LATERAL DIFFERENCES IN NORMAL MAN AND LATERALIZATION OF BRAIN FUNCTION*

The historical development of work on lateral differences (LDs) is described and an evaluation is attempted of the present state of the study and of the conceptual and methodological problems which it encounters. Contemporary work is mainly motivated by the notion that LDs reflect hemispheric specialization and thus provide a means of studying hemisphere function in easily available normal subjects. Work on LDs in visual recognition has first been conducted, however, within a completely different conceptual framework. Right visual field advantage (RVFA) for words and letters was first thought of as reflecting the left-to-right direction of latin writing which produces opportunity for differential perceptual learning in the two hemifields or, as was later considered, creates left-to-right scanning habits. On the contrary, right ear advantage (REA) in dichotic listening was, from its discovery by Kimura, shown to be linked to lateralization of speech control in the left hemisphere. The possibility that visual field effects could also be related to hemispheric specialization was then examined by looking for effects of handedness and for correlations between LDs and pathological data. After considerable initial resistance, the notion has become widely accepted. While the early work tended to deal with broad categories of tasks or of stimulating materials, more analytical approaches have been developed and investigators have tried to specify the component operations which are responsible for observed LDs. It has been shown that a same task can give rise to different patterns of asymmetry according to the particular operating mode which is adopted. Chronometric methods have been used to isolate processing stages with different lateralizations, and specific hypotheses have been advanced concerning levels at which lateralization originates. Regarding the mechanism of LDs, the most often considered interpretation is based on the notion of direct access, i.e. of an advantage associated with primary projection in the competent hemisphere. According to whether localization of the critical operations is strict or relative, one of two alternative versions of direct access, called respectively the callosal relay and efficiency models, apply. Other mechanisms based on hemispheric activation or priming and on interference between operations competing for hemispheric processing capacity have also been considered and presumably contribute, beside direct access, to observed LDs. There are also other sources for LDs than brain asymmetry, a fact which tends to be overlooked with the current focusing of interest on hemispheric interpretations. Even in cases where the role of hemispheric specialization has been established, other determinants can play amplifying or masking roles, as has been shown for auditory laterality effects. Current attempts at providing simple dichotomous general characterizations of hemisphere function, such as the one in terms of holistic vs. analytic processing, are described. The opinion is offered that such attempts are premature and that they reflect an underestimation of the complexity of the problem. It is argued (1) that the present catalogue of registered LDs is both too narrow and too subject to selective biases to provide a basis for ambitious generalizations; (2) that a convincing account of lateralization would require an understanding of its evolutionary origin; (3) that it would also require a more advanced understanding of the various mechanisms, including interhemispheric interactions, intervening between hemisphere competence and performance.     

Hemispheric function and linguistic skill in the deaf

The nature of hemispheric processing in the prelingually deaf was examined in a picture-letter matching task. It was hypothesized that linguistic competence in the deaf would be associated with normal or near-normal laterality (i.e., a left hemisphere advantage for analytic linguistic tasks). Subjects were shown a simple picture of a common object (e.g., lamp), followed by brief unilateral presentation of a manually signed or orthographic letter, and they had to indicate as quickly as possible whether the letter was present in the spelling of the object's label. While hearing subjects showed a marked left hemisphere advantage, no such superiority was found for either a linguistically skilled or unskilled group of deaf students. In the skilled group, however, there was a suggestion of a right hemisphere advantage for manually signed letters. It was concluded that while hemispheric asymmetry of function does not develop normally in the deaf, the absence of this normal pattern does not preclude the development of the analytic skills needed to deal with the structure of language.     

A Voxel-based lesion study on facial emotion recognition after circumscribed prefrontal cortex damage

Previous studies have shown inconsistent findings regarding the contribution of the different prefrontal regions in emotion recognition. Moreover, the hemispheric lateralization hypothesis posits that the right hemisphere is dominant for processing all emotions regardless of affective valence, whereas the valence specificity hypothesis posits that the left hemisphere is specialized for processing positive emotions while the right hemisphere is specialized for negative emotions. However, recent findings suggest that the evidence for such lateralization has been less consistent. In this study, we investigated emotion recognition of fear, surprise, happiness, sadness, disgust, and anger in 30 patients with focal prefrontal cortex lesions and 30 control subjects. We also examined the impact of lesion laterality on recognition of the six basic emotions. The results showed that compared to control subjects, the frontal subgroups were impaired in recognition of three negative basic emotions of fear, sadness, and anger – regardless of the lesion laterality. Therefore, our findings did not establish that each hemisphere is specialized for processing specific emotions. Moreover, the voxel-based lesion symptom mapping analysis showed that recognition of fear, sadness, and anger draws on a partially common bilaterally distributed prefrontal network.     

