Focal changes in cerebral blood flow produced by a test of right-left discrimination

A sensorimotor control activation task was used to isolate the focal cerebral blood flow changes resulting from the visual and cognitive processing of a right-left discrimination task. Eleven normal right-handed males participated. The sensorimotor control task produced significant bilateral increases in flow in most cortical channels. Significant bilateral parieto-occipital activation was found for the right-left discrimination task over and above the flow changes produced by the sensorimotor control task. The left occipital flow increase resulting from the right-left discrimination task was found to be negatively related to task performance. An inverse relationship was also found between WAIS Performance IQ and the blood flow change in the left parietal channel. These results suggest areas for further testing concerning potential individual differences in cognitive processing during the performance of a right-left discrimination task.     

Children's preference for body or external object on a task requiring transposition and discrimination of right-left relations.

First graders were tested on trials requiring the discrimination and transposition of right-left relations, with stimuli similar to (dolls) or different from (toy planes) the body. On an initial series of training and practice trials children were allowed to respond to body and external objects as referents providing right-left cues. A later series of test trials was then used to determine the referent the child actually preferred. Results of the test trials indicated that 24 children tested with the dolls preferred the body as referent, while 24 children tested with planes preferred the external referent. The results were interpreted as suggesting two alternative systems through which children develop an understanding of right-left relations and possibly other concepts as well.     

Right-left confusion in the adult: A verbal labeling effect

Directional judgments are typically slower when relative location is described by the words “east” and “west” or “right” and “left” than when described by the words “north” and “south” or “up” and “down.” A series of experiments are reported that disentangle verbal from perceptual encoding explanations for right-left difficulty. Overall, our results support a verbal encoding explanation for right-left confusion in the adult. Experiments 1-3 demonstrate that in a response-differentiation task, it is response to the labels “north,” “east,” “south,” and “west” that is responsible for right-left confusion. In addition, Experiments 4-6 demonstrate that right-left difficulty in a mirror image discrimination task is contingent on the use of directional labels. (The data also suggest that it may be more difficult to deal with “up,” “down,” “left,” and “right” than with “north,” “south,” “east,” and “west”) The data are interpreted as inconsistent with a bilateral symmetry explanation for right-left confusion.     

A code-theoretical note on object handedness

This study is a theoretical exercise dealing with discrimination between images and mirror-images. It focuses on the way codes of shapes represent their handedness. We compare two code systems with different reference frames. These frames determine the specific sensitivity of each system. One system uses an asymmetric reference frame. It is called the H-system and was inspired by an idea of Corballis (1988 Psychological Review 95 115-123). The other system, being our proposal, uses a symmetric reference frame and we have named it the M-system. We demonstrate the following. A code of the H-system provides a cue for the handedness of a shape, but not for rotation, i.e. no cue for the appropriate kind of code rotation which should be tested in case images and mirror-images are discerned by mental rotation. The M-system is the converse in both respects. A code of this system does not provide a handedness cue but, instead, a rotation cue. Thus, for handedness discrimination, the H-system neither needs nor guides mental rotation, whereas the M-system does both. This M-system generates object-centred structural codes enriched with viewpoint information. Various visual experiments reported in the literature favour the M-system over the H-system, implying that perception does not make use of an asymmetric but of a symmetric reference frame.     

Right-nostril advantage for discrimination of odors

Olfactory discrimination was tested with eight pairs of odors presented to each nostril of each subject. Ninety-nine subjects were tested; there were equal numbers of left- and right-handers, as well as both sexes. Detection thresholds for phenylethyl alcohol were measured separately in each nostril using a forced-choice staircase procedure. In addition, a verbal dichotic listening test known to be sensitive to language lateralization was administered. Results indicated that discrimination performance was significantly better when the stimuli were presented to the right nostril than when they were presented to the left, but no differences between the nostrils in detection thresholds were found. The right-nostril advantage did not vary as a function of sex or handedness, and did not bear any relation to language lateralization as measured via dichotic listening. The asymmetry for olfactory discrimination replicates an earlier study and is interpreted in terms of a possible specialization of function within the right cerebral hemisphere.     

Neuropsychological evaluation and rehabilitation in mental retardation

A discussion is provided of issues related to neuropsychological assessment of mentally retarded individuals. The first matters considered are test selection in relation to mental age and diagnostic issues. Various methods of evaluation are then considered, including assessment of orientation to sound and visual stimuli, language comprehension, right-left discrimination, and abstract and symbolic processes. Methods are also reviewed for detailed evaluation of language functions including testing for aphasia and for academic skills such as reading, writing, and calculation, as well as for spatial and constructional abilities. A section is devoted to use of neuropsychological assessment for rehabilitation of various disorders such as congenital aphasia, alexia, and spatial-constructional disorders.     

