Journal of Abnormal Psychology 2008, Vol. 117, No.

4, 799 – 811

Copyright 2008 by the American Psychological Association 0021-843X/08/$12.00 DOI: 10.1037/a0013500

Grasping Behavior in Schizophrenia Suggests Selective Impairment in the Dorsal Visual Pathway
Jelena P. King and Bruce K. Christensen
St. Joseph’s Healthcare Hamilton and McMaster University

David A. Westwood
Dalhousie University

This study frames anomalous functional brain organization in schizophrenia (SCZ) within an evolutionary model of brain development, the dual trends theory (DTT). The DTT argues that neural architecture develops along 2 separate pathways: the dorsal archicortical trend and the ventral paleocortical trend. The DTT dovetails with visual system organization, which is also composed of 2 independent pathways: a dorsal stream dedicated to visuomotor action and a ventral stream dedicated to perceptual processing. The present study examined the integrity of these pathways using a size-contrast visual illusion. Prior research has shown that, normally, perceptual estimations of object size are susceptible to visual illusions, whereas goal-directed actions are resistant. The authors hypothesized that, unlike control participants, SCZ patients’ actions would be susceptible to the illusion, reflecting dorsal stream dysfunction. Here, 42 SCZ patients and 42 healthy controls grasped and estimated the size of target blocks in control and illusion conditions. During estimation, both groups were equally perturbed by the illusion; however, grasping movements of patients alone were influenced by the illusion. These results suggest disrupted dorsal brain circuitry in SCZ but relatively intact ventral circuitry. Keywords: schizophrenia, visual illusion, action and perception, dual cytoarchitectonic trends, evolution

Schizophrenia (SCZ) is associated with pervasive neuropathology (Akbarian et al., 1996; Gur et al., 1998). These brain abnormalities are considered to be developmental in origin and to mediate the behavioral and cognitive consequences of this illness. Attempts to understand the functional consequences of SCZrelated neuropathology, however, have relied on traditional neuropsychological models of acquired localized brain damage and largely ignored models of brain development and evolution. There are several problems with this approach. First, patients with localized lesions differ substantially from patients with SCZ on measures of symptomatology and cognition (e.g., Pantelis et al., 1999). Second, neuropathological studies of SCZ have failed to reveal the macroscopic lesions that typify acquired brain pathology (Harrison, 1999). Third, the lesions associated with acquired brain damage are focal, whereas those associated with SCZ are distributed across larger neural systems. Fourth, adult animals with acquired

Jelena P. King and Bruce K. Christensen, St. Joseph’s Healthcare Hamilton, Hamilton, Ontario, Canada, and Department of Psychiatry and Behavioural Neurosciences, McMaster University, Hamilton, Ontario, Canada; David A. Westwood, School of Health and Human Performance, Dalhousie University, Halifax, Nova Scotia, Canada. This research, in part, was conducted in partial fulfillment of requirements for the doctoral degree in the Department of Psychology at the University of Waterloo, Waterloo, Ontario, Canada, for Jelena P. King and was funded by an operating grant to Bruce K. Christensen from the Canadian Psychiatric Research Foundation. Results of the current study were presented at the 17th annual conference of the Society for Research in Psychopathology, September 2002, San Francisco, CA. We gratefully acknowledge the programming assistance of Yee Hong Chia. Correspondence concerning this article should be addressed to Bruce K. Christensen, St. Joseph’s Healthcare Hamilton, 100 West 5th Street, Hamilton, Ontario, Canada L8N 3K7. E-mail: Bruce.Christensen@mcmaster.ca 799

brain lesions do not exhibit behavior that is prototypically schizophrenic (Lipska, Jaskiw, & Weinberger, 1993), and neurodevelopmental lesions affecting frontolimbic connectivity are superior to acquired lesion models in their ability to produce SCZ-like syndromes and biochemical responses (Joel, Weiner, & Feldon, 1997). Collectively, these problems highlight the need for a valid neurodevelopmental framework within which the functional consequences of SCZ-related neuropathology can be better integrated and understood across broader neural systems. The current research uses the dual trends theory (DTT; Pandya & Barnes, 1987; Sanides, 1969) as a neurobiological framework for understanding the range of cognitive and behavioral impairment associated with SCZ. The DTT argues that neural architecture develops along two separate pathways: the archicortical trend and the paleocortical trend. Cytoarchitectonic methods have revealed that cross-species cortical differentiation emanates from the hippocampus (i.e., archicortex) and the pyriform cortex (i.e., paleocortex) and progresses through distinct cortical regions (Pandya & Barnes, 1987; Pandya & Yeterian, 1985). Anatomically, new regions are organized in relationship to these two primitive cortical areas and occupy different cortical territories. Those regions deriving from the archicortex are stationed largely in dorsal–medial aspects of the primate brain, whereas those regions originating from the paleocortex occupy ventral–lateral regions of the primate brain. Consistent with this structural duality, researchers have suggested that the archicortical and paleocortical trends subserve two different functions. The archicortical system is responsible for the stable execution of task-relevant, volitional control functions and subserved by cortical networks preferentially responsive to redundancy in the environment. In contrast, the ventral paleocortical system subserves arousal functions in response to novel environmental contingencies (Goldberg, 1987; Pribram & McGuinness,

