不同运动图象同时刺激左右眼时的交替视动震颤(OKN)现象

采用了同时对左右眼分别以不同的运动图象刺激的实验方法,来测量及分析其OKN眼动反应,探索在OKN反映中两眼之间的输入关系以及眼动控制机制。实验结果发现在两眼的刺激图象不一致时,眼动反应为交替的OKN反应,即中枢神经系统根据各眼的刺激速度,交替地控制产生OKN眼动反应。本文还从闭环控制上讨论了视网膜上速度误差信号的作用。     

不同运动图象同时刺激左右眼时的交替视动震颤(OKN)现象

采用了同时对左右眼分别以不同的运动图象刺激的实验方法,来测量及分析其OKN眼动反应,探索在OKN反映中两眼之间的输入关系以及眼动控制机制。实验结果发现在两眼的刺激图象不一致时,眼动反应为交替的OKN反应,即中枢神经系统根据各眼的刺激速度,交替地控制产生OKN眼动反应。本文还从闭环控制上讨论了视网膜上速度误差信号的作用。     

Behavioral genetic approaches to visual system development and function in zebrafish

The zebrafish is a recent vertebrate model system that shows great potential for a genetic analysis of behavior. Early development is extraordinarily rapid, so that larvae already display a range of behaviors 5 days after fertilization. In particular the visual system develops precociously, supporting a number of visually mediated behaviors in the larva. This provides the opportunity to use these visually mediated behaviors to screen chemically mutagenized strains for defects in vision. Larval optokinetic and optomotor responses have already been successfully employed to screen for mutant strains with defects in the visual system. In the adult zebrafish a visually mediated escape response has proved useful for screening for dominant mutations of the visual system. Here, I summarize visually mediated behaviors of both larval and adult zebrafish and their applicability for genetic screens, and present, the approaches and results of visual behavior carried out to date.     

Stabilization of the gaze in chameleons: visual and vestibular reflexes

Some visual, vestibular and proprioceptive reflexes which contribute to gaze (head + eye) stabilization were quantified in the chameleon. All the reflexes were analysed in the horizontal plane, and the visual reflexes were also studied in the vertical plane. In restrained-head animals, both the optokinetic nystagmus (OKN) and the vestibulo-ocular reflex (VOR) had low gains. In free-head animals, the head (opto-collic or vestibulo-collic reflex) and eye (OKN or VOR) responses added their effects, thus improving gaze stabilization, especially during vestibular stimulation. Cervical stimulation provoked both a cervico-ocular reflex (COR) in the compensatory direction and a large number of saccades. The saccadic response was especially marked in the presence of patterned visual surroundings.     

Optokinetic horizontal head nystagmus in the frog after regeneration of transected retinal fibres

The recovery of optokinetic responses during regeneration of transected retinal fibres was studied. Regenerated optic fibres were demonstrated by the cobaltfilling technique. After transection of the chiasma in the midline in most of the cases regenerating retinal fibres did not cross, but terminated in ipsilateral visual centres. Aberrant fibres were found in the telencephalon, the periventricular region of the diencephalon, anterior-, posterior- and ventral tegmental commissures and in the isthmic nucleus. In one group of animals optokinetic responses did not return after regeneration of retinal fibres. Re-innervation was either symmetrical on both sides from one eye, or terminal fibres were disorganized. In the second group of animals normal optokinetic nystagmus (OKN) returned. In these cases the fibres were symmetrically distributed on both sides or, alternatively, the crossed fibres outnumbered the ipsilateral ones. The third group of animals showed reversed OKN, spontaneous turning of the head, circling or head nystagmus. Here the majority of regenerated fibres termined ipsilaterally. When the OKN returned, the retinal projection was always restored in the pretectal region. Contrary to our earlier work, we concluded that the basal optic nucleus is not essential for horizontal optokinetic head nystagmus because in a few animals, retinal fibres did not invated this nucleus, and in spite of this the OKN could be evoked. We think now that the most important structure (as an input channel) for horizontal OKN of frogs is the pretectal region.     

