Eye as a key element of conspecific image eliciting lateralized response in fish

Visual lateralization in different aspects of social behaviour has been found for numerous species of vertebrates ranging from fish to mammals. For inspection of a shoal mate, many fishes show a left eye–right hemisphere preference. Here, we tested the hypothesis that in fish, there is a key cue in the conspecific appearance, which elicits lateralized response to the whole image of the conspecific. In a series of eight experiments, we explored eye preferences in cryptic-coloured Amur sleeper, Perccottus glenii, fry. Fish displayed left-eye preferences at the population level for inspection of a group of conspecifics, their own mirror image, and a motionless flat model of a conspecific. In contrast, no population bias was found for scrutinizing an empty environment or a moving cylinder. When fry were showed a model of a conspecific in a lateral view with the eye displaced from the head to the tail, they again showed a significant preference for left-eye use. On the other hand, ‘eyeless’ conspecific model elicited no lateralized viewing in fry. Finally, the left-eye preference was revealed for scrutiny of the image of a conspecific eye alone. We argue that in Amur sleeper fry, eye is the element of the conspecific image, which can serve as a ‘key’ for the initiation of lateralized social response. This key element may serve as a trigger for the rapid recognition of conspecifics in the left eye–right hemisphere system. Possible causes and advantages of lateralized perception of social stimuli and their key elements are discussed in the context of current theories of brain lateralization.     

Functional lateralization, interhemispheric transfer and position bias in serial reversal learning in pigeons (Columba livia)

In the present study we investigated lateralization of color reversal learning in pigeons. After monocular acquisition of a simple color discrimination with either the left or right eye, birds were tested in a serial reversal procedure. While there was only a slight and non-significant difference in choice accuracy during original color discrimination, a stable superiority of birds using the right eye emerged in serial reversals. Both groups showed a characteristic ‘learning-to-learn’ effect, but right-eyed subjects improved faster and reached a lower asymptotic error rate. Subsequent testing for interocular transfer demonstrated a difference between pre- and post-shift choice accuracy in pigeons switching from right to left eye but not vice versa. This can be accounted for by differences in maximum performance using either the left or right eye along with an equally efficient but incomplete interocular transfer in both directions. Detailed analysis of the birds’ response patterns during serial reversals revealed a preference for the right of two response keys in both groups. This bias was most pronounced at the beginning of a session. It decreased within sessions, but became more pronounced in late reversals, thus indicating a successful strategy for mastering the serial reversal task. Interocular transfer of response patterns revealed an unexpected asymmetry. Birds switching from right to left eye continued to prefer the right side, whereas pigeons shifting from left to right eye were now biased towards the left side. The results suggest that lateralized performance during reversal learning in pigeons rests on a complex interplay of learning about individual stimuli, stimulus dimensions, and lateralized response strategies. Received: 4 June 1999 / Accepted after revision: 18 August 1999     

A simple technique for lateralizing visual input that allows prolonged viewing

A simplified technique is described for obtaining lateralization of visual input with prolonged viewing. This technique is based on the presence of constant normal lateral limits for horizontal rotation of the eyes with reference to the head. With head movement prevented by use of a standard bite bar, and the eyes rotated to the left and held at their lateral limit, the temporal half of the visual field of the left eye may be used for lateralized input to the right hemisphere or vice versa for input to the left hemisphere. Any form of visual stimuli or visually monitored task can be used if confined within one of the extreme temporal hemifields. In comparison to previous methods, this technique is technically simple, inexpensive, without significant risk or discomfort to the subject, readily applicable to normal and various brain-lesioned subjects, and permits prolonged in-depth viewing. An alternative version of this technique uses a stabilized spectacle frame fitted with adjustable central occluders set to allow vision through only one or both of the extreme temporal hemifields.     