In the PAM. Hence. 2006) have proposed that SCZ may be the result of selective aberrations in the development of these pathways. SCZ patients demonstrate deficits on cognitive tasks that require volitional. Selemon. are affected by visual illusions. 2001). The ventral or parvocellular pathway projects primarily from parvocellular layers of V1 to inferotemporal regions and mediates perceptual processing necessary for object identification.. Esiri. 2000. automatic. In this regard. or intuitive behavior is comparatively intact (Danion. 1990). 1990). Goodale. whereas the archicortical trend terminates in areas of the visual cortex responsible for peripheral vision (Pandya & Yeterian. visual information regarding object identity and significance forms a key aspect of the arousal function ascribed to ventral brain regions. whereby emotional saliency is overassigned to otherwise innocuous stimuli (Kapur. On a functional level. 1991). Ventral processing for perception is thought to use relative rather than absolute metrics because relative metrics are more useful for representing stable features of the environment. 1988).e.. DeSouza. dorsal visual information regarding object spatial location is used to guide manipulatory object-directed actions (Pandya & Yeterian. and goal-directed behavior (Bilder et al. Giaccio. which lie at the core of perceptual experience. a defect in the archicortical trend is consistent with many of the identified abnormalities of brain structure. low spatial resolution. the paleocortical trend appears to terminate in areas of the visual cortex subserving central vision.e. 2000. negative symptoms) can be conceptualized as deficits in willed action and willed intention (Frith & Done. the dorsal stream is often referred to as the visionfor-action system whereas the ventral stream is referred to as the vision-for-perception system. 2006).e. That is. & Goodale. 2003). & Carey. identification. many of the clinical symptoms in SCZ (e. high spatial resolution.. & Vermaat. or explicit visual perception. In contrast. Livingston & Hubel. 1982).800 KING. In addition. Finally. but grasping under identical illusion conditions remained consistently scaled to the true size of the target disk. preferential archicortical connectivity relating to peripheral vision— crucial for accurate spatial perception—to dorsal trunk and limb regions of somatosensory association cortex and dorsal and medial prefrontal regions is involved in the integration of visuospatial and motivational processes.e. a key implication of the PAM is that vision and visually guided behavior can be used as a window into the relative integrity of dorsal and ventral brain circuitry. Giaccio. Kleinman. Milner and Goodale (1995) have proposed an alternative interpretation of the functions of the dorsal and ventral visual streams that differs somewhat from those outlined by the dual-processing model (e.g. Milner. have a pronounced effect . color discriminating.. whereas performance should be spared in tasks that depend primarily on ventral stream processing (i. The connectivity and function outlined by the DTT dovetails with the dual-processing model of primate vision. Ventral stream visual processing permits the formation of perceptual representations required for cognitive processes like recognition. Kauffmann-Muller. Psychophysical studies using pictorial visual illusions provide support for the relative independence of vision for perception and vision for action in normal human visual behavior. which appear to disproportionately involve dorsal brain regions (i.e. Mrzljak.. The two distinct architectonic trends. Haffenden & Goodale. 2002. the archicortical trend progresses ventrolaterally whereas the paleocortical trend progresses ventromedially. vision for action) should be disrupted in individuals with disorders—such as SCZ—that selectively target dorsal brain regions. and neck regions of somatosensory association cortex and ventrolateral prefrontal regions is closely tied to the synthesis of object– emotion relationships in a behavioral context (Pandya & Yeterian. AND WESTWOOD 1975). Rajkowska. which are important for identifying objects and selecting appropriate responses. The dorsal or magnocellular pathway projects primarily from magnocellular layers of primary visual cortex (V1) to occipitoparietal regions and is devoted to processing spatial information and movement control. Meulemans. 1987). Thus. hippocampus. as explicated by the DTT. 2001). whereas performance on tasks requiring responsive.. The evidence to support these observations is briefly summarized below but reviewed in greater detail elsewhere (Christensen & Bilder. 2003).. The mag- nocellular pathway is subserved by cells with physiological properties specifically tuned to process motion and spatial information (i. 1995) but ventral sparing (i. For the purposes of the present investigation. illusions. Jakobson. Specifically. 2004). For example. color blind. Haffenden and Goodale (1998) used the Ebbinghaus illusion to demonstrate that healthy participants’ manual estimates of circular disks were impacted by the illusion. First. ventrolateral prefrontal cortex. & Crow. ventral sparing in the absence of contextual regulation from the dorsal system may explain the development and/or maintenance of positive symptoms.. Selemon. Gur et al. 1995.. amygdala. have also been traced within the cortical visual system. effortful. vision for perception). and memory. 2000).e. 1983). which also distinguishes between two independent cortical visual pathways (Ungerleider & Mishkin. In the context of the PAM. head. whose effects arise from obligatory comparisons of the target object to other objects in the surrounding context. investigators (Christensen & Bilder. CHRISTENSEN. The crux of Milner and Goodale’s perception–action model (PAM) is that the dorsal and ventral streams do not differ in the types of visual information they process but rather in how that information is linked to the control of behavior. although both trends originate in the temporal lobe (Galaburda & Pandya. slow responding. Livingston & Hubel. More recently. Cells in the ventral parvocellular pathway exhibit properties selective for high-resolution form perception necessary for object identification (i. such results are interpreted in terms of differences in the underlying visual processes that take place in the ventral perception and dorsal action streams. preferential paleocortical connectivity between inferotemporal regions subserving central vision— crucial for accurate object perception—to ventral face. This pattern of visual connectivity is associated with two key functional behavioral implications. whereas goal-directed actions like grasping are immune to these illusions (Aglioti. Chance. In contrast. 1987). & Goldman-Rakic. Arguably the most important contribution of the PAM is acknowledgment that visual control of action is the primary function of the dorsal visual stream. More specifically. medial– dorsal frontal cortex.. Conversely. & Goldman-Rakic.g. 1990). which are key functional roles of the dorsal trend. fast responding. performance on tasks that depend on dorsal stream processing (i. Herman. 2000. visual transformations in the dorsal stream lead to the programming and control of skilled actions directed toward objects (reviewed in Goodale & Westwood. 1998. Studies of this type have consistently demonstrated that perceptual judgments of object size. reviewed in Carey.

Elliot. claims about specific dorsal stream impairment are somewhat premature. Spitzer.28) 32. with the majority working in labor– clerical fields or pursuing postsecondary education. 2001).e. values are means (and standard deviations). Inc. Collectively.76 (11. As reported below. Quebec. Drawing on the PAM.36) 14. Thus. the visual control of action must be concerned with the absolute rather than relative metrics of objects because actions could not be accurate or efficient without such precise information. The abnormalities in movement execution appear to involve fine motor control during reaching. visual transformations in the dorsal stream must ignore. like conscious perceptual experience. In contrast.. 1996). tricyclic antidepressant. Elliott. SCZ patients can use redundant object information and cues to facilitate movement in much the same way as healthy controls (Carnahan et al.888 Note. Aguilar. may be rendered susceptible to the influence of visual illusions. such as reaction time (RT).. All participants denied a history of neurological illness.e.e... All clinical interview (i.. However. According to the PAM. SCZ patients can integrate intrinsic and extrinsic object information into their actions (Carnahan et al. to some extent. loss of consciousness for greater than 20 min from a blow to the head). Oldfield. Montreal. & Velamoor. SCZ patients do not show gross motor abnormalities. this pattern of findings provides tentative support for the idea that SCZ is associated with dysfunction in dorsal circuitry with relatively preserved ventral processing.g. the results of this experiment support these predictions. brain injury (i. or history of medical conditions that could affect cognitive performance (e. current or past Diagnostic and Statistical Manual of Mental Disorders (4th ed. this hypothesis leads to the prediction that SCZ patients should show selective deficits in the visual control of action in the face of spared visual perception. myocardial infarction. Although deficits in motor functioning are commonly reported in SCZ (e. 26 men) with SCZ or schizoaffective disorder and 42 healthy controls (23 women.48) 14. Elliott.g. albeit less so. More specifically. reviewed in Heinrichs & Zakzanis.30 mg (SD 271. By and large. The majority of control (n 40) and patient (n 38) participants were right-handed (as assessed using the Edinburgh Handedness Inventory. last 6 months) DSM–IV substance abuse disorder. 1994) and. 1998). in movement execution. with one of these qualifying as neuroleptic naive. & Velamoor..36) 2. actions like grasping are not sensitive to the same pictorial illusions that have such a robust effect on conscious perceptual experience. with the majority of patients (i. inTable 1 Demographic Characteristics of Patient and Healthy Control Samples Variable Age (years) Education (years) Controls (n 42) Patients (n 42) t(82) p 38. Only a few studies have examined visuomotor functioning in SCZ using more sensitive kinematic measures. because these experiments did not specifically dissociate conscious visual perception and visual control of action. . 1995).88 (9. but rather more subtle difficulties linking vision to the control of action. reflecting intact ventral stream processing. Finally.36 (2. or sedative agents at the time of research participation. these findings suggest a somewhat specific deficit in the ability of SCZ patients to properly integrate visual object information into the control of their actions. it is expected that grasping behavior. Two patients were unmedicated.. The mean chlorpromazine equivalent for the sample was 371. American Psychiatric Association. Gibbon.141 . Patients were significantly younger than controls but matched on education level.15).e. 1997) and have been shown to properly scale the size of their grasping hand to the size of the target object (Carnahan. Canada). learning disability. All participants had normal or corrected 20/20 vision confirmed by acuity testing (Snellen Test. These studies have revealed that patients with SCZ have deficits at the level of movement planning (Carnahan. Chua. Unless otherwise noted. and all participants used their preferred hand to complete the experimental tasks. we sought to test the hypothesis of disproportionate dorsal trend disruption in SCZ by examining the relative effects of a visual illusion on perceptual experience and control of action. 1995). see Table 1 for demographic characteristics of the sample). All patient participants were taking atypical antipsychotic medications.016 0. Velamoor.451 . DSM–IV. In the current study. reflecting disruption in dorsal stream processing... Because of this target-centered style of processing. Patient diagnoses were confirmed by the Structured Clinical Interview for DSM–IV–TR Axis I Disorders (SCID I. thyroid disease. and (b) only SCZ patients will demonstrate an effect of the illusion on grasping behavior. (a) SCZ patients and healthy controls will both demonstrate the typical effect of the size-contrast illusion on perceptual judgments of object size. Method Participants Participants were 42 patients (16 women. First. those objects in the visual scene that are not the target for intended action. 1994) substance dependence or current (i. & Norman. possibly reflecting a reduced ability to integrate visual feedback into the control of ongoing movements (Carnahan.. 1971). 19 men). No patients were taking anticholinergic.GRASPING BEHAVIOR IN SCHIZOPHRENIA 801 on perceptual experience. However. Participants in both groups were generally middle aged and high school educated. 1997 Carnahan. & Carnahan. & Williams.43 (2. Canada by posted advertisements and referrals from hospital staff. Control participants were also excluded if they reported having a firstdegree relative with a SCZ-spectrum illness. SCZ patients were recruited from an academic psychiatric hospital in Toronto. The purpose of the present study was to investigate the relative integrity of the dorsal and ventral visual pathways in SCZ using a visual illusion paradigm to test the hypothesis that SCZ is associated with selective dysfunction of neural structures in the dorsal trend. these findings have been based on conventional methods. Consequently. in individuals with impairments in dorsal stream functioning but spared ventral stream functioning. Malla. 86%) receiving only one neuroleptic. AMG Medical. Ontario. Control participants were recruited from the community using fliers posted in the neighborhood surrounding the psychiatric hospital and local newspaper. Materials Clinical Scales and Cognitive Instruments A number of clinical and cognitive instruments were administered to characterize the sample.

