海水鱼趋光特性的电生理研究——Ⅰ.蓝圆鲹、鲐鱼视网膜电图的一般特性

1.记录和分析了我国主要中上层趋光鱼——蓝圆鲹、鲐鱼在位完整眼的视网膜电图(ERG)。ERG 显示典型的混合型视网膜的特征。ERG 对缺氧极敏感,在阻断循环水几分钟后,b 波迅速减小,留下PIII 和c 波。2.暗视b 波振幅与刺激强度的关系可用非线性方程V/V_(最大)=I~a/(I~a+K)描述。其中V为相应于光强Ⅰ时b 波振幅;V_(最大)为b 波最大振幅;a 和K 均为常数。3.在强光明适应后,鲐b 波阈值在1小时内回复至暗视水平,但鲹却极缓慢,整个暗适应进程超过3小时。4.在明适应过程中,b 波振幅有两种不同类型的变化:当背景光强低时,随适应时间振幅逐渐下降,但当背景光强超过一定水平(40微瓦/平方厘米)后,却随适应时间而增大,这可能是明视系统的适应特点。5.鲹、鲐辨增阈曲线显示相似的特点:当背景光强低时,光强的变化并不引起辨增阈的明显变化,但超过一定水平后两者就是线性关系。不同波长的曲线随着背景光强的增加呈现交叉的趋势,表明存在着几种光谱敏感度不同的感受系统。     

鲫鱼(Carassius auratus)视网膜敏感度在明适应过程中的变化

在明适应过程中,随着适应光强的不同,鲫鱼视网膜电图b波的敏感度呈现两种不同类型的变化,分别定名为D型(下降型)和R型(回复型)。当适应光较弱时为D型——敏感度随适应时间逐渐降低,在几分钟内达到稳定。但当适应光强超过一定水平后(1.95×10~(-6)瓦/平方厘米,网膜水平),即为R型——敏感度一开始急剧降低(5～8对数单位),随后逐渐回复,回复的时间进程近似为指数函数。适应光愈强,回复的幅度愈大,达到稳定的时间愈长,当适应光强为1.95x10~(-4)瓦/平方厘米时,分别为2.3对数单位和30分。在中间的适应水平是一种过渡的类型,兼有D型和R型的某些特点。对不同适应水平时b波的光谱敏感特性、辨增阈、振幅-强度曲线和闪光融合频率的分析表明,网膜活动由杆细胞系统向锥细胞系统过渡的明适应水平恰与由D型变化向R型变化的水平相当,这一事实强烈地提示,D型和R型变化分别表示杆细胞系统和锥细胞系统的适应特性。R型变化基本上与网膜内色素颗粒的漂移无关,可能归因于锥细胞活动以及它从杆细胞的抑制中解脱出来。对若干有关的工作进行了讨论。     

鲫鱼(Carassius auratus)视网膜敏感度在明适应过程中的变化

在明适应过程中,随着适应光强的不同,鲫鱼视网膜电图b波的敏感度呈现两种不同类型的变化,分别定名为D型(下降型)和R型(回复型)。当适应光较弱时为D型——敏感度随适应时间逐渐降低,在几分钟内达到稳定。但当适应光强超过一定水平后(1.95×10~(-6)瓦/平方厘米,网膜水平),即为R型——敏感度一开始急剧降低(5～8对数单位),随后逐渐回复,回复的时间进程近似为指数函数。适应光愈强,回复的幅度愈大,达到稳定的时间愈长,当适应光强为1.95×10~(-4)瓦/平方厘米时,分别为2.3对数单位和30分。在中间的适应水平是一种过渡的类型,兼有D型和R型的某些特点。对不同适应水平时b波的光谱敏感特性、辨增阈、振幅-强度曲线和闪光融合频率的分析表明,网膜活动由杆细胞系统向锥细胞系统过渡的明适应水平恰与由D型变化向R型变化的水平相当,这一事实强烈地提示,D型和R型变化分别表示杆细胞系统和锥细胞系统的适应特性。R型变化基本上与网膜内色素颗粒的漂移无关,可能归因子锥细胞活动以及它从杆细胞的抑制中解脱出来。对若干有关的工作进行了讨论。     

