Task demands and binocular eye movements

Humans make rapid movements of their eyes several times a second that enable them to examine objects located at different positions in space with both of their eyes. Much of our understanding of these binocular movements comes from studies using experienced observers performing repetitive, unnatural tasks. But what eye movements are made when na?ve observers perform tasks demanding specific binocular visual information? We examined the binocular eye movements produced by observers performing two tasks differing in the visual information needed for their completion. Our motivation for doing this was to examine the role and function of binocular eye movements when making decisions. We considered the fixation strategies adopted by observers, the effects of the task on the dynamics of saccadic eye movements, and the combination of vergence and version in gaze shifts. We report that the task-dependent use of visual information can have a strong influence on the patterns of fixations, whilst not influencing saccade dynamics. Our data provide some support for the notion that observers choose and fixate a notional reference point in the scene when making judgments about depth structure.     

Latency of saccades, vergence, and combined movements in children with early onset convergent or divergent strabismus

The goal of this study was to examine latency of horizontal eye movements in the natural space (saccades, vergence, and combined saccade-vergence movements) in children with early onset convergent or divergent strabismus. Ten children were tested (8-11 years old): three with divergent strabismus, seven with convergent strabismus. A paradigm was used to elicit pure lateral saccades at far and near distance, pure vergence (convergence and divergence) and saccades combined with vergence movements. Horizontal eye movements from both eyes were recorded simultaneously by a photoelectric device (Oculometer, Dr. Bouis). The latency of saccades (at far and near distance), of vergence (convergence and divergence), and of combined movements greatly varies among subjects and has tendency to be longer than that observed in normal children of matched age, however, these differences reach significance in only a few cases. Children with divergent strabismus and residual gross binocular vision show abnormally longer vergence latencies than children with convergent strabismus without binocular vision. The initiation of combined movements does not show a dominant pattern, such as preceding vergence, as is found in normal children. Finally, strabismus surgery has no major effect on latencies. We conclude that there is no overall deficiency in latencies of eye movements in 3D space in children with early onset strabismus. Most likely, monocular visual input can be efficient as normal binocular vision for vergence movements. In a few subjects with divergent strabismus and fragile, intermittent binocular vision, latencies can be abnormally long, just because of the fragile binocular input and/or attention effort needs to use it. The absence of a pattern of initiation similar to normal children could be due to attention and fixation capabilities.     

Maldevelopment of convergence eye movements in macaque monkeys with small- and large-angle infantile esotropia

PURPOSE: To describe symmetric convergence eye movements evoked by disparity and/or accommodative cues in esotropic macaque monkeys, with the goal of determining whether these animals have the vergence deficits found in humans with esotropia. METHODS: Physical far and near targets were used to evoke large (approximately 8 degrees) symmetric convergence eye movements in four adult macaque monkeys (two with strabismus, two normal), using positive-feedback rewards. One strabismic monkey had infantile-onset, small-angle esotropia (small-eso approximately 2 degrees) induced by alternating occlusion from birth to age 9 months. The other strabismic monkey had naturally occurring, large-angle (approximately 25 degrees) infantile-onset esotropia (large-eso). Visual acuity was normal in each eye as measured by spatial sweep visually evoked potentials (VEPs). Eye movements were recorded using magnetic search coils. RESULTS: When viewing binocularly, both normal monkeys exhibited accurate, stereotyped symmetric convergence movements that achieved 87% to 96% of the required change in vergence angle by the end of the initial movement. In contrast, the small-eso monkey's convergence response when viewing binocularly was variable, strikingly asymmetric, usually accompanied by a disjunctive saccade, and subnormal, achieving only 56% of required vergence. The convergence response of the large-eso monkey was also asymmetric and weak, achieving 18% of the required vergence and employing conjugate saccades to refixate the near target. Monocular viewing (i.e., accommodative vergence) caused substantial reductions in both convergence amplitudes and velocities in the normal monkeys, but had a minor effect on the vergence behavior of the strabismic animals. CONCLUSIONS: Monkeys with small- and large-angle infantile esotropia have striking maldevelopments of binocular (disparity-driven) convergence and use accommodative vergence and saccades to refixate near targets. Their vergence behavior resembles that in esotropic humans. The maldevelopment may be explained in large part by the paucity of binocular connections recently described in the visual cortex of esotropic macaques.     