Dep Depression Scale.91) Note. & Opler. Gen General Psychopathology Scale T score. Although differences between groups in depressive symptomatology and alcohol and drug use were anticipated (Baynes et al. Par Paranoid Belligerence Scale T score.001 . but no more so than typical SCZ patient Table 2 Clinical and Neuropsychological Characteristics of Patient and Healthy Control Samples Variable Neuropsychological Estimated FSIQ WRAT–III Reading HVLT-R Clinical PAI Dep PAI Alc PAI Drg PAI PIM PAI NIM PANSS Pos PANSS Neg PANSS Com PANSS Gen PANSS An PANSS Ac PANSS Th PANSS Par PANSS De Controls (n 42) Patients (n 42) t(82) t(82) t(70.574a 5. Fiszbein.60) 45.93) 109.53 (5.54) 103.33) 45.41 (10.045a 4. PIM Positive Impression Management Scale. euthymic. symptom. Morey. and Drug Problems scales. 1989). Clinical and Neuropsychological Characteristics of Sample Patients and controls did not differ across neuropsychological variables.34) 38.77) 104.434 .48 (12. Cognitive instruments included the Matrix Reasoning and Information subtests of the Wechsler Adult Intelligence Scale—Third Edition (Wechsler. Ac Activation Scale T score.108) t(64.290a p . HVLT-R). Kay.85) 53. PAI Personality Assessment Inventory. with 40 mm separating the two objects. Alcohol Problems.49) 36. 1993) Reading subtest.. Morey. respectively. Drg Drug Problems Scale.31 (8. each consisting of a target block and a single flanker block. 2000).802 KING. Clinically insignificant movement abnormalities and akathesia were observed for patients (as determined by the AIMS and Barnes Akathesia Rating Scale).356) t 1. and Positive Impression Management (PIM) scales from the Personality Assessment Inventory (PAI. which included practice video and live supervised ratings.25) 56. with the flanker occupying the opposite location. statistically controlling for these variables in a covariate analysis did not alter any of the experimental results. De Depression Scale T score.47) 39. Brandt & Benedict.004 . 1991.001 .133 . Clinical scales included the Positive and Negative Syndrome Scale (PANSS.03) 41. Depression. Sattler.69 (12.35 (5. Guy. the Abnormal Involuntary Movement Scale (AIMS. Clinically.001 113. and 80 for Depression.91) 40. Th Thought Disturbance Scale T score. NIM Negative Impression Management Scale.78 (6. flanked on either the left or right side (see Figure 1) by a smaller (20 mm 20 mm for the 40-mm target and 40 mm 40 mm for the 60-mm target) or a larger (60 mm 60 mm for the 40-mm target and 80 mm 80 mm for the 60-mm target) flanker. see Table 2 for clinical and neuropsychological characteristics of the sample). Additionally. .50 (11. values are means (and standard deviations).g. groups (as determined from PANSS scores). and the Depression. Two target sizes were used (40 mm 40 mm and 60 mm 60 mm). and the Wide Range Achievement Test—Third Edition (WRAT–III. PANSS Positive and Negative Syndrome Scale. including the NIM.50 (6. CHRISTENSEN.74 (4. FSIQ Estimated full scale IQ from the Matrix Reasoning and Information subtest of the Wechsler Adult Intelligence Scale—Third Edition (validity of . Targets were presented to the left or right of a central location on an equal number of trials. the Hopkins Verbal Learning Test–Revised (HVLT-R. HVLT-R Hopkin’s Verbal Learning Test–Revised total T score.80) 48. 1991).517 0.77) 53. WRAT–III Reading Wide Range Achievement Test—Third Edition Reading subtest scaled score.48 (6.571 3.10 (11. 1976).51) t(77) t(59. 1987).70 (4. and mildly psychotic. a Degrees of freedom have been adjusted because of heterogeneity of variance.79) 35. Drug Problems. 1997). The target object was marked with a small red dot.. 2001).52) 47.69) 50.468) t(49. AND WESTWOOD cluding SCID I).93) 57.26 (7. Negative Impression Management (NIM). Com Composite Scale T score.26) 40. Individuals were randomly assigned to each rater.69 (9.867 for the dyad selected.47) 59.81 (11. 84.600a 3. Alcohol Problems. patients were medicated.71) 46.50 (8. Neg Negative Scale T score.02) 40. such as estimated Full Scale IQ and WRAT–III reading level. no patient score across any of the PAI scales was in a clinically elevated range (T score cutoffs 96. and Drug Problems. the Barnes Akathesia Rating Scale (Barnes.97 (6. Patients demonstrated significantly higher scores on several indexes of the PAI.001 . but significantly lower scores on the PIM scale. Unless otherwise noted. Experimental Apparatus Four target arrays were used. Pos Positive Scale T score. 1999).17 (8.85) 45.001 .71 (9.81 (5.007a 4.09 (7. Alcohol Problems. and cognitive instruments were administered by two graduate students who were trained via formal seminars and clinical practica.786 5. and any questionable responses were reviewed with a registered clinical psychologist (Bruce K.35 (4. Alc Alcohol Problems Scale. Christensen). Wilkinson.585) t(56.50) 35. but patients were impaired on a measure of verbal memory (e. An Anergia Scale T score.45 (8.20 (3.