Hemispheric predominance assessment of phonology and semantics: a divided visual field experiment

The aim of the present behavioural experiment was to evaluate the most lateralized among two phonological (phoneme vs. rhyme detection) and the most lateralized among two semantic ("living" vs. "edible" categorization) tasks, within the dominant hemisphere for language. The reason of addressing this question was a practical one: to evaluate the degree of the hemispheric lateralization for several language tasks, by using the divided visual presentation of stimuli, and then choose the most lateralized semantic and phonological for mapping language in patients by using fMRI in future studies. During the divided visual field experiment by using words (semantic tasks) and pseudo-words (phonological tasks) as stimuli, thirty-nine right-handed participants were examined. Our results have shown that all tasks were significantly left hemisphere lateralized. Furthermore, the rhyme was significantly more lateralized than phoneme detection and "living" was significantly more lateralized than "edible" categorization. The rhyme decision and "living" categorization will be used in future fMRI studies for assessing hemispheric predominance and cerebral substrate for semantics and phonology in patients. Our results also suggest that the characteristics of stimuli could influence the degree of the hemispheric lateralization (i.e., the emotional charge of stimuli for words and the position of the phoneme to be detected, for pseudo-words).     

Lateralized interhemispheric transfer of color cues: evidence for dynamic coding principles of visual lateralization in pigeons

Visual feature discrimination tasks in pigeons reveal a right eye/left hemisphere dominance at the population level. Anatomical studies and lesion data show this visual lateralization to be related to asymmetries of the tectofugal system, which ascends from the tectum over the n. rotundus to the forebrain. Anatomically, this system is characterized by numerous morphological and connectional asymmetries which result in a bilateral visual representation in the dominant left hemisphere and a mostly contralateral representation in the subdominant right hemisphere. Ontogenetically, visual lateralization starts with an asymmetrical embryonic position within the egg, which leads to asymmetries of light stimulation. Differences in exposure to light stimulation between the eyes result in activity differences between the ascending tectofugal pathways of the left and the right hemisphere, which are transcribed during a critical time span into morphological asymmetries. The asymmetries established after this transient period finally start to determine the lateralized processes of the visual system for the entire life span of the individual. We now can show that these anatomical lateralizations are accompanied by asymmetries of interocular transfer, which enable a faster shift of learned color cues from the dominant right to the left eye than vice versa. In summary, our data provide evidence that cerebral asymmetries are based both on "static" anatomical and on "dynamic" process-dependent principles.     

Aesthetic preference and lateral dominance

Aesthetic preferences for photographs with the main focal content either to the left or right of the photograph's center were examined in right- and left-handed subjects. Verbal responses or manual responses were required. In one experiment with 261 introductory psychology student-subjects, left-handers more often preferred photographs with the more important part on the left ("left-geared") than did right-handers. Exp. 2, involving 84 right-handed student subjects, showed that left-geared photographs presented on the left side were preferred more often than left-geared photographs presented on the right side, and left-geared photographs presented on the left side were more often chosen when a left-handed manual response was required. Interactions between handedness, position of the stimulus, language hemisphere, and response mode make it extremely difficult to ascertain whether the right hemisphere is really more involved in aesthetic decisions.     

Human gestures trigger different attentional shifts in chimpanzees (Pan troglodytes) and humans (Homo sapiens)

Brain lateralization has been the matter of extensive research over the last centuries, but it remains an unsolved issue. While hand preferences have been extensively studied, very few studies have investigated laterality of eye use in non-human primates. We examined eye preference in 14 Campbell’s monkeys (Cercopithecus c. campbelli). We assessed eye preference to look at a seed placed inside a tube using monocular vision. Eye use was recorded for 100 independent and non-rewarded trials per individual. All of the 14 monkeys showed very strong preferences in the choice of the eye used to look inside the tube (mean preference: 97.6%). Eight subjects preferred the right eye and six subjects preferred the left eye. The results are discussed in light of previous data on eye preference in primates, and compared to data on hand preference from these subjects. Our findings would support the hypothesis for an early emergence of lateralization for perceptual processes compared to manual motor functions.     

