Ocular Compensation Research Articles

Effects of varying illumination on ocular aberrations and aberration compensation before and after small incision Lenticule extraction: a prospective cohort study

BackgroundThere are few reports regarding the influence of varying illumination on the compensation effect before and after corneal refractive surgery. We aimed to evaluate the changes in refraction, higher-order aberrations, and aberration compensation between mesopic and photopic illumination before and after small incision lenticule extraction.MethodsIn this prospective cohort study, only the right eyes of patients who underwent small incision lenticule extraction for the correction of myopia and myopic astigmatism at the Tianjin Eye Hospital were included. Wavefront refraction and higher-order aberrations were measured preoperatively and 3 months postoperatively under mesopic and photopic illumination. Compensation factors were calculated as 1 − (aberration of the whole eye/aberration of the anterior corneal surface).ResultsForty patients undergoing small incision lenticule extraction were enrolled. All surgeries were completed without postoperative complications. Preoperatively, the eyes only had a statistically significantly higher (t = − 4.589, p < .001) spherical refractive error under mesopic vs. photopic illumination (median [interquartile range], − 6.146 [2.356] vs. − 6.030 [2.619] diopters [D]), whereas postoperatively, the eyes also exhibited statistically significantly higher (t = − 3.013, p = .005) astigmatism (− 0.608 [0.414] vs. − 0.382 [0.319] D). Differences in spherical refraction between the two illuminations were the highest in postoperative eyes (Δ > 0.5 D). Only postoperative eyes exhibited statistically significant elevations (t ≥ 4.081, p < .001) in higher-order aberrations under mesopic illumination, and only preoperative eyes exhibited statistically significantly enhanced (χ2 = 6.373, p = .01 for fourth-order and χ2 = 11.850, p = .001 for primary spherical aberrations) and decreased (χ2 = 13.653, p = .001 for horizontal trefoil) compensation factors under mesopic illumination.ConclusionsExaggerations in higher-order aberrations and myopic shift after small incision lenticule extraction became apparent under mesopic illumination. Slight undercorrection may have an enhanced effect under low illumination and may reduce night vision. The specific changes in compensation effects in preoperative eyes may improve optical quality under mesopic illumination. Postoperative eyes have reduced compensation ability, specifically for spherical aberrations, under mesopic illumination, which may diminish night vision. Further studies that include the measurement of subjective night vision parameters should be conducted.

Evidence of top-down modulation of the Brentano illusion but not of the glare effect by transcranial direct current stimulation.

Transcranial direct current stimulation (tDCS) has been widely used for modulating sensory, motor and cognitive functions, but there are only few attempts to induce and change illusory perception. Visual illusions have been the most traditional and effective way to investigate visual processing through the comparison between physical reality and subjective reports. Here we used tDCS to modulate two different visual illusions, namely the Brentano illusion and the glare effect, with the aim of uncovering the influence of top-down mechanisms on bottom-up visual perception in two experiments. In Experiment 1, to a first group of subjects, real and sham cathodal tDCS (2mA, 10min) were applied over the left and right posterior parietal cortices (PPC). In Experiment 2, real and sham cathodal tDCS were applied to the left and right occipital cortices (OC) to a second group of participants. Results showed that tDCS was effective in modulating only the Brentano illusion, but not the glare effect. tDCS increased the Brentano illusion but specifically for the stimulated cortical area (right PPC), illusion direction (leftward), visual hemispace (left), and illusion length (160mm). These findings suggest the existence of an inhibitory modulation of top-down mechanisms on bottom-up visual processing specifically for the Brentano illusion, but not for the glare effect. The lack of effect of occipital tDCS should consider the possible role of ocular compensation or of the unstimulated hemisphere, which deserves further investigations.

The Interaction of Pre-programmed Eye Movements With the Vestibulo-Ocular Reflex.