GA was defined as the vector distance between the receivers on the index finger and thumb and measured at 10% intervals of MT from 10%–100%. GA reflects the opening and closing of the fingers that occurs when grasping an object. For the grasping trials. This parameter is normally scaled to the distance of the target object from the actor. in random order) were carried out followed by 40 target flanker experimental trials (target and flanker sizes and target positions were randomized). Ten target-only control trials were carried out for the 40-mm and 60-mm target objects in the grasping and sizeestimation tasks. GA Procedure Participants performed two tasks (grasping and size estimation) in separate sets of trials. increased TPGA can indicate that more of the movement was programmed in advance of movement onset. Goggles were occluded between trials and then cleared at the beginning of the next trial as the participant’s cue to respond. and wrist of the preferred hand. 1987). target array. and environment during the entire response. Overhead schematic of the stimulus display.500 ms after trial initiation. Movement Time (MT) Target arrays were presented on a flat table surface 15 cm distal to a central starting position. Milton. Time to Peak Grip Aperture (TPGA) TPGA was the time taken. Toronto. Milton. VT).GRASPING BEHAVIOR IN SCHIZOPHRENIA 803 800. Offline position data were filtered using a second-order dual pass Butterworth filter with a low-pass cutoff set at 10 Hz. Participants received as many practice trials as necessary to adequately perform all tasks. thumb. thumb. Goggle lenses remained clear for 2.. to reach PGA after movement onset. participants were instructed to lift the hand off the starting position and adjust the finger–thumb aperture to indicate the length of the target object. for a total of 80 target flanker trials in the entire study. one each on the index finger. This phase of the movement is thought to be programmed in advance of movement onset and is relatively uninfluenced by feedback processing. grasp the target object about its length and lift it a few centimeters off the table surface. Grasping Variables Figure 1. thumb. thus providing a good indication of visuomotor transformation characteristics. calculated as offset time minus onset time. For each task. VT) with three position sensors. Targets (40 mm or 60 mm) were marked with red circles and presented on either the left or right side of the display. For the size-estimation trials. Thus. Motion Analysis and Data Reduction Finger. without moving the hand toward the target. participants were instructed to quickly reach out. and illustrated right is a 60-mm target flanked by a smaller object. in milliseconds. Ascension Technologies Corporation. permitting full vision of the hand. PGA was established by Jeannerod (1986) as a clearly identifiable landmark that occurs within 60%–70% of the duration of the reach and is highly correlated with object size. Movement onset was defined as the point where wrist velocity exceeded 25 mm/s for five continuous frames. Peak Grip Aperture (PGA) PGA was defined as the maximum vector distance between the receivers on the index finger and thumb. Ascension Technologies Corporation. for a total of 40 target-only control trials in the entire study. Ascension Technologies Corporation. Milgram. MiniBirds receivers (MiniBirdsTM 800. The order of tasks was counterbalanced across participants. flanked on the other side by smaller or larger objects. Peak Wrist Velocity (PV) PV was defined as the fastest velocity reached during movement in the x direction for the marker on the wrist.5 s synchronized with the clearing of the goggle lenses. Canada.e. and wrist (sampling rate 144 Hz). VT) were taped to the index finger. thereby establishing PGA as the standard in terms of operationalizing and measuring grasping behavior. Many authors have since replicated Jeannerod’s work (reviewed in Castiello. and movement offset was determined as the point where grip aperture (GA) became stable (i. Hand kinematics were measured using a magnetic motion-tracking system (MiniBirds™ 800. 20 target-only control trials (40-mm and 60-mm targets. Visual events were controlled using goggles fitted with liquid-crystal lenses (PLATO Translucent Technologies. Position data were then differentiated using a 3-point central finite difference algorithm. Milton. 2005). with greater velocities associated with greater target distances. Threedimensional position data were sampled at 144 Hz for 2. MT was the duration of the movement. RT RT was a measure of response initiation that was calculated from the point of goggle clearing to the moment the hand was lifted from the home key position. Ten trials were carried out for each of the four target arrays (40-mm or 60-mm target. and wrist positions were tracked during the experiment with a magnetic motion-tracking system (MiniBirds™ . small or large flankers) in the grasping and sizeestimation tasks. Illustrated left is a 40-mm target flanked by a larger object. 0 change in aperture for five continuous frames). Ontario.

Only the kinematic features of the grasp are predicted to be affected by the illusion in SCZ patients because it is this component of grasping that is typically sensitive to the size of the target object (Jeannerod. The fourth section explores the relationship Patients’ and controls’ perception of the illusion were assessed by analyzing the SGA in the size-estimation trials.05. 82) 604. to reach PV after movement onset. and (c) grasp kinematics (PGA. There was no significant between-group difference. larger) mixed factorial analysis of variance (ANOVA).5 mm or 38%) target object. this phase of the movement is thought to be programmed prior to movement onset. defined by four consecutive points within a maximum tolerable percentage range of deviation of . participants perceived the larger targets to be larger than the smaller targets. and flanker size. (b) reaching kinematics (PV and TTPV). F(1.10.006.822. and the targets presented beside smaller flankers to be larger than the targets presented beside larger flankers. 82) 43. the dependent measure was SGA.50. Performance in the sizeestimation trials was taken as an indication of participants’ conscious perceptual experience of the illusion. AND WESTWOOD TTPV was the time. The third section explores the relationship of temporal aspects of prehension to illusion effects for patients and controls. A. SGA was analyzed with a Group (patient. 40-mm target) and flanker effect (larger size estimates for the smaller vs.001. 82) 8. Stable grip aperture (in millimeters) for size-estimation responses as a function of target and flanker size. or Group Target Size Flanker Size interaction. that is. the control and SCZ participants were equally sensitive to the size-contrast illusion in terms of their conscious perceptual experience.09%. p . 1986).05. The second section explores the effects of object size and flanker size on (a) temporal aspects of the prehension responses (RT and MT). The first section explores the effect of the size-contrast illusion on participants’ manual estimates of object size to determine whether the illusion had the expected effect on size perception.804 Time to Peak Wrist Velocity (TTPV) KING. control) Target Size (40 mm. p . TPGA. 82) 0. p . SGA was significantly affected by target size.64. larger flanker) are consistent with the well-established perceptual effects of this stimulus configuration. F(1. and significantly larger stable grip aperture was observed for smaller versus larger flanker trials. p . F(1. Perceptual Effects of the Size-Contrast Illusion SGA Perceptual Variables—Stable Grip Aperture (SGA) For size-estimation trials. A Target Size Flanker Size interaction. In summary. F(1. Results The experimental results are presented in several sections. p .3 mm or 55%) versus smaller (1.981. p . 40 mm target Large Flanker Small Flanker B. Bars represent within-subjects standard error of the mean. between symptoms and neurocognitive measures to illusion effects. Much like TPGA. F(1. executed with little influence from feedback processing. .01. 60 mm target Large Flanker Small Flanker 72 94 * 70 92 * * Stable Grip Aperture (mm) Stable Grip Aperture (mm) * 68 90 88 66 86 64 84 62 82 60 80 Patient Group Control Patient Group Control Figure 2. in milliseconds. 82) 0.001 (see Figure 2). The above-noted target effect (larger size estimates for the 60-mm vs. Significantly larger stable grip aperture was observed for the 60-mm target (Panel B) as compared with the 40-mm target (Panel A). 60 mm) Flanker Size (smaller. and GA at 10 relative time points during the action). indicated that the magnitude of the flanker effect was greater for the larger (3. CHRISTENSEN.