An investigation of bilateral asymmetries in electrodermal activity

Research in cerebral laterality supports the idea that functional differences between the left and right hemispheres exist with respect to cognitive style and perceptual ability. Related research, which has examined autonomic nervous system (ANS) correlates of cerebral laterality, suggests that a component of the ANS, electrodermal activity (EDA), is also lateralized. Some findings in the literature report the occurrence of bilateral asymmetries in phasic and tonic EDA as a function of a left or right hemisphere preference for information processing. This experiment used normal male subjects who were either left or right movers in a test of Conjugate Lateral Eye Movement (CLEM). This selection procedure served to maximize a subject’s preference for either a right or left hemisphere mode of information processing. Bilateral EDA was recorded continuously while subjects performed a visual recognition task using word (left hemisphere) and shape (right hemisphere) stimuli. The data do not support the contention that performance on a procedure chosen to selectively activate a given hemisphere elicits asymmetric tonic and/or phasic EDA. The results show no significant difference in the frequency of elicited skin conductance responses under either of the experimental conditions. Bilateral tonic EDA rose continuously over time and did not vary in either hand as a function of task. The present results fail to offer support for either of the hypotheses which argue for I) increased contralateralexcitation with selective hemispheric activation, or 2) increased contralateralinhibition of the EDR with hemispheric arousal.     

Role of task factors in visual field asymmetries

Any tachistoscopic study of cerebral hemisphere asymmetry imposes a variety of task demands on the participants, ranging from demands imposed by specific viewing conditions to demands imposed by response output requirements. The present article discusses the role of several task factors that influence processing after the initial reception of the stimulus input, suggests a theoretical rationale for some of the effects of these task factors, and considers implications for future studies of visual laterality. The task factors discussed include both those that are relevant for minimizing artifacts that have nothing to do with hemispheric asymmetry and those that are relevant for interpreting hemispheric asymmetry in terms of specific perceptual and cognitive processes.     

Recognition of Facial Expression in Diverging Socioeconomic Levels

Cerebral organization has been seen to vary in different social or ethnic groups; also, sociocultural retardation is a serious obstacle to development in poorer countries. The ability of the right hemisphere to recognize facial emotions in a tachistoscopic task was tested in university and low-socioeconomic-level subjects. All were male and right-handed. The university students recognized facial emotions projected to their left field-right hemisphere significantly better than with their right field-left hemisphere; the low-socioeconomic-level subjects showed no differences in hemispheric performance. The explanation of laterality as a function of social class may be of importance in dealing with sociocultural retardation.     

Uncoupled leftward asymmetries for planum morphology and functional language processing

Explanations for left hemisphere language laterality have often focused on hemispheric structural asymmetry of the planum temporale. We examined the association between an index of language laterality and brain morphology in 99 normal adults whose degree of laterality was established using a functional MRI single-word comprehension task. The index of language laterality was derived from the difference in volume of activation between the left and right hemispheres. Planum temporale and brain volume measures were made using structural MRI scans, blind to the functional data. Although both planum temporale asymmetry (t(1,99) = 6.86, p < .001) and language laterality (t(1,99) = 15.26, p < .001) were significantly left hemisphere biased, there was not a significant association between these variables (r(99) = .01,ns). Brain volume, a control variable for the planum temporale analyses, was related to language laterality in a multiple regression (beta = -.30, t = -2.25, p < .05). Individuals with small brains were more likely to demonstrate strong left hemisphere language laterality. These results suggest that language laterality is a multidimensional construct with complex neurological origins.     

Socially dependent auditory laterality in domestic horses (Equus caballus)

Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener’s arousal on auditory processing and laterality.     