两种等亮度不同运动图形刺激下的交替OKN眼动

等亮度彩色运动条纹产生视动震颤（ＯＫＮ）眼动反应揭示了ＯＫＮ眼动控制系统中存在颜色通道;而在两个相反方向运动的等亮度彩色运动条纹图象同时显示刺激时能引起交替的两个方向的ＯＫＮ跟踪眼动。故表明了在ＯＫＮ眼动控制系统中各个颜色通道交替地控制产生ＯＫＮ眼动反应。     

运动图形刺激时家兔的视动震颤反应

旨在用实验方法研究家兔的视动震颤（ＯＫＮ）眼动特点以及单侧前庭迷路损伤对ＯＫＮ的影响,结果表明：单眼刺激时,家兔的ＯＫＮ反应存在着从颞侧到鼻侧方向的方向优势;恒定速度刺激时,刺激开始后,家兔的ＯＫＮ眼动跟踪速度具有从小到大最后趋于稳态的建立过程,刺激消失后,存在眼动速度由大到小直到消失的视动后震颤（ＯＫＡＮ）反应,这两个过程反应了ＯＫＮ系统中可能存在速度存储机制及其对ＯＫＮ眼动的控制作用;单侧前庭     

Forward genetic analysis of visual behavior in zebrafish

The visual system converts the distribution and wavelengths of photons entering the eye into patterns of neuronal activity, which then drive motor and endocrine behavioral responses. The gene products important for visual processing by a living and behaving vertebrate animal have not been identified in an unbiased fashion. Likewise, the genes that affect development of the nervous system to shape visual function later in life are largely unknown. Here we have set out to close this gap in our understanding by using a forward genetic approach in zebrafish. Moving stimuli evoke two innate reflexes in zebrafish larvae, the optomotor and the optokinetic response, providing two rapid and quantitative tests to assess visual function in wild-type (WT) and mutant animals. These behavioral assays were used in a high-throughput screen, encompassing over half a million fish. In almost 2,000 F2 families mutagenized with ethylnitrosourea, we discovered 53 recessive mutations in 41 genes. These new mutations have generated a broad spectrum of phenotypes, which vary in specificity and severity, but can be placed into only a handful of classes. Developmental phenotypes include complete absence or abnormal morphogenesis of photoreceptors, and deficits in ganglion cell differentiation or axon targeting. Other mutations evidently leave neuronal circuits intact, but disrupt phototransduction, light adaptation, or behavior-specific responses. Almost all of the mutants are morphologically indistinguishable from WT, and many survive to adulthood. Genetic linkage mapping and initial molecular analyses show that our approach was effective in identifying genes with functions specific to the visual system. This collection of zebrafish behavioral mutants provides a novel resource for the study of normal vision and its genetic disorders.     

The interstitial nucleus of Cajal of the cat. II. Effects of kainic acid lesion on vertical optokinetic nystagmus and after-nystagmus

The effects of bilateral lesions of the interstitial nucleus of Cajal (INC) by ka?nic acid on vertical optokinetic nystagmus (OKN) and after-nystagmus (OKAN) were studied in four cats: in three cats, in the acute stage from 1 to 60 days after the lesions; in the fourth cat, they were studied 3 years after the lesions were made. Histological control of lesions showed that the whole INC was bilaterally destroyed in two cats of the acute group and only the upper part of INC in the third cat. In the chronic cat, the density of cell bodies in both INC was lower than normal. In the acute group, the cats exhibited a spontaneous downward eye drift in light and in darkness. During an upward optokinetic stimulation, the effect of INC lesions was dramatic: upward slow phases and downward quick phases of OKN were abolished. Sixty days post lesions, small upward slow eye movements were again observed. During a downward optokinetic stimulation, the defect was much less; in particular, after a slight impairment of downward slow phases, during the first days post lesions, they recovered quickly. The secondary optokinetic after nystagmus (OKAN II) ensuing a downward OKN was cancelled and did not reappear 60 days post lesions. In the chronic stage, three years after the lesions, during an upward optokinetic stimulation, the cat showed upward slow phases with velocities close to normal. However, upward slow phases were curved: the velocity at the end of the slow phases was lower than at the beginning. After an upward OKN (the direction of slow phases gives the direction of the OKN and OKAN), the ensuing OKAN was present but abnormal.     