音高模式加工的大脑偏侧化研究

作为一种超音段成分,音高在言语的理解和产生过程中承担着重要作用,这些作用主要包括:区别不同声调的词汇意义,区别陈述、疑问等不同的语调,标志话语的情绪色彩等。音段、语义和句法信息的加工具有左脑优势,但音高模式加工的单侧优势及其机制还存在很多争论,本文从音高模式加工的两个主要理论———功能假设和声音信号假设出发,对相关研究加以评述。     

Lateralization of visuospatial attention across face regions varies with emotional prosody

It is well-established that linguistic processing is primarily a left-hemisphere activity, while emotional prosody processing is lateralized to the right hemisphere. Does attention, directed at different regions of the talker’s face, reflect this pattern of lateralization? We investigated visuospatial attention across a talker’s face with a dual-task paradigm, using dot detection and language comprehension measures. A static image of a talker was shown while participants listened to speeches spoken in two prosodic formats, emotional or neutral. A single dot was superimposed on the speaker’s face in one of 4 facial regions on half of the trials. Dot detection effects depended on emotion condition—in the neutral condition, discriminability was greater for the right-, than for the left-, side of the face image, and at the mouth, compared to the eye region. The opposite effects occurred in the emotional prosody condition. The results support a model wherein visuospatial attention used during language comprehension is directed by the left hemisphere given neutral emotional prosody, and by the right hemisphere given primarily negative emotional prosodic cues.     

Divided Visual Attention and Left Hemisphere Activation Among Psychopathic and Nonpsychopathic Offenders

Recent divided attention studies suggest the impulsive antisocial behavior of psychopaths may be related to deficient processing of information under conditions that place substantial demands on left-hemisphere-specific processing resources (left hemisphere activation (LHA) hypothesis; D. S. Kosson [1996, 1998]). To examine performance under conditions constraining eye movements and covert shifts of attention, 26 psychopathic and 46 nonpsychopathic inmates completed a divided attention task in which lateralized stimuli appeared briefly and simultaneously in the left visual field (LVF) and right visual field (RVF) of a monitor. Targets were either presented primarily in the RVF to induce LHA or were equiprobable in LVF and RVF to promote equal activation (EA) of left and right hemisphere resources. Psychopaths were less accurate than nonpsychopaths only under LHA conditions; within-group comparisons also revealed a substantial decrement from EA to LHA only in psychopaths. Thus, psychopaths’ performance deficits were specific to conditions priming left hemisphere resources asymmetrically.     

Differential visual field-interhemispheric transfer: can it explain sex and handedness differences in lateralization?

104 men and women were tested for visual field-hemispheric transfer of spatial information on a dot-localization task. Right-handed subjects showed significant improvement when stimuli were presented to the left visual field of the right hemisphere (LVF-RH) after practice on the same task presented to the right visual field of the left hemisphere (RVF-LH) first. No improvement was found when the task was presented in the reverse order (LVF-RH first followed by RVF-LH). It was concluded that, for right-handers, transfer of spatial information to the right hemisphere is facilitated while transfer to the left hemisphere is inhibited. Left-handed subjects demonstrated no significant improvement in either condition, suggesting inhibition or lack of transfer of spatial information in either direction. No sex differences were found in either right-handed or left-handed subjects. The findings suggest that there may be different mechanisms underlying the similarities in functional lateralization of women and left-handers.     

A right visual field advantage for visual processing of manipulable objects

Information about object-associated manipulations is lateralized to left parietal regions, while information about the visual form of tools is represented bilaterally in ventral occipito-temporal cortex. It is unknown how lateralization of motor-relevant information in left-hemisphere dorsal stream regions may affect the visual processing of manipulable objects. We used a lateralized masked priming paradigm to test for a right visual field (RVF) advantage in tool processing. Target stimuli were tools and animals, and briefly presented primes were identical to or scrambled versions of the targets. In Experiment 1, primes were presented either to the left or to the right of the centrally presented target, while in Experiment 2, primes were presented in one of eight locations arranged radially around the target. In both experiments, there was a RVF advantage in priming effects for tool but not for animal targets. Control experiments showed that participants were at chance for matching the identity of the lateralized primes in a picture?Cword matching experiment and also ruled out a general RVF speed-of-processing advantage for tool images. These results indicate that the overrepresentation of tool knowledge in the left hemisphere affects visual object recognition and suggests that interactions between the dorsal and ventral streams occurs during object categorization.     