Right Hemisphere Involvement in Processing Later-Learned Languages in Multilinguals

In two experiments, multilingual Papua New Guinean subjects were tested using a divided visual field technique designed to determine hemispheric laterality for English and for Tok Pisin. Various factors including age of acquisition, proficiency, mode of instruction, and number of years that the language had been used were considered in relation to language laterality. Only age of acquisition proved to be a significant contributor to the laterality effects obtained; older acquirers of both English and Tok Pisin showed greater right hemisphere involvement than early acquirers. Although proficiency did not seem to be related to language laterality, it too was systematically affected by acquisition age. Older acquirers of English performed significantly poorer than younger acquirers on all four of the language-usage tests given. The strong influence of acquisition age on cerebral laterality for language and proficiency is interpreted as supporting a critical period for language learning.     

Lateralization of cognitive processes in the brain

The lateralization of cognitive processes in the brain is discussed. The traditional view of a language-visuo/spatial dichotomy of function between the hemispheres has been replaced by more subtle distinctions. The use of magnetic resonance imaging (MRI) to study brain morphology has resulted in a renewed focus on the relationship between structural and functional asymmetry. Focus has been on the role played by the planum temporale area in the posterior part of the superior temporal gyrus for language asymmetry, and the possible significance of the larger left planum. The dichotic listening technique is used to illustrate the difference between bottom-up, or stimulus-driven laterality versus top-down, or instruction-driven laterality. It is suggested that the hemispheric dominance observed at any time is the sum result of the dynamic interaction between bottom-up and top-down processing tendencies. Stimulus-driven laterality dominance is always monitored and modulated through top-down cognitive processes, like shifting of attention and changes in arousal. A model of top-down modulation of bottom-up laterality is presented with special reference to the understanding of psychiatric disorders.     

Motivation, affect, and hemispheric asymmetry: power versus affiliation

In 4 experiments, the authors examined to what extent information related to different social needs (i.e., power vs. affiliation) is associated with hemispheric laterality. Response latencies to a lateralized dot-probe task following lateralized pictures or verbal labels that were associated with positive or negative episodes related to power, affiliation, or achievement revealed clear-cut laterality effects. These effects were a function of need content rather than of valence: Power-related stimuli were associated with right visual field (left hemisphere) superiority, whereas affiliation-related stimuli were associated with left visual field (right hemisphere) superiority. Additional results demonstrated that in contrast to power, affiliation primes were associated with better discrimination between coherent word triads (e.g., goat, pass, and green, all related to mountain) and noncoherent triads, a remote associate task known to activate areas of the right hemisphere.     

Hemispheric specialization for reading

Behavioral laterality tasks with linguistic stimuli were used to assess the differential processing efficiencies of the cerebral hemispheres in right- and left-handed adults. Findings from a lateralized lexical decision task with concrete nouns supported Zaidel's (1983) "direct access" model of hemispheric functioning. A dual task consisting of oral and silent reading indicated that the right hand was significantly more disrupted than the left during unimanual finger tapping; however, some bilateral interference was observed. Taken together the findings suggest that although the left hemisphere was relatively more efficient, the right hemisphere was dynamically involved in the reading process.     

Laterality of cerebral function: relations with anxiety, repression, and perception

Based on two factor analyses of data from 67 right-handed undergraduates, a laterality quotient based on forward memory spans, spatial minus digit, divided by their sum, was devised. The hemispheric balance factor for which this quotient served as the principal marker had significant positive loadings by the laterality quotient for the Edinburgh Handedness Questionnaire and by a measure of repressive tendencies, the Marlowe-Crowne Social Desirability Scale. Scores from this factor were positively related to global as opposed to analytic perception as measured by the Navon tachistoscopic task. This factor was unrelated to the trait anxiety and working short-term memory factors which also emerged. Trait anxiety was negatively related to global perception. The results raise the possibility that greater degrees of right-handedness may be related to the release of the nondominant right hemisphere from inhibition and that working short-term memory may not be lateralized.     

Sensory laterality in affiliative interactions in domestic horses and ponies (<Emphasis Type="Italic">Equus caballus</Emphasis>)

Many studies have been carried out into both motor and sensory laterality of horses in agonistic and stressful situations. Here we examine sensory laterality in affiliative interactions within four groups of domestic horses and ponies (N?=?31), living in stable social groups, housed at a single complex close to Vienna, Austria, and demonstrate for the first time a significant population preference for the left side in affiliative approaches and interactions. No effects were observed for gender, rank, sociability, phenotype, group, or age. Our results suggest that right hemisphere specialization in horses is not limited to the processing of stressful or agonistic situations, but rather appears to be the norm for processing in all social interactions, as has been demonstrated in other species including chicks and a range of vertebrates. In domestic horses, hemispheric specialization for sensory input appears not to be based on a designation of positive versus negative, but more on the perceived need to respond quickly and appropriately in any given situation.     