The role of verbal labels in the judgment of orientation and location

In an earlier study it was found that judgments of right-left orientations and locations were more difficult than judgments of up-down only when spatial words were used in the tasks. Experiments are reported in which pictures of many objects were presented to eliminate the possibility that subjects in previous studies had used strategies specific to single-stimulus tasks. In experiment 1, right-left orientations were judged more slowly than up-down orientations both when the spatial words were used and when arbitrary letters replaced the spatial words. In experiment 2, judgments of the right-left locations of pictures took longer than judgments of their up-down locations only when spatial words were used in the task; the right-left difficulty was eliminated when arbitrary letters replaced the words. The differential effect of words and letters in location judgments seems to be due to the different coding strategies adopted by subjects under the two conditions. It is concluded that a right-left difficulty does not depend on the use of spatial terms: word and letter conditions yield different results only when the task permits different judgments to be made under the two conditions.     

Preparation for horizontal or vertical dimensions affects the right-left prevalence effect

When stimulus and response simultaneously vary in both horizontal and vertical dimensions, the stimulus-response compatibility effect is often larger for the horizontal dimension. We investigated the role of preparation for each dimension in this right-left prevalence. In Experiment 1, tasks based on horizontal and vertical dimensions were mixed in random order, and the relevant dimension in each trial was cued with a variable cue-target stimulus onset asynchrony (SOA). A right-left prevalence effect was observed only when participants prepared for the upcoming task. Experiment 2 replicated the absence of the prevalence effect for the simultaneous presentation of cue and target using a fixed SOA of 0 msec. In Experiment 3, the right-left prevalence emerged with a 0-msec SOA when participants prepared for e achdimension basedon its frequency. These resultssuggest that participants' internal set can be greater for the horizontal dimension, leading to the right-left prevalence effect.     

Spatial and non-spatial visual discrimination learning in zebrafish (Danio rerio)

Zebrafish (Danio rerio) provide an excellent model for assessment of molecular processes of neurodevelopment. To determine the functional importance of molecular events during neurodevelopment, we have developed methods for assessing learning in zebrafish in a three-chambered fish tank. In the first study, simple escape response was assessed. Zebrafish tested with a moving net learned to escape to another chamber more rapidly over the six sessions of training than the fish with the still net which did not learn. Upon reversal of the contingencies, the fish switched to the inactive net rapidly learned to suppress the escape response and fish formerly in the inactive net condition learned to avoid the moving net. In the second study, spatial discrimination learning was assessed. Zebrafish were trained on a right-left position discrimination to avoid the active net. Zebrafish showed significant improvement in escape responses over six sessions of training with three trials per session. In the third study, red-blue non-spatial discrimination learning was assessed. There was a significant improvement over the first six training sessions. With the reversal of contingencies, there was a significant decline of performance. With continued training, the fish again significantly improved avoidance. These studies found an effective motivational stimulus and procedure for studying escape behavior in zebrafish; a procedure whereby zebrafish would learn both spatial and non-spatial discrimination. These methods are being developed to help determine the functional importance of molecular events during zebrafish neurodevelopment. Accepted after revision: 20 August 2001 Electronic Publication     

Is saccade-induced retrieval enhancement a potential means of improving eyewitness evidence?

Saccade-induced retrieval enhancement (SIRE) is the effect whereby making bilateral saccades enhances the subsequent retrieval of memories. Two experiments explored SIRE's potential to improve eyewitness evidence. Participants viewed slideshows depicting crimes, and received contradictory and additive misinformation about event details either once (Experiment 1) or three times (Experiment 2). Participants then performed saccades or a fixation control task before being tested on their memory for the slideshows and making confidence judgements. Saccades increased discrimination between seen and unseen event details regardless of whether or what type of misinformation was presented. Because prior studies indicated that SIRE might be more robust for individuals who are strongly right-handed versus not, we examined SIRE as a function of handedness and found that saccades improved memory for event details regardless of participants' handedness. However, participants who were not strongly right-handed had fewer false memories than participants who were strongly right-handed, extending previous findings of superior memory among individuals who are not strongly right-handed. Saccades also increased confidence in true memories (Experiment 1) and decreased confidence in false memories (Experiment 2). The results support SIRE's potential to improve eyewitness evidence.     