003.89) 583.52) 187. In other words. .26. F(1. 82) 2.73) 394. Results for MT indicated no significant between-group main effect. p . F(1. The only significant effect was a main effect for target size.006. p . An asterisk indicates that group means were significantly different at p .21.72) 410.40) 248. MT was greater for the 60-mm as compared with the 40-mm target object.500 ms for every trial. p . All values are in milliseconds. Reach Kinematics PV.001. 82) 1.23) 204.59 (104. The ANOVA for PV revealed no significant betweengroup main effect.57 (118.58) 638. there was no significant betweengroup difference.31.84 (55.17. TPGA time to peak grip aperture. although RT was comparable for both flanker conditions for the 40-mm target. the grasping behavior of the control participants was insensitive to the size-contrast illusion.568.64) 391.68) 402. and flanker size. Moreover.76) 441. F(1. All SCZ patients responded well within the allotted 2.10. both groups showed similar scaling of their actions to features of the target and flanker objects. p . no RT data for any patient trial were excluded from the analysis.75) 191. 82) 3.746. TTPV time to peak wrist velocity.89) 249. Patients demonstrated comparable RT performance to controls.006. Remaining effects were not significant. and 60-mm.68) 451.236. RT reaction time.27 (123. The results for the temporal aspects of grasping indicate that there are no fundamental differences in the gross movement abilities of control and SCZ participants because the two groups initiated movements with similar latencies and took about the same amount of time to complete those movements. or the 60-mm target.71) 251.64) 197. t(42) 5.46 (73.00.249.29) 197. F(1. a significant effect of flanker size.17) 363. 82) 29.05) 590.617. p . t(42) 6. Simple effects testing revealed that for controls.70 (121.40) 232. TPGA. p . t(42) 0. TTPV was significantly greater for patients compared with controls for the 60-mm target object presented beside small flankers (see Table 3). PV was larger for the 60-mm compared with the 40-mm target object.76 (88. p . In addition.001. F(1. 1986). indicating that both groups demonstrated greater SGA for the 60-mm target compared with the 40-mm target.73 (47.00. for the 10 target-only control trials carried out for the 40-mm and 60-mm target objects in the size-estimation task.18) 439. F(1.54) 424.41) 395. MT movement time.08.92 (36. 60L 60-mm target with large flanker.19 (88.76.94 (120. these results suggest that patients and controls are fundamentally similar in the control of reaching. and. F(1. and MT was greater for targets presented beside large compared with small flankers (see Table 3).10 (112. Visuomotor Effects of the Size-Contrast Illusion Temporal Aspects of Prehension RT. p . F(1. MT. 82) 7. No other results were significant.58 (40.96 (40. 82) 0. but significant within-group main effects for target size. Results for TTPV indicated no significant betweengroup main effect. 82) 1. the lack of group differences is not readily accounted for by data that were eliminated for slow patients.13) 386.09 (124. 40S 40-mm target with small flanker.001 (see Figure 3).671.17 (82.320.32) 634.12. 82) 3. and a significant Target Size Flanker Size interaction. RT was greater for the 60-mm compared with the 40-mm target object and when targets were presented beside large compared with small flankers. and a Flanker Size Group interaction. or interaction.39 (56. Remaining interactions were not significant. 82) 9.913.651.79) 192. 60S 60-mm target with small flanker. targets. F(1.08 (103.001. t(42) 1.272.674.14) 188.019. F(1.53 (131.45) 595. p .66 (31.81. The TPGA analysis revealed a significant betweengroup effect.88) 593. F(1.76 (34. and significant main Note. 82) 3.06) 231.35. F(1. TTPV. F(1.28 (73. whereas robust illusion effects were seen in the grasping movements of the SCZ participants. Results for the PGA analysis indicated a significant effect of target size. PV peak wrist velocity.02) 374.35 (59.85 (94. 40L 40-mm target with large flanker. Table 3 Group Means (and Standard Deviations) for Temporal Aspects of Prehension and Kinematics of Wrist Motion Variable RT-40L MT-40L PV-40L TTPV-40L TPGA-40L RT-40S MT-40S PV-40S TTPV-40S TPGA-40S RT-60L MT-60L PV-60L TTPV-60L TPGA-60L RT-60S MT-60S PV-60S TTPV-60S TPGA-60S Controls 226. Because PV is related to the reaching rather than the grasping component of prehension (Jeannerod. 82) 1.12) 644.77) 632. In contrast.110. consequently. 82) 0.97. Remaining interactions were not significant.15 (78.579.38. but a significant within-group main effect for target size.60 (27.114. despite the very robust effect of the flankers on perceptual size judgments reported earlier. 82) 735.001.64 (98.82 (111. Results for RT revealed no significant between-group effect. PGA was not affected by the flankers in the stimulus array for either the 40-mm target.55 (120.28.30) Patients 236.21 (51. p .81 (101.88 (86. Therefore.92 (124. F(1. PGA was affected by flanker size in a similar magnitude to the effect observed in the size-estimation task for both the 40-mm.53 (42. p .001.62) 196. F(1. 82) 41.001.GRASPING BEHAVIOR IN SCHIZOPHRENIA 805 Perceptual Control Trials As further support for the similar perceptual experiences of the SCZ and control participants.001. but a Target Size Flanker Size Group interaction.88.90 (81. p . p . p . p . and flanker size.89 (96.55) Grasp Kinematics PGA. p .88 (125.23) 377.17 (84. 82) 7.32. 82) 25.50. but significant within-group main effects for target size.05. However.001.127.78) 227. F(1. F(1. p . p . 82) 3. p .26 (119.06. for the patient group. p .41 (31. TTPV was comparable between groups for the 40-mm target object presented beside the small and large flankers and the 60-mm target object presented beside large flankers.001. p . 82) 5.42) 422.26) 374. which would have conceivably biased the RT data and other temporal aspects of prehension data. RT was significantly greater for the 60-mm target object when presented beside large compared with small flankers (see Table 3). F(1.15) 433. p .

82) 0. Rather. with the exception of a significant Group Target Size interaction for PV. For PGA. In other words.88. p .907. Remaining interactions were also not significant. Peak grip aperture (in millimeters) for grasping responses as a function of target and flanker size.. F(1.72. Because no overall group difference was found for MT. RT. group differences were analyzed for the 10 target-only control trials in which the 40-mm and 60-mm target objects were presented without flankers.29. p . F(1.32.335. These results suggest that although patients and controls are similar in the control of reaching (as indicated from comparable TTPV data). 40 mm target 100 B. Grasping control trials. F(1.001. p . Similarly. CHRISTENSEN.806 KING. p .e. p . SD 9. 80) 0. The similar grasping performance of SCZ and control participants in the target-only control trials indicates that the different effects of the size-contrast illusion on grasping for the two groups— central to our experimental hypothesis—is not merely a reflection of gross differences in grasping ability for the two groups. 60 mm target 115 Large Flanker Small Flanker Large Flanker Small Flanker 95 * Peak Grip Aperture (mm) 110 * Peak Grip Aperture (mm) 90 105 85 100 80 Patient Group Control 95 Patient Group Control Figure 3. F(1.51. No interactions were significant. For the patient group. However. effects for target size. This subtle difference permits an interpretation of the differences in grasping behavior between SCZ and control participants in terms of dorsal and ventral stream functions. F(1. p . F(1. there were no significant between-group differences or interactions across any of the temporal aspects of prehension (i. 80) 2.006. regardless of perceptual illusions).494. of importance. which is the phase of the action where feedback-based control is most important.01. or Group Order interaction. There was a significant main effect for target size. 80) 0. a smaller proportion of the movement duration for the patient group was spent under feedback control.001. indicating that perceptual estimates were greater for individuals who performed the grasping task first (M 81. Simple effects testing revealed that patients demonstrated significantly slower PV for the 40-mm target but comparable PV with controls for the 60-mm target. there was no significant Group Order interaction.46. significantly larger peak grip aperture was observed for the 60-mm target (Panel B) as compared with the 40-mm target (Panel A). and flanker size. To establish whether task order modulated the illusion effect.08. SD 10. p . and significantly larger peak grip aperture was observed for smaller versus larger flanker trials. F(1. To determine whether SCZ and control participants simply grasp all objects in a different manner (i. Remaining interactions were not significant.91. . 82) 5. 82) 871.013. Bars represent within-subjects standard error of the mean. F(1. Order effects. there were no significant between-group differences or interactions noted for the kinematics of wrist motion. F(1.95) versus those who performed the perceptual task first (M 76. The results for PGA indicated no significant effect of order. Although significantly larger peak grip aperture was observed for the 60-mm target (Panel B) versus the 40-mm target (Panel A). the different effects of the illusion can be attributed to a difference in the way that contextual visual cues are used for the control of grasping in the two groups. indicating that both groups used larger PGAs for the 60-mm target compared with the 40-mm target.121. we performed two separate Order Group Target Size Flanker Size factorial ANOVAs with PGA and SGA as separate dependent variables. MT). 80) 6. there was no significant between-group difference.47.024. or Group Target Size interaction.e. p . F(1. with patients spending more time prior to reaching PGA. In addition. p . 82) 7. no significant effect of flanker size was detected for the control group.. Results for SGA indicated a significant between-group effect of order.781. 82) 20.94. 82) 0. indicating that the order effect influenced both groups equally.94.05. this means that patients would also have spent less time after PGA.19). they differ somewhat in the control of grasping. p . p . AND WESTWOOD A.