The Vestibulo-Ocular Reflex (VOR) works to stabilize gaze during unexpected head movements. However, even subjects who lack a VOR (e.g., vestibulopathic patients) can achieve gaze stability during planned head movements by using pre-programmed eye movements (PPEM). The extent to which PPEM are used by healthy intact subjects and how they interact with the VOR is still unclear. We propose a model of gaze stabilization which makes several claims: (1) the VOR provides ocular stability during unexpected (i.e., passive) head movements; (2) PPEM are used by both healthy and vestibulopathic subjects during planned (i.e., active) head movements; and (3) when a passive perturbation interrupts an active head movement in intact animals (i.e., combined passive and active head movement) the VOR works with PPEM to provide compensation. First, we show how our model can reconcile some seemingly conflicting findings in earlier literature. We then test the above-mentioned predictions against data we collected from both healthy and vestibular-lesioned guinea pigs. We found that (1) vestibular-lesioned animals showed a dramatic decrease in compensatory eye movements during passive head movements, (2) both populations showed improved ocular compensation during active vs. passive head movements, and (3) during combined active and passive head movements, eye movements compensated for both the active and passive component of head velocity. These results support our hypothesis that while the VOR provides compensation during passive head movements, PPEM are used by both intact and lesioned subjects during active movements and further, that PPEM work together with the VOR to achieve gaze stability.

TRAVEL TO HIGH MOUNTAIN ELEVATIONS FOLLOWING VITRECTOMY WITH INTRAOCULAR GAS

To evaluate the effects and safety of travel by land through high mountainous elevations after surgery in patients who have undergone pars plana vitrectomy with intraocular gas. A retrospective cohort study of 75 patients post pars plana vitrectomy with intraocular gas who traveled by land through mountain elevations of up to 3895 feet above sea level within 1 day of surgery. The average rate of ascent through the mountains was 29 ft/min and the maximum theoretical ocular compensation was 0.57 cubic centimeters (cc), which occurred at 4259 feet above sea level. A statistically significant change in postoperative intraocular pressure was found when compared with preoperative (P = 0.010), with two factors influencing this change: the type of gas (C3F8, P = 0.038) and lens status (pseudophakic, P = 0.010), with a mean final intraocular pressure still within the safe range. There were no cases of retinal vascular occlusion, acute elevations of intraocular pressure requiring surgical intervention, or symptomatic visual field loss attributable to elevated intraocular pressure. It seems that patients with a complete fill of intraocular gas after pars plana vitrectomy may travel safely by land through mountains with a peak ascent of 3895 feet, final ascent of 2787 feet, and a mean rate of 29 ft/min. These findings can significantly impact patient costs and convenience.

High Resolution Retinal Imaging Based on Ocular Aberration Measurement and Compensation

Ocular retinal imaging is a major diagnostic modality for retinal disease, and can play a critical role for diagnosing systemic diseases such as diabetes and eye-specific diseases such as macular degeneration and diabetic retinopathy, the leading causes of blindness. In order to get high-resolution retinal imaging and develop the low-cost and compact retinal imaging system, we employ micro adaptive optics, which is consisted of wavefront sensor, wavefront corrector and control system. In this paper, the theory, design and testing of the ocular retinal microscopy is detailed, with an emphasis on the eye wavefront aberration describing, aberration detecting method with Hartmann-Shack wavefront sensing and close-loop aberration compensating by micromachined membrane deformable mirrors(MMDM).The ocular retinal microscopy experimental setup is built, the retinal cell imaging had been snapped. It is showed in this work that the ocular retinal microscopy based on adaptive optics system can enable diffraction-limited imaging of micro-scale features of the retina, through real-time compensation of aberrations introduced by the eye.

Effects of ocular rotation on parapapillary retinal nerve fiber layer thickness analysis measured with spectral-domain optical coherence tomography

To evaluate the effects of ocular rotation on parapapillary retinal nerve fiber layer (RNFL) thickness measured by spectral-domain optical coherence tomography (SD-OCT). Eighty-eight normal and 205 glaucomatous eyes were studied. RNFL thickness was measured by 3D OCT. Ocular rotation angle was measured from a fundus image obtained by a non-mydriatic fundus camera equipped with 3D OCT. The average, hemi-superior, and hemi-inferior RNFL thicknesses as well as those in the 4 quadrants and the 16-segmented superotemporal (ST-1) and inferotemporal (IT-4) sectors were compared both before and after correcting for ocular rotation. Receiver operating characteristic curves and the areas under the curve (AUC) for the RNFL thicknesses were calculated on the basis of the data from glaucomatous and control eyes. The relationships between RNFL thickness and retinal sensitivity in the corresponding visual field were analyzed using a Humphrey field analyzer. Correction for ocular rotation did not affect the AUCs of the hemi-superior and hemi-inferior RNFL thicknesses. RNFL thicknesses in all of the quadrants and in ST-1 and IT-4 were significantly changed by correcting for ocular rotation. The correlations between all RNFL sectors and retinal sensitivity were not changed by correcting for the ocular rotation angle. Ocular rotation compensation affected RNFL thickness measurement with 3D OCT. However, the effect was clinically negligible in the diagnosis of glaucoma.