NO对鲫鱼视网膜视杆和视锥信号传递的不同作用

在离体、灌流的鲫鱼视网膜上,考察了一氧化氮(NO)对视网膜电图(ERG)和水平细胞的作用.NO的供体硝普钠(SNP)压抑暗视ERGb波,却增大明视b波,这些作用均可为血红蛋白所阻遏.进而,SNP使视杆水平细胞对光反应减小,而增强L型视锥水平细胞的对光反应.这些结果表明,NO在外视网膜可压抑视杆系统的活动而增强视锥系统的活动.此外,可溶性鸟苷酸环化酶的抑制剂亚甲基蓝对视杆、视锥水平细胞的作用恰与SNP作用相反,这提示NO的作用可能由cGMP所介导.对NO的作用位点提出了工作假设.     

海水鱼趋光特性的电生理研究——Ⅲ.鲐鱼视顶盖诱发电位的一般特性和光谱敏感性

1.记录和分析了鲐鱼中脑视顶盖诱发电位(VETP)。波形基本特征同淡水鱼。2.在暗视条件下,光强和振幅间关系无简单规律可循,但峰潜伏期和光强关系满足关系式log(IT)=αlogⅠ+β,其中IT 为峰潜伏期;Ⅰ为刺激光强;α、β为常数。在明视时,此关系式在一定光强范围内依然成立。3.不同强度明适应时和暗适应过程中峰潜伏期的变化表明,决定峰潜伏期的主要因素是刺激光强,和网膜的适应状态关系不大。4.VETP 的暗适应曲线和ERG 的相同。5.VETP 暗视光谱敏感曲线和杆细胞视色素吸收光谱吻合得很好,提示在暗视条件下VETP 单一地反映了杆细胞的活动。在背景光强达4.07微瓦/平方厘米(角膜水平),短波段相对敏感度降低,长波段升高,但峰值并未向长波段位移。结合ERG 的结果推测,在更高背景光强时发生光谱敏感性的浦肯野位移是可能的。     

鲫鱼视网膜明、暗适应状态NADPH黄递酶染色的比较

在鲫鱼视网膜上考察了明、暗适应两种状态下视网膜细胞的NADPH黄递酶染色的不同.在暗适应5h的视网膜中,视杆和视锥的内段、部分水平细胞呈深度染色.部分无长突细胞和神经节细胞也有染色.此外,Muller细胞对NADPH黄递酶染色是强阳性.在明适应的视网膜中,光感受器的内段、神经节细胞的染色则较浅,而Muller细胞和水平细胞则呈阴性反应.此外,视杆去极化型双极细胞则清楚地被染色.无长突细胞在两种状态下的染色没有显著差异.Muller细胞,部分水平细胞和视杆去极化型双极细胞中存在诱导型一氧化氮合成酶,其诱导与否取决于视网膜的适应状态.     

真鲷视网膜结构及其视觉特性研究：Ⅱ视网膜光感受细胞超微结构研究

对真鲷光感受细胞的超微结构进行观察,结果表明：视杆外段膜盘为游离膜盘,视锥外段膜盘则为连续的膜结构,视锥和视杆均含有连接纤毛和辅助外段。花萼状突起起源于内段。椭体内充满线粒体,无球状小体。双锥椭圆体并生膜为六层,视锥内段无鳍状突起,视锥突触带,在明适应视网膜中数量增多,在暗适应视网中数量减少,视杆突触带在这两种适应网膜中数量不变,每一杆小球只有一个突触带,而锥小足有４－６个突触带。     