and TPGA and illusion effects in grasping.. 60 mm) Flanker Size (smaller. a number of cognitive variables.080. However. PV. 738) 289. suggesting that slowness of the movement was not a confound for the illusion effects observed in grasping for the patient group. Haffenden & Goodale. the enhanced effect of the sizecontrast illusion in SCZ patients’ grasping movements does not appear to be an artifact of clinical or demographic factors.5 to . Similarly. The analysis of PGA presented earlier indicates that the grasping movements of SCZ patients were affected by the size-contrast illusion. In the current study. 1998). 1995. because RT is a measure of response initiation. SCZ participants were not grossly impaired in all aspects of prehensile behavior. a measure that indicates the relative proportion of the grasping component that is programmed in advance of the response and executed without online feedback control. 30%. if SCZ is associated with dorsal stream impairment. In this regard.14. However.e. 40%.4) compared with medium to large effects (Cohen’s d ranging from . Among healthy participants. particularly on measures of self-reported psychopathology. Nevertheless. Results indicated a trend toward a significant betweengroup effect. greater illusion effects in estimating would be negatively correlated with positive symptoms and positively correlated with negative symptoms.295. It is important to note that these observations indicate that the increased illusion effect on grasp formation in SCZ participants cannot be attributed to faster or slower responding or to an increased or decreased proportion of time spent using online visual feedback. or other kinematic parameters associated with the reaching component of the movement (i. Discussion The current experiment used a visual illusion paradigm to test the hypothesis that SCZ is associated with a defect in dorsal brain functioning and spared ventral brain functioning.64. To our knowledge. p .028. 70%. were weakly to moderately related (i. which has often neglected existing models of brain development and evolution.058) for the patient group only. most notably verbal memory and IQ. 90%. it is more directly related to overall MT than PGA—from which illusion effects are determined. This approach is relatively novel and improves on previous SCZ research. Conversely.e. whereby early and late time points of movement were indistinguishable for patients and controls compared with midpoint values. grasping has been shown to resist the effects of visual illusions whereas perceptual estimates have been shown to be highly sensitive (Aglioti et al. . To explore more carefully where group differences in GA occurred. this is also the first study to investigate perceptual and motor functions in SCZ using a behavioral paradigm designed to dissociate dorsal versus ventral stream processing.. p . p . there appeared to be a consistent pattern. However. and 100% of MT) mixed factorial ANOVA. group differences in TPGA could not account for the different illusion effects in grasping because no significant correlation was observed between the two measures for either patients or controls. F(9. at a macroscopic level. 50%. the results did not change when age was modeled as a covariate. then greater illusion effects in grasping should be positively correlated with negative symptoms and negatively correlated with positive symptoms. 80%. TTPV) in the illusion trials. Neurocognitive Measures In general. TPGA is more directly related to PGA. a number of clinical covariates were identified for the SCZ group. larger) Relative Time (10%. control) Target Size (40 mm. However. with the exception of significant differences at the 40%.. MT. but perception was not. In contrast to the controls. Pearson r values ranging from . the reaching component of prehension is similar for patients and controls. A trend-level significant negative correlation was observed for RT with illusion effects for the small target (r . no clear associations were observed between negative and positive symptoms for SCZ patients with illusion effects in grasping or estimating consistent with this pattern.8) for midpoints of movement. 20%. These findings suggest that. Although patients were significantly younger than controls. a majority of these significant effects did not survive strict Bonferroni adjustments. Symptoms and Neurocognitive Measures: Relationship to Grasping and Size Estimation Clinical Symptoms On the basis of the proposed functional divisions of dorsal and ventral brain circuitry. calculation of effect sizes revealed consistently smaller effects for early and late time points of movement (Cohen’s d ranging from . Although SCZ Relationship of Temporal Aspects of Prehension to Illusion Effects in Grasping To establish whether temporal aspects of prehension might have influenced illusion effects in grasping. we conducted separate within-group Pearson r correlations between RT. Nonetheless. 82) 3.21) to illusion effects in grasping and perception. statistically controlling for these variables did not affect the results. This pattern of results is consistent with a relatively selective SCZ deficit in dorsal stream function with sparing of ventral stream function. Results indicated that neither MT nor TPGA were significantly related to illusion effects across both target sizes for either group. F(1. and a significant Group Relative Time interaction.GRASPING BEHAVIOR IN SCHIZOPHRENIA 807 GA throughout the movement. Therefore. we sought to interpret our findings as reflecting important neuroanatomical organization consequent to neurodevelopment anomalies.01 to . Indeed. Simple effects testing revealed that patients and controls demonstrated comparable GA across all time points. no significant relationships between TPGA and illusion effects were observed for patients or controls. 60%. 50%. we analyzed GA with a Group (patient.01 to . and no consistent pattern was observed across all variables in each group. and 60% points of the movement profile. MT. the SCZ participants were sensitive to the size-contrast illusion in both the grasping and perception tasks. however. control participants in the current study showed the typical pattern: Grasping was immune to the size-contrast illusion. Consistent with previous findings. No group differences were seen in control nonillusory trials or in RT. indicating that patients who were slower to initiate the grasping movement demonstrated larger illusion effects. There were also no clinical or demographic differences between our comparison groups that could account for these findings. The only group effect obtained was for TPGA in the illusion trials.