Human eye-head gaze shifts preserve their accuracy and spatiotemporal trajectory profiles despite long-duration torque perturbations that assist or oppose head motion

Humans routinely use coordinated eye-head gaze saccades to rapidly and accurately redirect the line of sight (Land MF. Vis Neurosci 26: 51-62, 2009). With a fixed body, the gaze control system combines visual, vestibular, and neck proprioceptive sensory information and coordinates two moving platforms, the eyes and head. Classic engineering tools have investigated the structure of motor systems by testing their ability to compensate for perturbations. When a reaching movement of the hand is subjected to an unexpected force field of random direction and strength, the trajectory is deviated and its final position is inaccurate. Here, we found that the gaze control system behaves differently. We perturbed horizontal gaze shifts with long-duration torques applied to the head that unpredictably either assisted or opposed head motion and very significantly altered the intended head trajectory. We found, as others have with brief head perturbations, that gaze accuracy was preserved. Unexpectedly, we found also that the eye compensated well--with saccadic and rollback movements--for long-duration head perturbations such that resulting gaze trajectories remained close to that when the head was not perturbed. However, the ocular compensation was best when torques assisted, compared with opposed, head motion. If the vestibuloocular reflex (VOR) is suppressed during gaze shifts, as currently thought, what caused invariant gaze trajectories and accuracy, early eye-direction reversals, and asymmetric compensations? We propose three mechanisms: a gaze feedback loop that generates a gaze-position error signal; a vestibular-to-oculomotor signal that dissociates self-generated from passively imposed head motion; and a saturation element that limits orbital eye excursion.

Temporal constraints on experimental emmetropization in infant monkeys.

To characterize the temporal integration properties of the emmetropization process, the authors investigated the effects of brief daily interruptions of lens wear on the ocular compensation for negative lenses in infant rhesus monkeys. Eighteen monkeys wore -3 D lenses binocularly starting from approximately 3 weeks of age. Six of these monkeys wore the lenses continuously. For the other animals, the -3 D lenses were removed for four 15-minute periods each day. During these periods, the monkeys viewed through either zero-power lenses (n = 6) or +4.5 D lenses (n = 6). Three monkeys reared with binocular plano lenses and 16 monkeys reared normally served as controls. Refractive development was assessed by cycloplegic retinoscopy and A-scan ultrasonography. As expected, the group of animals that wore the -3 D lenses continuously exhibited clear evidence of compensating axial myopia. These predictable myopic changes were mostly eliminated by the brief, daily periods of viewing through plano lenses. Interestingly, brief periods of viewing through +4.5 D lenses produced weaker protective effects. Brief periods of unrestricted vision can prevent the axial myopia normally produced by long daily periods of imposed hyperopic defocus. Thus, the temporal integration properties of the emmetropization process normally reduce the likelihood that transient periods of hyperopic defocus will cause myopia.

Ocular compensation for alternating myopic and hyperopic defocus

During development, the eye grows under visual feedback control, as shown by its compensating for defocus imposed by spectacle lenses. Under normal conditions the sign and magnitude of defocus vary with viewing distance, accommodative status and other factors. To explore how periods of myopic and hyperopic defocus are integrated over time we presented rapidly alternating episodes of myopic and hyperopic defocus by sequentially illuminating a nearby scrim and the wall beyond it to chick eyes wearing lenses that put the far point between the two surfaces. We found that equal periods of myopic and hyperopic defocus generally led to compensatory hyperopia, showing that myopic defocus had a disproportionate effect. Furthermore, the degree of hyperopia depended on the frequency of alternation: low frequencies (1 cycle/30 min) resulted in more hyperopia, whereas at high frequencies (1 cycle/s) the myopic and hyperopic defocus nearly cancelled each other. If similar temporal integration effects apply to humans, they may help explain why brief accommodation events may not influence lens-compensation and why a child’s total reading time may be a poor predictor of myopic progression.