家兔的开环及闭环视动震颤（OKN）研究

研究了家兔闭环及开环状态的ＯＫＮ反应。闭环时,ＯＫＮ眼动速度与刺激速度成正比,增益接近１。当固定受刺激眼形成开环状态时,ＯＫＮ眼动速度远大于刺激速度,增益可达１０２左右,说明ＯＫＮ系统是由运动跟踪的负反馈机制控制的;刺激范围缩小则眼动反应减弱,表明ＯＫＮ系统对运动信息具有空间总和作用;以视野中不同高度的局部刺激时,发现视网膜中央视带比周边部对ＯＫＮ刺激的敏感性高;单光点刺激视带中央可诱发出清楚的ＯＫＮ反应,这为临床上应用ＯＫＮ客观测定视网膜功能和运动感知提供了可能性。     

正常及异常双眼视觉的视动震颤（OKN）反应特性研究

为探讨不同双眼视觉状态下ＯＫＮ反应的特性,对正常人和不同类型的双眼视觉异常者的单眼鼻向及颞向ＯＫＮ反应进行了研究。实验发现：单眼视觉抑制者表现出鼻向与颞向ＯＫＮ反应不对称特性;两眼皆因视剥夺造成双眼视觉异常者主要以ＯＫＮ眼动增益降低为特点;双眼视觉正常者鼻、颞向ＯＫＮ反应是对称的。结果表明：单眼ＯＫＮ眼动反应的不对称特性及增益改变对探讨双眼视觉异常机制有重要意义,为视皮层双眼细胞异常导致单眼ＯＫＮ不对称的假设提供了支持性证据,并对弱视早期诊断及其分类有重要价值。     

Measuring the optokinetic response of zebrafish larvae

Our laboratory screens for visual mutants by examining larval eye movements in response to rotating illuminated stripes. This behavior, which is termed an optokinetic response (OKR), is a reflex that appears in zebrafish at the same time as the development of the visual system. The OKR can be accurately measured by 4 d post-fertilization, which is the age when larvae begin foraging for food. The OKR requires approximately 1 min per larva analyzed. After identifying fish with defective eye movements, we conduct secondary screens (such as histological analysis and electroretinography) to identify the subset of fish with disruptions in the function of the outer retina. This paper describes our protocol for the OKR. Our setup is simple to construct and the materials needed are inexpensive. This makes our system especially useful for new undergraduate and graduate students, as well as introductory science lecturers.     

经直接与间接通路的视动震颤(OKN)眼动反应

采用鼻向及颞光栅图形刺激猫的半侧视网膜,以探讨鼻侧半神网膜至视束核的视动震颤直接通路及颞侧半视网膜经皮层至ＮＯＴ的间接通路对ＯＫＮ眼动反应的神经控制特性。实验结果显示：高速度刺激的闭环ＯＫＮ反应及开环ＯＫＮ反应均对鼻向运动刺激有明显的方向选择性;半侧视网膜刺激时,鼻侧半视网膜的鼻向ＯＫＮ增益明显高于颞侧半视网膜的鼻向增益,说明猫的ＫＯＮ眼动反应以跟踪鼻向运动为主,并且这种对鼻向运动刺激选择性主要来     

The Development of Vision in the Zebrafish (Danio rerio)

We studied the development and maturation of the visual system by determining when zebrafish begin to see and to move their eyes. This information was correlated with the time courses of the development of the retina, the retinofugal projection, the retinal image, and the extraocular muscles, to obtain an integrated picture of early visual development. Two visual behaviors were monitored over 48–96 hr postfertilization (hpf). The startle response (body twitch) was evoked by an abrupt decrease in light intensity. The optokinetic response (tracking eye movements) was evoked by rotation of a striped drum. Visually evoked startle developed over 68–79 hpf, more than 20 hr after the onset of a touch-evoked startle. It was not seen in eyeless fish, excluding a role for nonretinal light senses. Tracking eye movements developed over 73–80 hpf. They were always in the direction of drum rotation, even when the fish had been light deprived from blastula stage, ruling out a “trial and error” period of learning to track the drum. The image formed by the ocular lens was examined in intact fish made transparent by suppressing the formation of melanin. The eye was initially far sighted and gradually improved, so that by 72 hpf the image plane coincided with the photoreceptor layer. The extraocular muscles assumed their adult configuration between 66 and 72 hpf. Thus, the retinal image and functional extraocular muscles appeared nearly simultaneously with the onset of tracking eye movements and probably represent the last events in the construction of this behavior.     