& Javitt. Doniger. parahippocampal gyrus (Brown et al. Flashman. Indeed. Ruf. & Milner. 2003). including aspects of face processing (Williams. 2002). high spatial frequency detection (O’Donnell et al. spatial localization and trajectory discrimination (O’Donnell et al.. magnocellular deficits have been extensively linked to dyslexia. 2000. Group differences in illusion effects were noted only during the midphase of the movement profile. 2002). The perceptual analysis of the target in the ventral stream falls prey to sizecontrast illusions. 1995. Nuechterlein. such as forward masking (Saccuzzo. Green. including deficits in global analysis (Place & Gilmore. why.G. illusion effects are seen toward the midpoint of the grasping actions of SCZ patients. which are intact (Selemon et al.. 1996). There- . stereologic evidence indicates that areas of dorsolateral PFC exhibit more pronounced pathologic abnormalities than ventral PFC regions.. 1995). Higgins. physiological disruptions in areas specialized for object processing (Doniger et al. Thus. CHRISTENSEN. but not for controls. It has been shown that disruptions in areas specialized for object processing might occur secondary to dorsal visual stream dysfunction (Doniger et al. 2002). the greatest morphometric changes in SCZ are consistently reported in dorsal brain structures like the hippocampus (Gur et al. 2002). 1998). Accordingly. 1994). A number of visual neurophysiological studies (Butler et al. Nelson. movement programming is instead guided by an intact representation of the target that is generated by the perceptual mechanisms located in the ventral visual stream. we interpret our findings as evidence for disrupted movement programming processes in SCZ—processes that depend on visuomotor transformations normally mediated by the dorsal visual stream (Milner & Goodale. Findings of impaired visuomotor functioning in SCZ are broadly consistent with literature revealing SCZ deficits on dorsally mediated processes and/or visual tasks.808 KING. her performance improved dramatically when the response was delayed for 5 s. Murray. 2003. additional research that directly focuses on understanding these differences as a function of time is needed. showed poor scaling of GA to object size when the action was carried out immediately upon seeing the target. Presumably. 2004). Serper. and the generation of saccadic eye movements (Chen.. spatial working memory (Tek et al. a consistent pattern emerged showing small effect sizes at early and late time points compared with medium to large effect sizes at middle time points. Loughland. Silverstein et al. 2000). some of these findings might reflect primary dorsal steam impairment. this response delay allowed I. 1996. 1995). 1997). was able to use her intact perceptual abilities to produce relatively normal grasping movements.. Nonetheless. & McHaffie. Stein.. 1999). In the middle stages of the action.. the processing of visual details over context (Place & Gilmore. This analysis revealed some important observations that speak to the specificity of the visuomotor deficit in SCZ patients. a relatively crude measure of grasping control that does not specify the point in time where this kinematic landmark occurs. & Braff. and areas of dorsolateral and dorsomedial prefrontal cortex (PFC.. 1998). low spatial frequency processing (Slaghuis. & Riordan. however. it should be emphasized that the current temporal comparison was exploratory and not the focus of the current study. Future research could illuminate the impact of visual feedback by using closed loop paradigms in which vision is occluded during reaching. In addition. Milner et al. The absence of group differences in later stages of the action suggests that SCZ participants may have used online feedback-based control to correct earlier effects of the size-contrast illusion (Desmurget & Grafton. 1994. In an effort to better understand the time course of this effect. 1999). However. Nakayama. Although a number of these effects did not survive strict multiple comparison alpha adjustments. However.. 1998). Levy. The absence of illusion effects in early stages of the action is probably related to inertia of the hand and arm. the current study is the first to link these motor deficits to a dysfunction in specific brain regions. do SCZ patients seem able to correct their grasping actions toward the end of the response? The answer probably lies in the observation that movement programming and online movement control may have distinct neural substrates within the dorsal stream (Desmurget & Grafton. 1998. to control her action using her perceptual memory for the previously viewed object. Tost. I. 2006. motion discrimination (Chen. it takes time for the motor system to move the hand and fingers. (2001) reported that I. The observed illusion effects in grasping for SCZ patients was indexed by PGA. perhaps only the former substrates are affected in SCZ. Given that the online feedback control of action is also mediated by dorsal stream mechanisms. 2003)— circuits that might be unaffected in SCZ. 2006). 1996). et al. Levy.. To compensate for this disruption. Palafox. 2002) have also demonstrated relatively preserved ventral stream function in the context of selective physiological disruptions in dorsal visual brain regions. whereby individuals with reading impairment demonstrate deficits on psychophysical measures subserved by the dorsal visual pathway (Revheim et al. Weber-Fahr. then. Saykin. 1980. 1980). and the use of nonmotion cues to process motion (Chen..G.. Overall.. Feedback control would have been possible because the hand and target remained visible throughout the movement. et al. AND WESTWOOD deficits in motor planning and execution have been previously reported (Carnahan et al. Matthysse. we conducted a temporal analysis of GA formation at 10 time points throughout the action. 1996). and reading disability (Revheim et al. 1999). an optic ataxic individual with bilateral damage to the dorsal visual stream.. Westwood. & Davidson. reduced volume in fusiform gyrus (Onitsuka et al. Foxe..G. In contrast. Cadenhead. intact SCZ performance has been reported across ventrally mediated visual processes. the effect of the size-contrast illusion for SCZ patients may reflect a deficiency in the visuomotor transformations that were carried out before the onset of the movement during response programming.. & Henn. rendering it difficult to detect subtle effects of the sizecontrast illusion.. 2000). which appears to be unrelated to medication (Braus. & Holzman. 2006). Alternatively. Nonetheless. online movement control may be implemented by cerebellar and/or subcortical circuits outside of the dorsal visual stream (Jiang. Gordon. 1999). which capitalize on the relative scene-based metrics that characterize perceptual experience... Selemon et al. & Mintz. In addition. 2002). backward masking (Cadenhead. SCZ impairments have been reported in ventrally mediated functions. there is precedent in the neuropsychological literature that the ventral stream can contribute to the control of grasping movements when dorsal stream processing is disrupted. 1986). & Braff. Talcott et al. Past research has found that control of actions to remembered objects depends on visual processing taking place in the ventral visual stream (Goodale.