Temporal constraints on lens compensation in chicks

If the effective focal length of a growing eye is modified by spectacle lenses, the eye compensates by altering its growth, thereby keeping images in focus, a process we presume is similar to normal emmetropization. Using chicks, we have investigated how much visual exposure the eye needs to exhibit the two principal components of ocular compensation: altered rate of elongation (a scleral mechanism) and altered choroidal thickness. We have found that surprisingly small amounts of vision through spectacle lenses can elicit robust scleral and choroidal compensation if other visual feedback is limited by keeping the animals in the dark when not wearing lenses. Furthermore, we have found that the amount of vision necessary to induce these responses can be summarized as three rules: First, several brief daily episodes are more effective than a single or a few longer daily episodes, even if the total amount of vision is the same. Second, extremely brief episodes, even if very frequent, are relatively ineffective. Third, when plus and minus lenses are worn successively on the same eye, the plus lens has the dominant effect, even if the minus lens is worn five times longer than the plus lens. In addition, we have shown that the elongation rate and choroidal thickness responses are dissociable, such that brief, infrequent lens-viewing produces only an elongation response in the case of plus lens-wear and only a choroid response in the case of minus lens-wear. We thus show that the emmetropization system does not integrate defocus in a simple, linear fashion. These non-linearities, if present in children, might explain why, although education and reading show an epidemiological correlation with myopia, the total time spent reading and doing other nearwork by individual children generally does not predict the degree of myopia. It may therefore be necessary to quantify more complex temporal patterns of nearwork over the day in order to measure the impact of nearwork on eye growth.

Ocular compensation due to labyrinthine input during natural motion.

With recent technical developments, it has become possible to study eye and head movements in all angular and linear degrees of freedom during natural activities such as standing and treadmill ambulation. Such studies have revealed that normal human gaze stabilization during ambulation is accomplished not only by ocular rotation in response to head rotation, but also by head translation coordinated with head rotation as appropriate to viewing distance. Typically, head translation in a given direction is coupled via an active mechanism with antiphase rotation in the opposite direction, accomplishing some gaze stabilization for near targets and supplementing the vestibulo-ocular reflex (VOR). Subunity angular VOR gain (eye velocity/head velocity) not only is observed during most natural activities, but also minimizes the disturbance to retinal image stability so that excellent visual acuity can be obtained. The generation of eye movements compensating for head motion during ambulation can be quantitatively modeled as an additive combination of semicircular canal responses, with otolith responses scaling by inverse viewing distance, so angular VOR gain correlates poorly with gaze stability. Compensatory eye movements during self-generated head rotations appear driven to a significant extent by nonvestibular mechanisms since they cannot be modeled as an additive combination of otolith and canal responses. People with unilateral vestibular deafferentation have normal gaze stability during natural activities such as standing, walking, and running, although gaze stability is impaired in bilateral deafferentation. Limitation of head motion during natural activities is a common strategy in those with reduced peripheral vestibular sensitivity.

A population-based survey of hospitalized work-related ocular injury: diagnoses, cause of injury, resource utilization, and hospitalization outcome

Occupational injury is a major source of ocular trauma and is often preventable. A statewide population-based survey of severe work-related ocular injury was generated by using the California Hospital discharge database to identify hospitalized ocular injury and workers compensation as principal payor to identify work-relatedness. Information concerning diagnoses, procedures, causes of injury, length of hospital stay, total hospital charges and disposition at hospital discharge were obtained for injuries occurring during the calendar year 1988. A total of 455 admissions for work-related ocular trauma were identified. The most common work-related ocular trauma diagnoses associated with hospitalizations were open globe injury (46%), adnexal wounds (20%), orbital fractures (11%), and traumatic hyphema (11%). The most common causes of work-related ocular trauma were foreign-body or projectile objects (19%), transport vehicles (18%), cutting or piercing objects (17%), and assaults (9%). Approximately 8% reported other than routine disposition at time of hospital discharge, including long-term nursing or rehabilitation services and death. Mean hospital stay when ocular trauma was the principal admitting diagnosis was 3.7 days. Results differed significantly for admissions reporting ocular trauma as the principal admitting diagnosis compared to admissions that did not. Hospitalized work-related ocular trauma is represented by a wide spectrum of injuries with substantial morbidity and economic costs. Projected to the United States population, these data indicate annual hospital charges excluding professional fees of $14.6 million when work-related ocular trauma is the principal admitting diagnosis and $40 million for admissions where ocular trauma is either a principal or secondary diagnosis.