急性低氧对鲫鱼视网膜视杆和视锥信号的不同影响

在保持完整血液循环的鲫鱼眼杯标本上,应用Ａｇ－ＡｇＣｌ电极记录视网膜电图（ＥＲＧ）,研究了急性低氧下不同适应状态ＥＲＧ反应变化的情况,以期分析视锥与视杆通路对急性低氧的敏感性是否不同。结果表明：１．急性低氧对明视ＥＲＧ－ｂ波的影响要远远快于对暗视ｂ波的影响,这说明视锥信号通路比视杆信号通路对缺氧敏感;２．在间视状态下,ＥＲＧ的ｂ波在低氧开始反几分钟内有一个明显的增大过程,而在明视或暗视中皆未观察到     

观察和思考

视觉的二元说(Duplicity Theory of Vision)是视觉生理的一块重要基石。根据这一理论,脊椎动物视网膜中二类形态不同的光感受器——视杆和视锥分别在不同的光强水平实施     

低浓度钴离子选择性压抑鲫鱼视网膜中视杆信号的传递

低浓度(10～30μmol/L)钴离子(Co ²⁺)选择性压抑鲫鱼视网膜中视杆信号的传递,在分离、灌流的鲫鱼视网膜,10μmol/L Co ²⁺超极化视杆水平细胞,并完全压抑其对光反应,但对视锥水平细胞并无影响.此外,低Co ²⁺几乎完全压抑暗视视网膜电图(ERG)b波,但不影响明视b波,低Co ²⁺并不影响光感受器电位(PⅢ),也不影响视杆水平细胞谷氨酸受体的反应能力.其它2价金属离子(Co ²⁺,Mn²⁺)并不显示相似的选择性压抑作用,但谷氨酸转运蛋白的阻遏剂(β-OH-Asp)显示相似的作用,提示低Co ²⁺的作用可能与谷氨酸重摄取的压抑有关.     

Adaptation in the Ventral Eye of Limulus is Functionally Independent of the Photochemical Cycle, Membrane Potential, and Membrane Resistance

The early receptor potential (ERP), membrane potential, membrane resistance, and sensitivity were measured during light and/or dark adaptation in the ventral eye of Limulus. After a bright flash, the ERP amplitude recovered with a time constant of 100 ms, whereas the sensitivity recovered with an initial time constant of 20 s. When a strong adapting light was turned off, the recovery of membrane potential and of membrane resistance had time-courses similar to each other, and both recovered more rapidly than the sensitivity. The receptor depolarization was compared during dark adaptation after strong illumination and during light adaptation with weaker illumination; at equal sensitivities the cell was more depolarized during light adaptation than during dark adaptation. Finally, the waveforms of responses to flashes were compared during dark adaptation after strong illumination and during light adaptation with weaker illumination. At equal sensitivities (equal amplitude responses for identical flashes), the responses during light adaptation had faster time-courses than the responses during dark adaptation. Thus neither the photochemical cycle nor the membrane potential nor the membrane resistance is related to sensitivity changes during dark adaptation in the photoreceptors of the ventral eye. By elimination, these results imply that there are (unknown) intermediate process(es) responsible for adaptation interposed between the photochemical cycle and the electrical properties of the photoreceptor.     

THE DARK ADAPTATION OF THE COLOR ANOMALOUS MEASURED WITH LIGHTS OF DIFFERENT HUES

Determinations of minimum light thresholds as a function of time in the dark have been made for four color normal, three deuteranopic (or deuteranomalous), and four protanopic (or protanomalous) subjects. Measurements were made with red, reddish orange, yellow, green, violet, and white test lights. Dark adaptation curves for the deuteranopes and deuteranomalous are essentially identical with those of the color normal for all colors. The cone portions of the protanopic dark adaptation curves measured with the red, reddish orange, yellow, and white lights are higher than the corresponding data for the color normal, the discrepancy between the two sets of data decreasing from the long to short wave lengths. Dark adaptation curves for the protanopes and protanomalous measured with green and violet light are essentially normal in appearance. A theoretical explanation is advanced to account for these findings in terms of the known sensitivity characteristics of the normal and color-anomalous eye.     