Evolution of hemispheric specialization: advantages and disadvantages

Lateralization of the brain appeared early in evolution and many of its features appear to have been retained, possibly even in humans. We now have a considerable amount of information on the different forms of lateralization in a number of species, and the commonalities of these are discussed, but there has been relatively little investigation of the advantages of being lateralized. This article reports new findings on the differences between lateralized and nonlateralized chicks. The lateralized chicks were exposed to light for 24 h on day 19 of incubation, a treatment known to lead to lateralization of a number of visually guided responses, and the nonlateralized chicks were incubated in the dark. When they were feeding, the lateralized chicks were found to detect a stimulus resembling a raptor with shorter latency than nonlateralized chicks. This difference was not a nonspecific effect caused by the light-exposed chicks being more distressed by the stimulus. Instead, it appears to be a genuine advantage conferred by having a lateralized brain. It is suggested that having a lateralized brain allows dual attention to the tasks of feeding (right eye and left hemisphere) and vigilance for predators (left eye and right hemisphere). Nonlateralized chicks appear to perform these dual tasks less efficiently than lateralized ones. Reference is made to other species in discussing these results.     

Ear differences in simple reaction time: The influence of attentional factors

Simple motor reaction times (right-handed) to tape-recorded consonant-vowel syllables were obtained from 12 subjects under two conditions of monotic stimulation: “expected” presentation (subject informed as to ear of presentation) and “random” presentation (ear of presentation varied randomly). Significantly faster left ear reaction times were obtained in the “expected” condition. The rank order of subjects' standard dichotic listening task scores correlated negatively with reaction time ear differences in the “expected” presentation condition. Results are discussed in terms of existing theories of response lateralization in simple reaction time tasks, and an expanded hypothesis is offered. Specifically, it is suggested that two or more distinct but potentially overlapping mechanisms may be responsible for observed asymmetries in monotic auditory perception. Both an attentional or lateralized motor response bias and an inherent lateralization of function may operate side by side, differentially activated by task demands, mode of stimulus presentation, and nature of stimuli.     

Functional lateralization of the human premotor cortex during sequential movements

A neurological truism is that each side of the brain controls movements on the opposite side of the body. Yet some left hemisphere brain lesions cause bilateral impairment of complex motor function and/or ideomotor apraxia. We report that the left dorsal premotor cortex of normal right-handed people plays a fundamental role in sequential movement of both right and left hands. Subjects performed sequential finger movements during functional magnetic resonance imaging of the motor cortices. In right-handed subjects, the volume of activated dorsal premotor cortex showed a left hemispheric predominance during hand movements. We suggest that the observed left premotor dominance contributes to the lateralization found in lesion studies.     

The laterality of dreaming

The cortical locus of dream generation could be lateralized to the right or left hemisphere, or be bilaterally represented with either equal or unequal contributions from each hemisphere. In this paper we review the neurological literature for cases of loss or alteration of dream report after brain damage. The distribution of lesion sites is used to test the various hypotheses concerning the laterality of dreaming. The hypothesis receiving best support is that dreaming is lateralized to the left hemisphere in individuals with typical neurologic organization.     