Tests of the mouse visual system

To apply the approach of forward genetics (e.g., gene identification with mutagenesis and screening, followed by positional cloning) to the mouse, it is necessary to have available screening tests that can be applied rapidly to individual mice and that give a reliable assessment of visual function. This paper reviews the strengths and limitations of two anatomical tests related to visual function, fundus examination and retinal histological examination. Two tests that do not depend on behavior of a conscious animal are reviewed: the electroretinogram and the visual evoked potentials of the cortex. Eight behavioral tests are also summarized: maze-based tests, cued fear conditioning, tests based on conditioned suppression, visual placing, optokinetic nystagmus, pupillary reflex, and light-induced shifts in circadian phase. It is recommended that retinal histology, the electroretinogram, and visual-evoked potentials be used at the present time for screening because they assess the function and structure of the visual system rapidly and reliably. In fact, the electroretinogram (or visually evoked potentials) can be recorded from several animals simultaneously in response to the same stimulus. It is also recommended that efforts be made to develop more appropriate, automated, behavioral tests of visual perception than are now available, particularly tests that rely solely on rewarding visually evoked behavior. Two other promising behavioral tests are cued fear conditioning and variants of maze tests. Received: 17 December 1999 / Accepted: 17 December 1999     

Effects of optokinetic stimulation on motor asymmetry in the goldfish

In goldfish fries, we examined the effect of the optomotor reaction (drive to swim toward moving images of vertical dark bars) on the behavioral motor asymmetry. Contralateral optokinetic stimulation of fishes (rotation of the bars against the direction preferred by fishes in their turnings) gradually smoothed and, later on, inverted the motor asymmetry, while the asymmetry underwent no modifications in the case of ipsilateral optokinetic stimulation (rotation of the bars in the direction similar to that preferred for turnings). Contralateral optokinetic stimulation also induced long-lasting inversion of the motor asymmetry of immobilized fishes deprived of the possibility to follow the movement of bar images. Ipsilateral optokinetic stimulation of fishes with the enucleation of the ipsilateral eye enhanced their motor asymmetry, while contralateral stimulation either did not modify the motor asymmetry of such individuals or inverted this feature. These data agree with the concept that, in fishes, one eye dominates and more actively provides tracking of the movement of bars, while another eye is a subdominant one. In general, we first found that the use of specific visual stimulation allows one to modify for a long time the behavioral motor asymmetry of the fishes, which, as is known, correlates with the morphofunctional asymmetry of Mauthner neurons (MNs). Visual information that activates MNs influences mostly the ventral dendrites of these neurons; thus, our findings allow us to believe that stimulations, which initiate the optomotor reaction, can serve as an adequate physiological model of natural visual stimulation of MNs (with projection of the respective influences on the ventral dendrites of the above cells). The use of such an experimental paradigm opens up new possibilities for studies of the role of these dendrites in the functions of MNs and of the plasticity of morphofunctional organization of these cells. Neirofiziologiya/Neurophysiology, Vol. 39, No. 2, pp. 133–145, March–April, 2007.     

Visual cortex contribution to the organization of eye movements produced by local electrical stimulation of the cat lateral geniculate body

Eye movements evoked by local electrical stimulation of the dorsal nucleus of the lateral geniculate body were analyzed after removal of the visual cortex and in intact animals during trials on awake cats. No significant difference was observed between the eye movement patterns of the two animal groups evoked by electrical stimulation. These movements could be classed into three main groups: those unassociated with the starting position of the eyes in orbit (or unidirectional movements), goal-directed, and centered movements, with direction depending on the initial position of the eyes in their orbits. Our findings indicate that the cortical visual areas are neither the principal nor an indispensable link in the chain for transmitting signals evoked by (electrically) stimulating the geniculate body from the cortical structures of the direct visual pathway towards the operative links of the oculomotor system. Potential pathways for conducting information from the dorsal nucleus of the lateral geniculate body to oculomotor system structures are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 164–170, March–April, 1987.     

Interocular motion combination for dichoptic moving stimuli

When gratings moving in different directions are presented separately to the two eyes, we typically perceive periods of the combination of motion in the two eyes as well as periods of one or the other monocular motions. To investigate whether such interocular motion combination is determined by the intersection-of-constraints (IOC) or vector average mechanism, we recorded both optokinetic nystagmus eye movements (OKN) and perception during dichoptic presentation of moving gratings and random-dot patterns with various differences of interocular motion direction. For moving gratings, OKN alternately tracks not only the direction of the two monocular motions but also the direction of their combined motion. The OKN in the combined motion direction is highly correlated with the perceived direction of combined motion; its velocity complies with the IOC rule rather than the vector average of the dichoptic motion stimuli. For moving random-dot patterns, both OKN and perceived motion alternate only between the directions of the two monocular motions. These results suggest that interocular motion combination in dichoptic gratings is determined by the IOC and depends on their form.