Effects on the compensatory responses to positive and negative lenses of intermittent lens wear and ciliary nerve section in chicks

This study examined the ocular compensation to lens-induced defocus in chick and the effect of interrupting lens wear on a daily basis. Eyes fitted with +10 D lenses at hatching compensated rapidly, with almost complete compensation after 4 days of lens wear; they had decreased vitreous chamber depth compared to normal eyes and were thus hyperopic when the lenses were removed. In contrast, adaptation to the −10 D lenses was much slower, was still incomplete after 9 days of lens wear, and in this case, eyes had increased vitreous chamber depth and were myopic without the lenses. Adaptation improved when lens wear was delayed until 7 days after hatching. The effect of interrupting lens wear by periods of normal vision varied with the sign of the lenses worn. Hyperopia was always seen in response to +10 D lenses, although the magnitude of the response decreased as the duration of lens wear was decreased. In contrast, even brief periods of normal vision, i.e., 3 hr, prevented the development of myopia in response to the −10 D lenses; this apparent sensitivity to normal vision is similar to that reported for form-deprivation myopia. Ciliary nerve section used here to eliminate accommodation did not alter these response patterns.

Ocular compensation for self-motion. Visual mechanisms.

In monkeys, there are several reflexes that generate eye movements to compensate for the observer's own movements. Two vestibuloocular reflexes compensate selectively for rotational (RVOR) and translational (TVOR) disturbances of the head, receiving their inputs from the semicircular canals and otolith organs, respectively. Two independent visual tracking systems that deal with residual disturbances of gaze are manifest in the two components of the optokinetic response: the indirect or delayed component (OKNd) and the direct or early component (OKNe). We hypothesize that OKNd--like the RVOR--is phylogenetically old, being found in all animals with mobile eyes, and that it evolved as a backup to the RVOR to compensate for rotational disturbances of gaze. Indeed, optically induced changes in the gain of the RVOR result in parallel changes in the gain of OKNd, consistent with the idea of shared pathways as well as shared functions. In contrast, OKNe--like the TVOR--seems to have evolved much more recently in frontal-eyed animals and, we suggest, acts as a backup to the TVOR to deal primarily with translational disturbances of gaze. Frontal-eyed animals with good binocular vision must be able to keep both eyes directed at the object of regard irrespective of proximity, and in order to achieve this during translational disturbances, the output of the TVOR is modulated inversely with the viewing distance. OKNe shares this sensitivity to absolute depth, consistent with the idea that it is synergistic with the TVOR and shares some of its central pathways. There is evidence that OKNe is also sensitive to relative depth cues such as motion parallax, which we suggest helps the system to segregate the object of regard from other elements in the scene. However, there are occasions when the global optic flow cannot be resolved into a single vector useful to the oculomotor system (e.g., when the moving observer looks towards the direction of heading). We suggest that on such occasions a third independent tracking mechanism, the smooth pursuit system, is deployed to stabilize gaze on the local feature of interest. In this scheme, the pursuit system has an attentional focusing mechanism that spatially filters the visual motion inputs driving the oculomotor system. The major distinguishing features of the 3 visual tracking mechanisms are summarized in Table 1.

Eye movement responses to linear head motion in the squirrel monkey. II. Visual-vestibular interactions and kinematic considerations