Field of vergence limits

The binocular visual field is the combined visual field obtained when the eyes are fixating a given point without head movements. The binocular field of fixation is the area within which central bifixation is possible by moving the eyes but not the head. This study introduces a new concept of the binocular field, namely, the vergence limit field, defined as the binocular field of fixation that allows for a specified amount of vergence eye movements at its limits. A newly developed instrument and technique were used to measure this field for a number of normal subjects.     

Individual differences in binocular coordination are uncovered by directly comparing monocular and binocular reading conditions

Purpose. We evaluated systematically binocular coordination during a reading task by comparing binocular and monocular reading, and considering the potential effects of individual heterophoria and eye dominance. Methods. A total of 13 participants (aged 19-29 years, refractive errors -0.5 to 0.125 diopters [D]) read single sentences in a haploscope while eye movements were measured with an EyeLinkII eyetracker. Results. When reading monocularly, saccade amplitudes increased by 0.04 degrees and first fixation durations became longer by approximately 10 ms. Furthermore, saccade disconjugacies increased, and compensatory vergence drifts during fixation turned into a divergent drift relative to the viewing distance. The vergence angle adjusted for the actual viewing distance became less convergent during monocular reading by 0.5 degrees. Moreover, in participants who were almost orthophoric, only the first fixation duration became longer (by 20 ms) when the reading conditions changed from binocular to monocular. For exophoric participants, all parameters of binocular coordination changed, and first fixation duration decreased by 20 ms. When reading monocularly, no differences between the dominant right eye and the nondominant left eye were found. Conclusions. Because of obvious differences in binocular coordination between monocular and binocular reading, some vergence adjustments are driven actively by fusional processes. Furthermore, higher demands on these binocular fusional processes can be uncovered only by a detailed evaluation of monocular reading conditions.     

角膜屈光手术对近视患者双眼视功能的影响

目的：探究角膜屈光手术对近视患者双眼视功能的影响及临床意义。方法：回顾性研究。选取2018年 1 月至2020 年4 月于长春爱尔眼科医院屈光手术中心行角膜屈光手术的患者82 例（164 眼）,年龄 18～32（23.5±4.2）岁。术前等效球镜度（SE）为-8.38～-0.88（-4.75±1.69）D。根据双眼SE差值 将患者分为2组：屈光参差组21例（42眼）,双眼SE差值≥1.00 D;非屈光参差组61例（122眼）,双眼 SE差值<1.00 D。所有患者行术前和术后1 周、1 个月、3 个月、6 个月常规眼科检查及调节功能（调 节幅度,正、负相对调节,单、双眼调节灵敏度和调节反应）与聚散功能（远、近水平隐斜视,正、 负融像性聚散范围和调节性集合/调节量）检测。组内比较采用重复测量方差分析,组间比较采用 双因素重复测量方差分析和独立样本t检验。相关性采用Spearman进行分析。结果：屈光参差组 的调节功能中左眼调节灵敏度在术后1 周较术前降低,术后1、3、6 个月均较术前增加（F=2.552, P=0.045）;聚散功能中近正融像破裂点和恢复点术后各时间点均较术前减小（F=4.245,P=0.016; F=3.686,P=0.009）。非屈光参差组调节功能中术后各时间点正相对调节和调节幅度较术前均增加 （F=16.118,P<0.001;F=14.839,P<0.001）;聚散功能中远隐斜在术后各时间点均较术前减小（F=2.898, P=0.029）,远负融像破裂点和恢复点、远正融像恢复点和近正融像破裂点术后各时间点均较术前减 小（F=7.310,P<0.001;F=5.131,P=0.001;F=8.184,P<0.001;F=10.919,P<0.001）。远正融像破裂 点和近正融像恢复点术后逐渐下降,术后1、6 个月相比术前减小,且差异有统计学意义（F=6.814, P<0.001;F=6.942,P<0.001）。近负融像恢复点术后1 个月相比术前明显增加,术后3、6 个月逐渐下 降,与术前差异无统计学意义（F=2.811,P=0.037）。屈光参差组和非屈光参差组右、左眼单眼调节 灵敏度和双眼调节灵敏度随时间变化差异均有统计学意义（右眼：F时间=0.555,P=0.655,F组别=4.639, P=0.034,F交互=0.555,P=0.655;左眼：F时间=2.611,P=0.048,F组别=5.709,P=0.019,F交互=1.287, P=0.279;双眼：F时间=2.256,P=0.086,F组别=4.314,P=0.041,F交互=0.847,P=0.465）。结论：近视患 者角膜屈光手术后双眼视功能均有不同程度改善或维持术前水平,证明角膜屈光手术对视功能有积 极作用。     