The Rat Electroretinogram : I. Contrasting effects of adaptation on the amplitude and latency of the b-wave

Characteristics of the electroretinogram (ERG) produced by the essentially all rod eye of the rat are presented as functions of the number of quanta absorbed by each rod per stimulus flash. The ERG's were obtained with 1.5 msec. stimulus flashes and uniform illumination of the entire retina. Under these conditions, distortions in the ERG due to stray light are minimized, and the ERG more accurately reflects the activity of its retinal sources. The effects of background light and two forms of dark adaptation were studied and compared. The results, especially for the b-wave, permit an interpretation in terms of two distinct processes. One process appears to determine the b-wave latency. This process is almost independent of the state of adaptation of the retina. The other process does not affect the latency, but determines the b-wave threshold and amplitude. This process strongly depends upon the state of adaptation. Moreover, the effects of dark adaptation on this amplitude-determining process are almost identical with the effects of background light.     

Activity of long-wavelength cones under scotopic conditions in the cyprinid fish Danio aequipinnatus

In carp (Cyprinus) and goldfish (Carassius), long-wavelength cones are reported to be active under scotopic conditions. Using the electroretinogram (ERG), we tested another cyprinid fish, Danio aequipinnatus, which contains A₁-based visual pigments and for which we had previously measured the spectral sensitivities of individual cones. Dark adaptation curves show a rod/cone break at about 45 min. When thoroughly dark-adapted, the spectral sensitivity function is broader than can be accounted for by self-screening of rhodopsin, but it can be modeled by an additive combination of rods and the 560-nm cones. Dim, red background light causes adaptation of rods and a broadening of the spectral sensitivity function, which can be simulated by increasing the proportion of cones in the model. Brighter red backgrounds adapt the 560-nm cones. Because of the effect of red adapting lights, the ERG evidence for the participation of long-wavelength cones close to visual threshold appears to be different in Danio than in the goldfish Carassius. Accepted: 14 June 1997     

Effect of blocking the Na+/K+ ATPase on Ca2+ extrusion and light adaptation in mammalian retinal rods.

Membrane current and light response were recorded from rods of monkey and guinea pig by means of suction electrodes. The correlation between adaptation and the Na+/K+ pump was investigated by measuring light-dependent changes in sensitivity with and without inhibition of Na+/K+ ATPase by strophanthidin. Strophanthidin was found to reduce the dark current, to slow the time course of the photoresponse, and to increase light sensitivity. At concentrations between 20 and 500 nM, the pump inhibitor suppressed in a reversible way the current re-activation occurring during prolonged illumination and modified the light-dependent decrease in sensitivity, which in control conditions approximates to a Weber-Fechner function. The effects of the pump inhibitor on the adaptive properties of rods are associated with an increased time constant of the membrane current attributed to the operation of the Na+:Ca2+,K+ exchanger. The effects of rapid application of the pump inhibitor on the current re-activation are consistent with the idea that significant changes in the internal sodium occur in rods of mammals during background illumination and that they play an important role in the process of light adaptation.     

Visual Adaptation in the Retina of the Skate

The electroretinogram (ERG) and single-unit ganglion cell activity were recorded from the eyecup of the skate (Raja erinacea and R. oscellata), and the adaptation properties of both types of response compared with in situ rhodopsin measurements obtained by fundus reflectometry. Under all conditions tested, the b-wave of the ERG and the ganglion cell discharge showed identical adaptation properties. For example, after flash adaptation that bleached 80% of the rhodopsin, neither ganglion cell nor b-wave activity could be elicited for 10–15 min. Following this unresponsive period, thresholds fell rapidly; by 20 min after the flash, sensitivity was within 3 log units of the dark-adapted level. Further recovery of threshold was slow, requiring an additional 70–90 min to reach absolute threshold. Measurements of rhodopsin levels showed a close correlation with the slow recovery of threshold that occurred between 20 and 120 min of dark adaptation; there is a linear relation between rhodopsin concentration and log threshold. Other experiments dealt with the initial unresponsive period induced by light adaptation. The duration of this unresponsive period depended on the brightness of the adapting field; with bright backgrounds, suppression of retinal activity lasted 20–25 min, but sensitivity subsequently returned and thresholds fell to a steady-state value. At all background levels tested, increment thresholds were linearly related to background luminance.     