Sex task and processing strategy effects in hemispheric alpha asymmetries for the recall and recognition of arousal words: results from perceptual and motor tasks in males and females

Hemispheric alpha asymmetries of males and females were observed during perceptual and motor tasks requiring recall and recognition of words controlled for level of arousal (positive, negative, and neutral). Verbal reports of individual processing strategy were collected and analyzed relative to hemispheric alpha ratios. Results showed greater alpha suppression in the left relative to right hemisphere for recall as compared to recognition tasks and for word presentation when contrasted with motor conditions. High positive correlations were found between narrative report of processing strategy and hemispheric alpha data. A separate analysis revealed that seven subjects identified as highly analytic processors showed greater alpha suppression in the left relative to right hemisphere across tasks, conditions, and stimuli than did seven highly visual processors who, in contrast, demonstrated greater right hemispheric alpha suppression. Task difficulty and individual differences in processing style that modify cerebral laterality effects are discussed.     

The effect of auditory input on cerebral laterality

Cerebral laterality was examined for third-, fourth-, and fifth-grade deaf and hearing subjects. The experimental task involved the processing of word and picture stimuli presented singly to the right and left visual hemifields. The analyses indicated the deaf children were faster than the hearing children in overall processing efficiency, and that they performed differently in regard to hemispheric lateralization. The deaf children processed the stimuli more efficiently in the right hemisphere, while the hearing children demonstrated a left-hemisphere proficiency. This finding is discussed in terms of the hypothesis that cerebral lateralization is influenced by auditory processing.     

阿拉伯数字与汉字大写数字认知的大脑功能一侧化实验研究

用半视野速示术检查阿拉伯数字和汉字大写数字在三种任务加工条件（认读、简单相加、奇偶概念异同判断）下进行认知的大脑两半球机能活动一侧化效应。结果表明：以被试正确反应的百分数为指标,两类数字的三种任务加工均没有表现出明显的视野（大脑半球）优势。以被试正确反应的反应时为指标,两类数字的认读和简单相加也未出现视野（大脑半球）的明显差别,而两类数字的奇偶概念异同判断却出现了差异显著的右视野（左半球）优势。     

Recognition of affective and noncanonical linguistic facial expressions in hearing and deaf subjects

This study explores the use of two types of facial expressions, linguistic and affective, in a lateralized recognition accuracy test with hearing and deaf subjects. The linguistic expressions represent unfamiliar facial expression for the hearing subjects whereas they serve as meaningful linguistic emblems for deaf signers. Hearing subjects showed left visual field advantages for both types of signals while deaf subjects' visual field asymmetries were greatly influenced by the order of presentation. The results suggest that for hearing persons, the right hemisphere may predominate in the recognition of all forms of facial expression. For deaf signers, hemispheric specialization for the processing of facial signals may be influenced by the differences these signals serve in this population. The use of noncanonical facial signals in laterality paradigms is encouraged as it provides an additional avenue of exploration into the underlying determinants of hemispheric specialization for recognition of facial expression.     

Valence specific laterality effects in free viewing conditions: the role of expectancy and gender of image

Recent research has looked at whether the expectancy of an emotion can account for subsequent valence specific laterality effects of prosodic emotion, though no research has examined this effect for facial emotion. In the study here (n=58), we investigated this issue using two tasks; an emotional face perception task and a novel word task that involved categorising positive and negative words. In the face perception task a valence specific laterality effect was found for surprise (positive) and anger (negative) faces in the control but not expectancy condition. Interestingly, lateralisation differed for face gender, revealing a left hemisphere advantage for male faces and a right hemisphere advantage for female faces. In the word task, an affective priming effect was found, with higher accuracy when valence of picture prime and word target were congruent. Target words were also responded to faster when presented to the LVF versus RVF in the expectancy but not control condition. These findings suggest that expecting an emotion influences laterality processing but that this differs in terms of the perceptual/experience dimension of the task. Further, that hemispheric processing of emotional expressions appear to differ in the gender of the image.