Splitting visual space with attention

If a right-left spatial compatibility effect is observed, it can be maintained that the space has been segmented into right- and left-side parts. The present study aimed at showing a spatial compatibility effect (and, by implication, a right-left subdivision of space) solely attributable to the orienting of attention. Five groups of 8 normal subjects were required to give right-left discriminative responses to stimuli presented within one of six empty boxes arranged in a horizontal row. Reaction times and errors were recorded. The first two experiments showed that a right-left grouping of the boxes occurred regardless of whether subjects' fixation was kept at the intermediate position (Experiment 1) or at one extremity (Experiment 2) of the row. In Experiments 3 and 4, subjects' attention was not aligned with a fixed position but was moved, through peripheral cues, from trial to trial and positioned between different pairs of adjacent boxes. The results showed that the display was again subdivided into two regions and that the reference point for the right-left subdivision was the focus of attention. In Experiment 5, eye position was instrumentally monitored, and subjects' attention was directed by central cues. The results confirmed that the focusing of attention leads to a right-left partitioning of space, with the boundary at the locus of focal attention. In conclusion, we demonstrated, by employing a right-left spatial compatibility paradigm, that directing attention to a position in space brings about a right-left perceptual organization that predominates over that provided by the other egocentric reference axes.     

Right-left prevalence with task-irrelevant spatial codes

The present work investigated the right-left prevalence effect caused by the automatic activation of horizontal and vertical spatial codes in a task (Simon task) in which spatial information is task-irrelevant. Experiment 1 showed a horizontal Simon effect and a vertical Simon effect with a two-dimensional stimulus-response set. In Experiments 2 and 3, the right-left prevalence was obtained in two-dimensional Simon tasks with two contralateral effectors and four effectors respectively. Experiment 4 showed that horizontal coding is based on multiple spatial codes, whereas only one spatial code was formed for vertical coding. On the whole, these results support the notion that the right-left prevalence effect is a general phenomenon affecting spatial coding, and suggest that the horizontal dimension is prevalent because it is based on multiple spatial codes.     

Effects of precuing horizontal and vertical dimensions on right—left prevalence

When stimuli and responses can be coded along horizontal and vertical dimensions simultaneously, a right-left prevalence effect is often obtained for which the advantage for a compatible mapping is larger on the horizontal dimension than on the vertical dimension. The present study investigated the role of preparatory processes in this right-left prevalence effect using a method in which the relevant dimension was cued at short and long intervals prior to presentation of the target stimulus. In three experiments, the right-left prevalence effect did not vary significantly in magnitude as a function of cue-target interval, suggesting that the effect is due primarily to relative salience of the horizontal and vertical codes, as determined by the task structure, and not to a greater ease of attending to the horizontal dimension.     

Mirror writing in pre-school children: a pilot study

Mirror writing refers to the production of individual letters, whole words or sentences in reverse direction. Unintentional mirror writing has been observed in young children learning to write and interpreted as the manifestation of different cognitive impairments. We report on mirror writing instances in a sample of 108 pre-school children. Results showed MW to be age-related but independent from handedness and left-right discrimination abilities. We propose an account of mirror writing as reflecting dissociation between acquired motor programmes for letter shape composition and unspecified spatial direction of hand movements. Before learning to write, the child’s directional cognitive system is assumed to be dichotomous, thus inducing the production of randomly oriented asymmetrical letters.     

Unimanual and bimanual tasks and the assessment of handedness in toddlers

The goal of this study was to examine the relations between three different measures of handedness: unimanual reaching, bimanual manipulation and unimanual manipulation. The appropriateness of the task chosen to evaluate handedness was also explored by contrasting different bimanual manipulation tasks for the more or less differentiated (passive\active) roles assigned to each hand. Forty children, between 18 and 36 months of age, were tested in the three conditions. The results show that the degree of bimanual handedness is greater on the bimanual tasks with a strong role differentiation than on the tasks with less differentiation. Bimanual tasks with a strong role differentiation elicited more right‐handedness than unimanual reaching. Among the children who showed handedness in reaching, the correlation between unimanual and bimanual handedness was high, especially for right‐handers. For some tasks, bimanual handedness appeared at the earliest age studied here (18 months), and there was little relationship between bimanual handedness and bimanual skill. In contrast with unimanual reaching, there was no age‐related change in the degree of handedness for either bimanual or unimanual manipulation. There was a bias toward the use of the right hand for unimanual manipulation. It was concluded that grasping is not the best task to employ to look for robust evidence of handedness, and that bimanual tasks offer a better way to estimate handedness in children, as long as the tasks are carefully chosen.     

Comparative and familial analysis of handedness in great apes

Historically, population-level handedness has been considered a hallmark of human evolution. Whether nonhuman primates exhibit population-level handedness remains a topic of considerable debate. This paper summarizes published data on handedness in great apes. Comparative analysis indicated that chimpanzees and bonobos show population-level right handedness, whereas gorillas and orangutans do not. All ape species showed evidence of population-level handedness when considering specific tasks. Familial analyses in chimpanzees indicated that offspring and maternal (but not paternal) handedness was significantly positively correlated, but this finding was contingent upon the classification criteria used to evaluate hand preference. Overall, the proportion of right handedness is lower in great apes compared with humans, and various methodological and theoretical explanations for this discrepancy are discussed.     