THE INFLUENCE OF LIGHT ADAPTATION ON SUBSEQUENT DARK ADAPTATION OF THE EYE

The course of dark adaptation of the human eye varies with the intensity used for the light adaptation which precedes it. Preadaptation to intensities below 200 photons is followed only by rod adaptation, while preadaptation to intensities above 4000 photons is followed first by cone adaptation and then by rod adaptation. With increasing intensities of preadaptation, cone dark adaptation remains essentially the same in form, but covers an increasing range of threshold intensities. At the highest preadaptation the range of the subsequent cone dark adaptation covers more than 3 log units. Rod dark adaptation appears in two types—a rapid and a delayed. The rapid rod dark adaptation is evident after preadaptations to low intensities corresponding to those usually associated with rod function. The delayed rod dark adaptation shows up only after preadaptation to intensities which are hundreds of times higher than those which produce the maximal function of the rods in flicker, intensity discrimination, and visual acuity. The delayed form remains essentially constant in shape following different intensities of preadaptation. However, its time of appearance increases with the preadaptation intensity; after the highest preadaptation, it appears only after 12 or 13 minutes in the dark. These two modes of rod dark adaptation are probably the expression of two methods of formation of visual purple in the rods after its bleaching by the preadaptation lights.     

长时间暗适应和弱背景光对鲤鱼视网膜视锥水平细胞的影响——形态和生理的相关研究

本工作利用电子显微镜及细胞内记录技术研究了在暗适应和弱背景光下鲤鱼视锥与水平细胞间突触部位超微结构及视锥水平细胞光反应的变化。在长时间暗适应(>2h)后,视锥水平细胞对光反应受到强烈压抑,其突起末端则外形光滑、圆钝。在施加弱背景光(15min)后,这些细胞的反应显著增大,其末端出现大量深陷于视锥小足内部的刺形结构,这种刺形结构增加了视锥与水平细胞间的突触传递效率,可能是视锥水平细胞光反应性增高的形态学基础。     

Light-induced reduction in excitation efficiency in the trp mutant of Drosophila

In the transient receptor potential (trp) mutant of Drosophila, the receptor potential appears almost normal in response to a flash but quickly decays to baseline during prolonged illumination. Photometric and early receptor potential measurements of the pigment suggest that the pigment is normal and that the decay of the trp response during illumination does not arise from a reduction in the available photopigment molecules. However, there is reduction in pigment concentration with age. Light adaptation cannot account for the decay of the trp response during illumination: in normal Drosophila a dim background light shortens the latency and rise time of the response and also shifts the intensity response function (V-log I curve) to higher levels of light intensity with relatively little reduction in the maximal amplitude (Vmax) of response. In the trp mutant, a dim background light or short, strong adapting light paradoxically lengthens the latency and rise time of the response and substantially reduces Vmax without a pronounced shift of the V-log I curve along the I axis. The effect of adapting light on the latency and V-log I curve seen in trp are associated with a reduction in effective stimulus intensity (reduction in excitation efficiency) rather than with light adaptation. Removing extracellular Ca+2 reduces light adaptation in normal Drosophila, as evidenced by the appearance of "square" responses to strong illumination. In the trp mutant, removing extracellular Ca+2 does not prevent the decay of the response during illumination.     

鳜鱼视觉特性及其对捕食习性适应的研究Ⅰ．视网膜电图光谱敏感性和适应特性

采用电生理方法研究了夜行性凶猛鱼类鳜鱼视网膜电图的一般特性、光谱敏感性和适应特性。顷鱼的视网膜电图不显示典型的混合型视网膜特征。明视和暗视视网膜电图的光谱敏感曲线形状基本相同,峰值都在５３０ｎｍ处,没有出现Ｐｕｒｋｉｎｊｅ氏位移。明适应曲线仅出现下降型变化,暗适应过程异常缓慢。鳜鱼的视网膜仅存在单一的光感受系统,即暗视系统,不可能形成色觉。但级鱼视网膜具有很高的光敏感性,适于弱光视觉。