Hemispheric lateralization of functions related to emotion

We have reviewed the evidence that processes and functions related to perception and expression of emotions are represented asymmetrically in the cerebral hemispheres. The literature describes three possible aspects of emotional lateralization: that emotions are better recognized by the right hemisphere; that control of emotional expression and related behaviors takes place principally in the right hemisphere; and that the right hemisphere is specialized for dealing with negative emotions, while the left is specialized for dealing with positive emotions. Evidence for the three hypotheses derives from methodologically diverse studies in unimpaired, brain-lesioned, and mood-disordered populations. Relatively little of the work has been precisely replicated, and conclusions rest on parallel lines of evidence from diverse sources. The present level of knowledge suggests a model of emotional control based on interactive inhibition between a right negatively biased and left positively biased hemisphere. However, the details of such a model, including the precise conditions under which emotion-related functions are lateralized, and the mechanisms of such lateralization have yet to be elucidated.     

The cost of action miscues: hemispheric asymmetries

Adaptive behaviors require preparation and when necessary inhibition or alteration of actions. The right hemisphere has been posited to be dominant for preparatory motor activation. This experiment was designed to learn if there are hemispheric asymmetries in the control of altered plans of actions. Cues, both valid and invalid, which indicate the hand most likely to be called onto respond, as well as the imperative stimuli that indicate the actual response hand, were presented to either the right or left visual fields of 14 normal right handed participants. The delay after a miscue is dependent on the time taken to inhibit the premotor and motor systems of the incorrectly activated hemisphere, as well as to activate the motor systems of the opposite hemisphere, which might have been interhemispherically inhibited by this miscue. Analyses of reaction times revealed that miscues presented in left hemispace (right hemisphere) cost more time than those miscues presented in right hemispace (left hemisphere), suggesting that activation of the preparatory systems controlled by the right hemisphere may take longer to reverse than those controlled by the left hemisphere. This asymmetry may be related to asymmetries in the strength of hemispheric activation with contralateral inhibition.     

Cerebral asymmetry in visual attention

The traditional view of cerebral lateralization of various cognitive functions has been challenged by results from recent experimental and clinical studies. Evidence has been gathered to suggest that a seemingly unitized cognitive function can be further broken down into various processing subcomponents which are distributed across the two hemispheres. For instance, according to such a more complex conceptualization of cerebral lateralization, language is seen not as a unitary ability, but rather as a collection of syntactic, semantic, and prosodic components, with each lateralized in particular manners. In much the same way, the present study attempts to examine the cerebral lateralization patterns of the seemingly unitary visual perception process. In a visual half-field experiment, 20 normal subjects were asked to make same/different judgments to laterally presented arrays of stimuli of the same type as previously studied by Treisman and her colleagues in experiments attempting to separate the preattentive and attentive stages of visual perception. Hemispheric differences were obtained only in tasks requiring attentive processing (i.e., Treisman's glueing). Results indicate a local attentional strategy for the left hemisphere and a global attentional strategy for the right hemisphere.     

Visual laterality in belugas (Delphinapterus leucas) and Pacific white-sided dolphins (Lagenorhynchus obliquidens) when viewing familiar and unfamiliar humans

Lateralization of cognitive processes and motor functions has been demonstrated in a number of species, including humans, elephants, and cetaceans. For example, bottlenose dolphins (Tursiops truncatus) have exhibited preferential eye use during a variety of cognitive tasks. The present study investigated the possibility of visual lateralization in 12 belugas (Delphinapterus leucas) and six Pacific white-sided dolphins (Lagenorhynchus obliquidens) located at two separate marine mammal facilities. During free swim periods, the belugas and Pacific white-sided dolphins were presented a familiar human, an unfamiliar human, or no human during 10–15 min sessions. Session videos were coded for gaze duration, eye presentation at approach, and eye preference while viewing each stimulus. Although we did not find any clear group level lateralization, we found individual left eye lateralized preferences related to social stimuli for most belugas and some Pacific white-sided dolphins. Differences in gaze durations were also observed. The majority of individual belugas had longer gaze durations for unfamiliar rather than familiar stimuli. These results suggest that lateralization occurs during visual processing of human stimuli in belugas and Pacific white-sided dolphins and that these species can distinguish between familiar and unfamiliar humans.