Eye-movement responses to disparity vergence stimuli with artificial monocular scotomas

The effects of artificial monocular scotomas on eye-movement responses to horizontal disparity vergence stimuli were studied in six subjects with normal binocular vision. Subjects viewed stereoscopic 1.5 degrees horizontal step disparity vergence stimuli through liquid crystal shutter glasses. The central portion of the stimulus presented to the right eye was removed to simulate monocular artificial scotomas of variable diameters (2 degrees to 10 degrees ). Eye movements were recorded with a binocular head-mounted eye tracker. Responses included pure vergence, vergence followed by saccades, and pure saccadic eye movements. The rate of responses with saccadic eye movements increased with the diameter of the artificial scotoma (p < 0.0001); there was an increase in the rate of responses starting with saccades (p < 0.0001), as well as an increase in the rate of saccades after initial vergence responses (p < 0.01). The probability of saccades after initial vergence responses was affected by the open-loop gain of the vergence response (p < 0.001). The open-loop gain decreased with increased diameters of the artificial scotomas (p < 0.0001). As the diameter of the artificial scotomas increased, the amplitude of the initial vergence eye-movement responses decreased, and the prevalence of saccadic eye movements and asymmetric vergence increased. The effects of the diameter of artificial monocular scotomas on eye-movement responses in subjects with normal binocular vision are consistent with the effects of diameter of suppression scotomas on eye-movement responses to disparity vergence stimuli in patients with infantile esotropia.     

Divergence limits within the field of vergence limits

The field of vergence limits has been defined as the binocular field of fixation that allows for a specified amount of vergence eye movements at its limits. Within this field, divergence amplitudes were measured in primary, secondary, and tertiary positions of gaze. Six subjects participated. It was found that divergence limits vary minimally with change in direction of gaze. The maximum amplitude of divergence was found 20 degrees below the primary straight ahead position.     

The influence of rapid prism adaptation upon fixation disparity.

Small errors of fusional vergence (fixation disparity) were examined as a function of the magnitude of horizontal prism stimulating convergence or divergence for a short (30sec) duration. Marked differences were observed between the amplitude of fixation disparity resulting from convergent and divergent stimulus disparities. In another experiment, subjects wore a horizontal prism for 30 sec after which time one eye was occluded for 40 sec. Measurements of vergence eye movements revealed an incomplete relaxation of fusional vergence (prism adaptation) after 40 sec of occlusion. Marked differences were observed between the amplitude of prism adaptation resulting from convergent and divergent stimuli. Maximum prism adaptationand minimum fixation disparity occurred with the same direction of prism, suggesting that a slow fusional vergence mechanism minimizes errors of binocular vergence.     

Are corresponding points fixed?

Several investigators have claimed that the retinal coordinates of corresponding points shift with vergence eye movements. Two kinds of shifts have been reported. First, global shifts that increase with retinal eccentricity; such shifts would cause a flattening of the horopter at all viewing distances and would facilitate fusion of flat surfaces. Second, local shifts that are centered on the fovea; such shifts would cause a dimple in the horopter near fixation and would facilitate fusion of points fixated at extreme viewing distances. Nearly all of the empirical evidence supporting shifts of corresponding points comes from horopter measurements and from comparisons of subjective and objective fixation disparity. In both cases, the experimenter must infer the retinal coordinates of corresponding points from external measurements. We describe four factors that could affect this inference: (1) changes in the projection from object to image points that accompany eye rotation and accommodation, (2) fixation errors during the experimental measurements, (3) non-uniform retinal stretching, and (4) changes in the perceived direction of a monocular point when presented adjacent to a binocular point. We conducted two experiments that eliminated or compensated for these potential errors. In the first experiment, observers aligned dichoptic test lines using an apparatus and procedure that eliminated all but the third error. In the second experiment, observers judged the alignment of dichoptic afterimages, and this technique eliminates all the errors. The results from both experiments show that the retinal coordinates of corresponding points do not change with vergence eye movements. We conclude that corresponding points are in fixed retinal positions for observers with normal retinal correspondence.     