1. Horizontal, vertical, and torsional eye movements were recorded (search coil technique) from five squirrel monkeys during horizontal linear oscillations at 0.5, 1.5, and 5.0 Hz, 0.36 g peak acceleration. Monkeys were positioned to produce linear motion in their nasooccipital (NO), interaural (IA), and dorsoventral (DV) axes. Responses of the linear vestibuloocular reflex (LVOR) were recorded in darkness and in the light with the subjects viewing a head-fixed field 22 or 9.2 cm from the eye. The latter condition provided a measure of "visual suppression" of the LVOR (VSLVOR). Responses were also recorded while monkeys viewed earth-fixed targets, which allowed visual enhancement of the LVOR (VLVOR). Vergence angle was recorded in two monkeys to assess directly the point of binocular fixation in space during linear motion. 2. Two LVOR response types, vertical responses during 0.5-Hz NO-axis translation (NO-vertical) and torsional responses at all frequencies during IA-axis oscillation (IA-torsional) could not be compensatory reflexes for head translation because they either move the eye off target (NO-vertical) or tort the eye relative to the visual world (IA-torsional), thereby degrading visual image stability. 3. Other response types are considered compensatory because they help maintain ocular fixation in space during linear head translation. These include horizontal responses to IA-axis motion (IA-horizontal), vertical responses to DV-axis translation (DV-vertical), and both horizontal and vertical responses to NO-axis oscillation (1.5 and 5 Hz). Observations focus on responses to 5-Hz oscillations, in which visual inputs are essentially ineffective in modifying the LVOR. 4. The kinematics of perfect ocular compensation during head translation indicate that the ideal ocular response is governed by the motion of the eye relative to target position. Relevant variables include target distance, which is crucial for all axes of motion, and target eccentricity, which is important only for head motion roughly parallel to the target (NO-axis translation). Findings are compatible with predictions based on ideal kinematics. However, it is the point of binocular fixation in space, not actual target position, that governs LVOR behavior. 5. The IA-horizontal and DV-vertical LVOR is in response to head motion roughly orthogonal to the line of sight. Responses under all stimulus conditions (LVOR, VSLVOR, and VLVOR) behaved similarly at 5 Hz, and were modulated linearly with vergence [in meter angles (MA), the reciprocal of binocular fixation distance].(ABSTRACT TRUNCATED AT 400 WORDS)

Local ocular compensation for imposed local refractive error

Chicks were raised in a low-ceiling environment to find out if their eye growth could compensate for locally imposed hyperopic refractive errors. These chicks became selectively more myopic in the upper visual field than chicks raised in a high-ceiling environment. The vitreous chamber in the low-ceiling birds showed a selective elongation in the ventral region that was not seen in the eyes of the high-ceiling birds. This morphological difference was small, but probably adequate to account for the additional myopia in the low-ceiling birds. These results are consistent with the idea of a visually mediated growth mechanism regulating the local refractive state across the entire visual field so that it matches the customary viewing conditions. Such a mechanism might account for the finding of Fitzke, Hayes, Hodos, Holden and Low ( Journal of Physiology, London, 369, 33–44, 1985) that the refractive errors in the lower field are exactly appropriate for focusing the image of the ground on the retina.

Vestibular mechanisms related to vertical and torsional eye movements.

1. A series of experiments concerning the organization of the vertical vestibuloocular reflex (VOR) of the rabbit are reviewed. 2. Unanesthetized, encéphale isolé and intact, alert rabbits have been submitted to natural stimulation of the labyrinth consisting in sinusoidal oscillations and static tilts around the longitudinal axis. The oculomotor responses have been recorded at the level of the III nucleus oculomotor neurons (OMN) and from the four vertical eye muscles. 3. The phase and gain of the OMN response at different frequencies of oscillation indicate a serial involvement of the macular ampullar labyrinthine receptors in the activation of the OMN. In the low frequency range of head movements, macular receptors alone can provide adequate ocular compensation. By raising the frequency of head movements, the threshold value of acceleration for recruiting second order vestibular neurons related to semicircular canals is reached and the vertical VOR is controlled predominantly by the ampullar receptors of the vertical canals. 4. These results have been confirmed and extended in the alert, intact rabbit. By orienting the animal in different positions with respect to the axis of oscillation it has been possible to activate selectively one or the other pair of complementary vertical canals. The EMG responses recorded from the vertical eye muscle exhibited different magnitudes in the different orientations, while the phase curve remained the same. 5. The static otolith-ocular reflex has also been investigated by tilting the alert animals oriented in the same way. The pattern of the EMG responses elicited by the pure macular input coincides with that obtained during dynamic stimulation. The existence of separate functional macular regions specifically connected with the different eye muscles and a close convergent action of the two types of labyrinthine receptors in ocular compensation, are suggested.

Eye stabilization by statocyst mediated oculomotor reflex inNautilus

Nautilus pompilius can rotate its eye relative to its body so as to compensate for changes in its body orientation and maintain its eye fixed with respect to gravity. An ocular compensation reflex stabilizes its eye about the pitch axis against the rocking motions that occur as the animal swims by jet propulsion. The eye is not held absolutely still, as 1–2° spontaneous rotations occur even if the animal is clamped.

