是它具有辨别颜色的能力。波长只要 有3-5nm的增减,就可被视觉系统分辨为不同的颜色 760 Trichromatic Theory 三 、Information Process in the Retina 四、Visual Pathway 五、Other Visual Phenomena (一)Visual Acuity Visual acuity refers to the clarity or clearness of one’s vision, a measure of how well a person sees. This may be thought of as the ability of the eye to see fine detail. Visual acuity test:Landolt C Snellen eye chart Landolt 环 Landolt C Snellen eye chart (二)Dark and Light Adaptation Dark Adaptation It occurs when going from a well light area to a dark area. Initially blackness is seen because our cones cease functioning in low intensity light. Also, all the rod pigments have been bleached out due to the bright light and the rods are initially nonfunctional. Light Adaptation This occurs when we move from the dark into bright light. The bright light momentarily dazzles us and all we see is white light because the sensitivity of the receptors is set to dim light. Rods and cones are both stimulated and large amounts of the photopigment are broken down instantaneously, producing a flood of signals resulting in the glare. (三)Visual Field The visual field is the total area in which objects can be seen in the peripheral vision while the eye is focused on a central point. (四)After Image and Fusion Phenomenon An image continuing to appear in one‘s vision after the exposure to the original image has ceased. One of the most common afterimages is the bright glow that seems to float before one’s eyes after staring at a light bulb or a headlight for a few seconds. The phenomenon of afterimages may be closely related to persistence of vision, which allows a rapid series of pictures to portray motion, which is the basis of animation and cinema. (五)Binocular Vision and Stereopsis Function of Binocular Vision Expand visual field Function of Binocular Vision Stereopsis Section 3 Function of the Auditory System 一、Auditory Threshold and Auditory Sensation Area F: 20—20000Hz, P: 0.0002—1000dyne/cm2 Fs: 1000--3000Hz, Language: 300—3000Hz 二、Functions of the External Ear Pinna (ear lobe): collects sound waves helps to distinguish the direction of sound Exteral auditory canal: sound waves conduction air pressure increase 一端封闭的管道对于波长为其长度4倍的声波能产生 最大的共振作用。 L=2.5cm λ=4L=0.1m λ = ν /ƒ=340/3000=0.1m 三、Functions of the Middle ear 1.Supercharging: 2.Protection: 四、Air and Ossicular Conduction Bone conduction Air conduction Sound waves Sound waves Vibration of temporal bone Exteral auditory canal Tympanic membrane Middle ear bones Air in the drum Oval window Round window cochlea cochlea Vibration of endolymph in cochlea 五、Functions of Cochlea (一)Anatomy of Cochlea helicotrema Scala vestibuli Reissner’s membrane Scala media basilar membrane Scala tympani Cochlea • A light microscopic image of a hair cell isolated from tissues normally surrounding it. • This hair cell is about 30 m long; the hair bundle is about 5 m wide. • Hair cells convert mechanical energy into an electrical energy through ion channels which open and close in sync with sound vibrations. •This slide from (A.J. Hudspeth, Science, 230:745-752, 1985) (二)Vibration of Basilar Membrane and Traveling Wave Theory Cochlear Representation of Sound basilar membrane • Frequency Theory Membrane vibrates at frequency of Sound source • Place Theory High Frequency sounds cause vibration near oval window Low frequency sounds cause vibration near helicotrema Place Theory Different parts of the basilar membrane respond most to different frequencies: high frequency at base; low frequency at apex. Cochlear Mechanics Actual Response to high frequency Passive Response Passive response caused by mechanical properties of membranes (e.g. width) Active Response is caused by outer hair cells, which inject energy back into vibration of cochlear membranes • Outer hair cells move (tilt slightly and change length) in response to sound (frequency specific). • The movement pushes on the basilar membrane which amplifies and sharpens its response a given location along the membrane. (三)Transduction Function of Hair cells 基底膜振动 ? 毛细胞 DISPLACEMENT OF THE BASILAR MEMBRANE TOWARD THE SCALA VESTIBULI (UPPER) RESULTS I DEPOLARIZATION OF THE OHC - ASSOCIATED WITH DECREASE OF OHC LENGTH DISPLACEMENT IN THE OPPOSITE DIRECTION PRODUCES HYPERPOLARIZATION AND AN INCREAS IN OHC LENGTH Inner hair cell Outer hair cell Cell number 3500 20000 Nerve innerved 90—95% 5—10% (四)Electrophysiology of the Cochlea 1. Endocochlear or endolymphatic potential +80mv -80mv 0mv 2. Cochler microphonic potential, CM 六、Action Potential in Auditory Nerves 七、Hearing Pathway Section 4 The Vestibular Sensation Of Equilibrium 一、Vestibular Apparatus and Adequate Stimulus -80mv -60mv -120mv 半规管壶腹脊的适宜刺激是正负角加速度 椭圆囊和球囊的适宜刺激是直线加速度运动 utricle saccule 二、Vestibular Reactions and Nystagmus (一) vestibular postural reflex (二)vestibular autonomic reactions (三)nystagmus 三、Equilibrium Pathway