Binocular fixation topography in patients with diabetic macular oedema

BACKGROUND: During retinal photocoagulation for diabetic maculopathy, there is a potential risk of foveal burns, and laser scars may later enlarge to be sight-threatening when involving retinal areas previously used during fixation. Since the retinal area used during binocular steady fixation has been found to vary considerably in the normal test person and central fixation may be even further compromised in patients with diabetic maculopathy, the sight-threatening side effects could possibly be reduced by taking into account the fixation area individually. However, no study has described and quantified the retinal area of fixation binocularly in patients with clinically significant macular oedema (CSME). METHODS: Sixteen diabetic patients with CSME in one or both eyes were examined. Each examination included visual acuity testing (ETDRS charts), a standard eye examination, central retinal thickness assessment by optical coherent tomography, fluorescein angiography and binocular quantification of fixational eye movements using an infrared recording technique. RESULTS: A negative correlation was found between visual acuity and mean microsaccadic amplitude (R=0.48, p=0.009). The maximal retinal extension of the fixation area ranged between 1.0 degrees and 3.0 degrees , and in two eyes with CSME, this area was estimated to exceed 800 mum on the retinal plane. No correlation was found between retinal thickness and visual acuity, retinal area of fixation, maximal extension of the fixation area or mean microsaccadic amplitude. CONCLUSION: Large interindividual differences in quantitative measures of binocular fixational eye movements were found. The mean amplitude of fixational eye movements was not correlated to central retinal thickness, and fixation area could only partly be predicted by visual acuity. Two eyes with CSME had an estimated maximal extension of the fixation area exceeding the central 800 mum on the retinal plane; thus, the possible benefit of individualising central photocoagulation according to precise measures of fixation area needs to be investigated on a larger population.     

Hering's law of equal innervation and the position of the binoculus

The movements of the eyes whilst changing fixation from near to distant vision were observed and objectively recorded. The magnitude of the binocular versional component was isolated from the change in vergence. The versional component was found to be greater in the dominant eye. This corresponds with the subjective observation of the physiological diplopic images. It is suggested that Hering's Law of equal innervation is applicable to asymmetrical movements only if the position of the binoculus is taken into account.     

Direction of heading and vestibular control of binocular eye movements.

To optimize visual fixation on near targets against translational disturbances, the eyes must move in compliance with geometrical constraints that are related to the distance as well as the speed and direction relative to the target. It is often assumed that the oculomotor system uses the vestibular signals during such movements mainly to stabilize the foveal image irrespective of the peripheral vision. To test this hypothesis, trained rhesus monkeys were asked to maintain fixation on isovergence targets at different horizontal eccentricities during 10 Hz oscillations along different horizontal directions. We found that the two eyes moved in compliance with the geometrical constraints of the gaze-stabilization hypothesis, although response gains were generally small ( approximately 0.5). The best agreement with the gaze stabilization hypothesis occurred for heading directions within +/-30 degrees from straight-ahead, whereas lateral movements exhibited greater variability and larger directional errors that reflected the statistical response variability inherent in the non-linear dependence on heading direction. In contrast to undercompensatory version (conjugate) components, the disjunctive part of the response (vergence) exhibited unity or higher than unity gains. The high vergence gains might reflect a strategy that aims at maintaining the binocular coordination of the gaze lines despite the low gain of the version movements.     

Is there a need for binocular vision evaluation in low vision?

The purpose of this study was to determine the need for a binocular vision evaluation in the management of low vision patients. Thirty subjects were selected with visual acuities of at least 6/60 in each eye. Twenty-three had age-related macular degeneration as the primary ocular pathology and seven had various other ocular conditions. The binocular vision evaluation comprised: detailed history and symptoms with an emphasis on the occurrence of near vision problems; modified binocular vision assessment which included cover test at near, ocular movements, convergence, Bagolini glasses, horizontal vergence reserves and prismatic measurement of any observed anomaly. History and symptoms revealed horizontal and vertical diplopia, jumbling of print, asthenopia, intermittent blurring and print being too small. Binocular vision assessment revealed exophoria at near, manifest horizontal deviations, convergence insufficiency, and vertical muscle imbalance. These anomalies occurred either singly or in combination. The results indicate that there is a need for binocular vision evaluation in low vision in order to reveal near vision disorders that may be partly attributable to binocular vision anomaly and not solely due to ocular pathology. This information should facilitate the implementation of the appropriate course of optometric management for these patients.     