Central control of postural orientation in flatfish. I. Postural change dependence on central neural changes.

ABSTRACT Flatfish metamorphose from a larval form that swims upright like a standard fish to an adult that lies on one side, with both eyes on the upper side, having rotated posture 90 ° relative to gravity and the former normal posture. Adult Citharichthys stigmaeus and Hypsopsetta guttulata were used in behavioural and physiological experiments to determine whether the postural change is a peripheral or central phenomenon. Cleared and sectioned specimens verify that the otolith organs, unlike the eyes, do not rotate within the skull, and so do not maintain the normal vertebrate orientation with respect to gravity. Ocular compensation to lateral tilt shows that tactile cues, vision, and the semicircular canals are inadequate to produce tilt responses, but elimination of otolith function abolishes tilt responses. The major postural role of the otolith organs is not lost. Selective removal of otoliths demonstrates that the flatfish utriculus has only a minor role in tilt responses, and that the sacculus-lagena is required, unlike the situation in other vertebrates. The details of the ocular compensation responses are similar to those of standard fishes. Each sacculus lies at an angle of up to 45 ° when in the normal position, but unilateral loss does not change the phase of the response curve, indicating that the null response is set for a non-zero value of gravitational shear, unlike the null at zero shear to the utriculus in other vertebrates. Hysteresis effects suggest a differential sensitivity between tilts near the normal and the upside-down null positions. The narrowness of the effect argues against mechanical restrictions. Possibly the vertical utriculus is useful only near the normal, as an accessory organ, like the vertical lagena in other vertebrates. Neural units recorded from both eighth nerve and medulla show the expected activity properties of regular and irregular rate, tonic and phasic responses to tilt, directional dependence and ‘multi-valuedness’, as in other vertebrates. No novel response types are found, nor any distinctive ‘into-level’ types described for some vertebrates. Vibration sensitivity is associated with irregular rate, and exclusively vibration-sensitive units are apparent only in the utriculus. The shift in functional relations of the otolith organs relative to gravity is not apparently compensated for by any major change in peripheral afferent gravistatic unit properties. An increasing distinction between the null at normal and the null upside-down is shown by limited data on ocular compensation in three year-classes of flatfish. A central change in vestibular function is suggested that might be dependent on experience, as is gradual compensation to a vestibular lesion. Since peripheral changes are not responsible for the postural change, alternative central mechanisms are proposed, including central weighting of input, recognition of a complex input pattern, and plasticity of connexions, all of which have received some supporting evidence from these results.

Central control of postural orientation in flatfish. II. Optic-vestibular efferent modification of gravistatic input.

ABSTRACT Visual influences on postural responses mediated by the otolith organs were studied in behavioural and physiological experiments on two species of flatfish, the bothid Citharichthys stigmaeus and the pleuronectid Hypsopsetta guttulata. Ocular compensation to lateral tilt is altered toward a directional light stimulus ; the light effect is independent of the tilt effect, and sums with it. Maximum light effect is roughly one-fourth the magnitude of the maximum tilt response. Single shocks to the optic tectum evoke up to three responses in the cut stump of the ipsilateral vestibular nerve, falling into three latency groups at roughly 5, 10 and 20 msec; all three follow stimulation of the anterior lateral tectum. The two later groups suggest by their delay and extinction at low stimulus rates that their path involves at least one synapse. Some units determined as primary otolith-organ afferents by their response characteristics to lateral tilts show a change in rate following a burst of shocks to the tectum, but no response to single shocks. The usual effect is excitatory, with a latency of 50 –100 msec, and lasting 200 –500 msec after the stimulus. Some of these units also show tonic alteration in rate while a light is shining into one eye. The alteration is either an increase with light to either eye, or a directiondependent increase or decrease, with a latency of 50 –100 msec, and lasting ionically during stimulation times of up to 1 min. The effect is independent of tilt. While the proportion of affected units is small, the elimination of mechanical, muscular, autonomic or antidromic effects and the specificity of many of the responses demonstrate that optic-vestibular integration can involve an efferent control system from the tectum to the otolith organs. The presence of such visual interactions with vestibular input in adult flatfish leads to the hypothesis that the central control of the 90 ° postural change occurring during flatfish metamorphosis may involve crucial optic influences during the period of eye migration.