Determinants of right-left and top-bottom prevalence for two-dimensional spatial compatibility

When stimulus and response sets vary along horizontal and vertical dimensions, the horizontal dimension is more dominant than the vertical one, an effect called right-left prevalence. Three accounts have been proposed that attribute the effect to a reduced ability to code vertical locations when horizontal codes are also present, the use of right-left effectors, or a difference in salience of the 2 dimensions. The accounts differ in terms of whether the ability to code and process the 2 dimensions is of limited capacity and whether the prevalence effect is a consequence of the effectors used for responding. The authors report 4 experiments that evaluated these issues. Results indicate that use of right-left effectors is important to the right-left prevalence effect because it increases the salience of the horizontal dimension. However, a top-bottom prevalence effect can be obtained if the vertical dimension is made more salient.     

The Specificity of Motor Skill and Manual Asymmetry: A Review of the Evidence and Its Implications

Results of investigations on gaining control of limb movements are reviewed, and their contribution to understanding the development of manual asymmetries is discussed in relation to the discrimination and programming of appropriate neuromuscular resources. An examination of the relevant evidence on number and types of manual asymmetries recorded provides strong grounds for concluding that where asymmetries occur, they simply represent a further example of the well-documented activity-specific nature of motor skills and of the extremely lengthy periods of learning or experience needed for their acquisition and perfection. This specificity of motor skill and manual asymmetry also readily accounts for most of the discrepancy usually reported between assessments of hand preference and performance differences between hands, because these alternative measures of handedness have rarely employed the same range or variety of tasks.     

Vertical versus horizontal spatial compatibility: Right-left prevalence with bimanual responses

For two-choice tasks in which stimulus and response locations vary along horizontal and vertical dimensions, the spatial compatibility effect is often stronger on the horizontal than vertical dimension. Umiltà and Nicoletti [(1990) Spatial stimulus-response compatibility (pp. 89–116). Amsterdam: North-Holland] attributed this right-left prevalence effect to an inability to code vertical location when horizontal codes are present simultaneously. Hommel [(1996) Perception & Psychophysics, 43, 102–110] suggested instead that it reflects a voluntary strategy. This study reports four experiments that examine this issue. Experiment 1 was a conceptual replication of Hommel's Experiment 1, with responses made on a numeric keypad and subjects instructed in terms of the vertical or horizontal dimension. The results replicated Hommel's findings that showed a right-left advantage with horizontal instructions; however, with vertical instructions, we found a benefit of vertical compatibility alone that he did not. This benefit for vertical compatibility alone was eliminated in Experiment 2 using a varied practice schedule similar to that used by Hommel. Experiment 3 showed right-left prevalence and a benefit of vertical compatibility alone, even with varied practice and vertical instructions, when subjects responded on perpendicularly arranged handgrips. These benefits were eliminated in Experiment 4 using Hommel's method of urging subjects to respond only in terms of the instructed dimension. With bimanual responses, right-left prevalence is a robust phenomenon that is evident when comparing across vertical and horizontal instructions and, when the right-left distinction is relatively salient, within the vertical instructions condition alone. Received: 4 February 2000 / Accepted: 5 May 2000     

On the evolution of handedness: a speculative analysis of Darwin's views and a review of early studies of handedness in "the nearest allies of man"

Scientists today who seek clues into the evolutionary origins of human handedness make extensive use of evidence from comparative studies, that is, studies that ask whether handedness occurs in other species, especially apes and monkeys, as the Darwinian principle of continuity would seem to imply, or whether it is uniquely human. Early investigations had the same goal and drew on much the same kind of evidence. In this article, I review studies of animal handedness in the period before 1859, when Darwin published On the Origin of Species, and afterward, through the 1st decade of the 20th century. Inasmuch as Darwin's published writings contain hardly any statements about handedness and none at all about its evolution and continuity across species, I also speculate about what Darwin himself might have said on the subject. To do this, I draw on his statements on related matters, such as the form and structure of the hand and the transition from a quadrupedal to bipedal stance, on other writers' reports and opinions about handedness with which he was familiar or likely to have been familiar, and finally, on clues from his own and only statement about animal handedness in an unpublished letter. I conclude by asking whether and how early investigators, lacking any statement by Darwin on the evolution of handedness, invoked his theory of evolution and his views on related matters in the interpretation of their findings.