Visual field defects for vergence eye movements and for stereomotion perception

An objective visual field can be mapped in terms of stimulus-induced eye movement. The authors used the scleral coil technique to record vergence and conjugate eye movements while stimulating different visual field locations with a 3 X 3 deg target whose image vergence was oscillated. For each of three subjects tested there was a visual field location where vergence eye movements were much weaker than in a control location of equal retinal eccentricity. On the other hand, conjugate eye movements driven from these two locations by lateral motion were similar. Field defects for ocular vergence coincided with regions in which oscillating retinal disparity failed to produce a sensation of motion in depth, although visual responses to static disparity were normal, and psychophysical thresholds for lateral motion showed no defect with either binocular or monocular viewing. It was concluded, therefore, that the perceptual stereomotion scotomata were not due to a monocular loss, but to a defective binocular interaction between motion signals from the left and right eyes, and that this defective interaction was specific for opposed rather than parallel motion in the two eyes. Furthermore, the visual loss was specific for motion rather than for position. The correlation between the field defects for ocular vergence and stereomotion perception leads the authors to suggest that the same defect in binocular interaction is responsible for both the eye movement and sensory abnormalities. Two candidate hypotheses are proposed: one is framed in terms of a single population, and the other in terms of two populations of cortical neurons.     

Dynamic visual fields of one-eyed observers.

BACKGROUND: The horizontal binocular visual field can extend to more than 200 degrees, while a monocular field is limited to 160 degrees. Additionally, the nose and other facial structures may block the monocular field further during certain eye movements. The purpose of this study was to compare the monocular against the binocular visual field and determine if head and eye movements can functionally overcome any measured deficit. METHODS: In Experiment 1, visual fields were measured monocularly with a bowl perimeter using 5 fixation positions. Binocular visual fields were calculated by combining the monocular visual field with its mirror image. In Experiment 2, subjects were allowed to make head, eye, and body movements to search for flashing lights 360 degrees around them, spaced every 45 degrees. The numbers of lights identified were compared for the subjects performing monocularly versus binocularly. RESULTS: The size of the overall monocular visual field was found to vary between 48% and 76% of the binocular visual field, depending on eye position. For the flashing light experiment, head and eye movements could not overcome the entire visual-field deficit with monocular viewing. Monocular performance remained 11.4% less than binocular performance. CONCLUSIONS: The visual-field deficit seen with monocular viewing is greatest with nasal fixation, and head and eye movements cannot totally compensate for this deficit when viewing time is limited. Vision standards that require full visual fields in each eye are more appropriate for occupations in which peripheral visual targets must be identified and visual search time is limited.     

Fixation disparity,accommodation, dark vergence and dark focus during inclined gaze*

We investigated several oculomotor functions at different angles of vertical inclination of the gaze direction from 15 deg upwards to 45 deg downwards. The mean accommodative resting state (measured in a dark visual field) increased when the eyes or the head were declined from 0 to 45 deg. Fixation disparity (the vergence error in minutes of arc relative to the principle visual directions) became more eso when a fusion target at a viewing distance of 40 cm was lowered: declining the gaze by 45 deg changed mean fixation disparity by 1.8 min arc with eye inclination (keeping the head upright), and by 0.9 min arc with head inclination (with eye position unchanged relative to the head). When the eyes were lowered, the individual rate of eso change in fixation disparity was correlated with the amount of the subjects’ near shifts in the resting position of vergence, measured in darkness. Significant test-retest correlations between repeated measurements showed that the effects of eye inclination on vergence varied in a reproducible way among individuals with good binocular vision.     

Asymmetrical adaptation of human saccades to anisometropic spectacles

We report the extent to which effective asymmetrical saccadic adaptation was achieved by a myopic subject, who was exposed to "long-term" adaptation as he wore anisometropic corrective spectacles for about 40 years and also the extent of "short-term" adaptation in this subject and two other subjects, who initially made conjugate saccades, when they wore newly fitted anisometropic spectacles for about 8 hr. Two-dimensional binocular eye positions were measured with an accurate and precise revolving magnetic field-sensor coil technique. We found that long-term adaptation of vertical saccades was virtually perfect (almost 100% of the asymmetry introduced by the spectacles was corrected). Long-term adaptation of horizontal saccades was less complete and increased with target separation from about 40% for saccadic amplitudes of 5 degrees to about 75% for amplitudes of 60 degrees. Short-term adaptation of vertical saccades was virtually complete (100%) in one newly fitted subject and only partially complete (40%) in the other two subjects. The persistence of the adaptive asymmetry of saccades during monocular viewing showed that adaptation derived from plasticity in the programming of saccades and not from modification of vergence responses. Without the anisometropic spectacles, 30 min of self-paced, one per second changes in binocular fixation between two targets, which required a version change of 45 degrees in combination with a vergence change of 11 degrees, did not induce any asymmetrical adaptation. This result shows that a specific repeated association of version and vergence eye movements was not sufficient to induce asymmetrical adaptation, leading us to suggest that the transient fixation disparities at saccade-offset might be the necessary stimulus for the asymmetrical saccadic adaptation we observed.     

Negative feedback control model of proximal convergence and accommodation

A comprehensive model has been developed to illustrate the interactions between the observer and the surrounding environment in the control of oculomotor responses to distance or 3-D space. Accommodation and vergence respond to both spatiotopic (body reference) proximal percepts and retinotopic (eye referenced) physical stimuli of blur and disparity. Both spatiotopic and retinotopic stimuli are derived respectively from perceptual and physical correlates of negative feedback for eye position. The spatiotopic and retinotopic stimulus errors are combined in the feedforward path and drive a common oculomotor controller which has a phasic-tonic organization. Spatiotopic and retinotopic stimuli are shown to be effective over complementary operating ranges. Perceptual spatiotopic errors of gaze provide optimal stimuli for near responses to large depth intervals whereas physical-retinotopic cues of blur and disparity provide quantitative information about small binocular fixation errors. Small dynamic variations of target distance are sensed both spatiotopically and retinotopically. Coarse and fine spatiotopic errors of gaze are processed differently. Large spatiotopic errors are sampled intermittently at the beginning of the near response, whereas small retinotopic position errors and spatiotopic velocity errors are sampled continuously throughout the near response. Former reports of empirically observed higher velocity of vergence responses to very large depth intervals is explained in terms of stimulus sampling modes rather than in terms of separate oculomotor control mechanisms. The model demonstrates a complementary function of top-down spatiotopic cues, which are used to initiate the near response, and bottom-up retinotopic cues, which are used to refine and complete the near response. Cross-couplings by vergence-accommodation and accommodative-vergence serve to coordinate the components of the near response when feedback from sensed response of one motor system (i.e. vergence) is more accurate than that of the other motor system (i.e. accommodation). The model presented here is concerned primarily with the near response mediated by accommodation and disjunctive eye movements and not by the independent vergence mediated by non-conjugate or yoked saccades of unequal amplitude.     

Inhibition of saccade and vergence eye movements in 3D space

Inhibitory capacity was investigated by measuring the eye movements of normal subjects asked to fixate a central point, and to suppress eye movements toward visual distracters appearing in the periphery or in depth. Eight right-handed young adults performed such a suppression or distracter task. In different conditions, the distracter could appear at 10 degrees left or right at a distance of 20, 40, or 150 cm (calling for horizontal saccades), or in a central position far or close (calling for convergence or divergence), or 7.5 degrees up or down at 40 or 150 cm (calling for vertical saccades). Eye movements were recorded binocularly with an infrared light eye-movement device. Results showed that (1) suppression performance was not perfect, as the subjects still produced eye movements; (2) errors were distributed unequally in three-dimensional space, with more frequent errors toward distracters calling for convergence, or leftward and downward saccades at a close distance; (3) distracters calling for saccade suppression yielded saccades in the direction of the distracter (that we called prosaccades), and saccades directed away from it (that we called spontaneous antisaccades); (4) for vergence, only distracters calling for convergence yielded errors, which were always promovements; (5) in addition, a small convergent drift was found for convergence distracters. Differences in the errors between saccade and vergence suggest that different inhibitory mechanisms may be involved in the two systems. Spatial left/right, up/down, and close/far asymmetries are interpreted in terms of attentional biases.