Moreover.. One possible explanation that could account for these findings is a restricted range of data. S. Washington.. Y. Current Biology. Schizophrenia Research. 157. J. a strength of the current study is that patient and control groups were studied at the same time using the same stimuli. G... (2001). 679 – 685. R. E. S. Goldman. Postmortem evidence of structural brain changes in schizophrenia. Thus. temporal horn area. & Jones. 43. This is potentially problematic given the strong links between cognitive functioning and level of functional impairment in SCZ (Green. D. R. Tost. unlike the current stable outpatient sample. Future studies using careful. Biological Psychiatry. 109 –113. Finally. 43. as well as differences in which brain regions mediate these processes and how this information is processed. as the SCZ patient sample was not highly symptomatic and was relatively high functioning in terms of cognition. D. L. J. (1989). who. which is consistent with the findings of the current experiment. Brown. These findings do not support the PAM... Amador. C. R. X. findings of illusory effects in grasping do not necessarily refute the hypothesis that perception and action are mediated by distinct visual systems. Cadenhead. H.. V.. R. (1994). Neuropsychology of first-episode schizophrenia: Initial characterization and clinical correlates. J. Extending this paradigm to include chronically ill patient groups would address the generalizability of this model. absence of a healthy control group comparison makes it difficult to anchor these results in a meaningful way. (1996). . Diagnostic and statistical manual of mental disorders (4th ed. FL: Psychological Assessment Resources. 59. (1994).. Archives of General Psychiatry. the only SCZ patients who demonstrated reduced susceptibility to the visual illusion were those who exhibited a high level of disorganized symptoms.. R.. Schizophrenia Research. Mitchell.. Depressive symptoms in stable chronic schizophrenia: Prevalence and relationship to psychopathology and treatment. L. Moreover.. Lynch. & Carnahan. Montgomery.. Mendoza. A. Size-contrast illusions deceive the eye but not the hand. References Aglioti. & Henn.. Butler. timing. 132–138. E.. Glover & Dixon. 36 – 42. including subtle differences in the timing of visual events. Ruf. J. Chua.. DeSouza. Sensory information processing in neuroleptic-naive first-episode schizophrenic patients: A functional magnetic resonance imaging study. P. J. Carnahan.. Meegan et al.. Franz. 2001.. (2003). Jr. suggesting that these deficits might be an index of illness severity. methodological differences across studies. D. Haffenden.GRASPING BEHAVIOR IN SCHIZOPHRENIA 809 fore. F.. Meegan. Hetrick. P. A. 5. and visual parameters. J. F. & Welsh. D.. L. Fahle.. Baynes. Differences in brain weight. Although most studies find that the grasping movements of healthy participants resist size-contrast illusions. Malla. S. might demonstrate a different pattern of perceptual and visuomotor behaviors. (1986). However. DC: Author. J. A.. Serper. and responses may lead to enhanced effects of illusory displays on visuomotor control. illusion-inducing elements are perceived as obstacles to the movement trajectory. Elliott. F.g. H. Hopkins Verbal Learning Test– Revised. most notably verbal memory performance.. Cooper.. 2006). Velamoor. Additionally. T... & Gegenfurtner. Bunney.. & Benedict. Reiter. Harkavy-Friedman.). a number of cognitive variables. D. H. Transient versus sustained visual channels in the visual backward masking deficits of schizophrenia patients. Gegenfurtner. An investigation into movement planning and execution deficits in individuals with schizophrenia. C. used an alternative forced-choice paradigm in which percent accuracy ranged from 43% to 57% across groups. A rating scale for drug-induced akathisia. Bell. Colter.. Bates. J. Odessa. C.g. DeSanti. D. & Goodale. Bulthoff. (2000). (2001). and Silverstein (2006). American Psychiatric Association. 2004.. Kim. W.. Akbarian. Phillips. G. methodological factors cannot account for observations that the control group resisted the illusion in grasping but the SCZ group did not. D. Archives of General Psychiatry.. Crow. N. G. the visual–visuomotor nature of the tasks may explain the weak relationship with verbal memory performance. Impaired perceptual grouping deficits have been linked to more chronic illness status and higher levels of disorganized symptoms in both SCZ and schizotypy.. 253–260. Barnes. The stimuli used to induce perceptual illusions may affect grasping movements for very different reasons (e. W. 1996). Carnahan.. 199 –209. 696 –701. Jagoe. The current research is limited in addressing whether the DTT is generalizable to more chronically ill SCZ groups. W. who reported that a subgroup of disorganized-symptom SCZ patients demonstrated less perceptual susceptibility relative to other patient groups to the Ebbinghaus illusion. A. 16.. Elliott. some studies do not (e. Methodological differences between our tasks and illness severity differences in patient samples likely contributed to the disparity. et al. Corsellis. Schizophrenia Research. & Braff. R. MacFlynn. et al. were only moderately related to performance on dorsal and ventral tasks. Potkin. In this regard. Goetz. & Fahle. 23. and parahippocampal gyrus compared with affective disorder. 5. S. R.g. 2001). 154. J. Uhlhaas et al. O. In addition. Schiff. the pathological process in SCZ appears to demonstrate a disproportionate regional specificity for dorsal brain circuitry compared with ventral regions.. time-locked transcranial magnetic stimulation would be useful in addressing some of these issues. Maldistribution of interstitial neurons in prefrontal white matter of the brains of schizophrenic patients. P... et al. when taken on balance. et al.. (1995). British Journal of Psychiatry. and alternative models have been proposed (e. R. F. Maddox. Mulholland. 213–221. 47–56. G. Braus. Finally. 59. R. G. Effects of schizophrenia and prefrontal leukotomy on movement preparation and generation.. & Goodale. T.. Bilder.. Archives of General Psychiatry.. R. Aguilar. Brandt. Robinson. Visual backward-masking deficits in schizophrenia: Relationship to visual pathway function and symptomatology. 425– 436. American Journal of Psychiatry. (1998). H. suggesting the possibility of floor effects due to difficulty confounds. 45. C. M. & Norman. M. 2001). R. Do action systems resist visual illusions? Trends in Cognitive Science. 53. Perceptual task results from the current study are at odds with the findings of Uhlhaas. (2000). Weber-Fahr. (2002). which rendered those patients insensitive to the surrounding context elements that induced a size distortion. M. A.. 672– 676.. Lyons. A. (1997)... M. Carey. Frith. S. R.. Franz.’s study. the current kinematic data are insufficient to address the possibility of crucial differences in the timing of when different aspects of action control are executed for patients and controls. These findings were interpreted as evidence for ventral deficits. In Uhlhaas et al. Journal of Clinical and Experimental Neuropsychology.. Bulthoff. S. D. the patients in the current sample did not demonstrate a high degree of disorganized symptoms. 2000. J. B. 549 –559. K. J. A potentially limiting aspect of the current experiment is failure to demonstrate clear differentiation between SCZ patient symptomatology with illusion effects in grasping and perceptual estimation.

R. Frith. Palafox. M. 153. Foxe. & Javitt.. Desmurget. Carnahan. D. Franz. S. Psychophysical evidence for separate channels for the perception of form. H. M. & Holzman. (2001).. B. New York: Biometrics Research. R. Goldberg. 136 –147. 6. Murray. & Velamoor.. & Bilder. M. V. K. (1996). M. P. The dual origin hypothesis: An evolutionary brain– behavior framework for analyzing psychiatric disorders... 56. Schiff. Chen. & Dixon. Neuroscience and Biobehavioral Reviews. M. 155. R. (2004).. W. S.. & Velamoor.. Latshaw. Dhillon. (2001). Washington. 17.. & Gegenfurtner. Journal of Neuroscience. Christensen. 1011– 1020. Dual cytoarchitectonic trends: An evolutionary model of frontal lobe functioning and its application to psychopathology. 13–23.. J. D. 4724 – 4729. Forward modeling allows feedback control for fast reaching movements. 158. Jaskiw. 247–256. 122. A. T. E. Archives of General Psychiatry. 1124 –1144. Perecman (Ed. P. K. Gegenfurtner. E.. D. 37– 47. N. June 26).. Archives of General Psychiatry. (2000). 437– 443.. Goodale. Impaired visual object recognition and dorsal/ventral stream interaction in schizophrenia. A neurological dissociation between perceiving objects and grasping them. Y. S. Danion. Meegan. B. Nakayama.. Influence of object size on prehension in leukotomized and unleukotomized individuals with schizophrenia. Matthysse. D. (1983). Psychosis as a state of aberrant salience: A framework linking biology. L. K. Jiang. I. Esiri. The dissociation between perception and action in the Ebbinghaus illusion: Nonillusory effects of pictorial cues on grasp. M. & Mintz. Dependence of impaired eye tracking on deficient velocity discrimination in schizophrenia. Experimental Brain Research. A. R. A. & Fahle. & Williams. (2006).. The neuropathology of schizophrenia: A critical review of the data and their interpretation.. Neuropsychopharmacology. M. C. E. B. Brain.. Joel. and depth. Fiszbein. 154 –156. K. A. D. & Zakzanis. 55. 51. Higgins. Psychological Science. The Positive and Negative Syndrome Scale for Schizophrenia. H. From intent to action: Evolution and function of the premotor systems of the frontal lobe. 67–75. Trends in Cognitive Science. R. In E.. J. D.. (2004).. & Hubel. H. R. Two distinct modes of control for object-directed action. D. L. Galaburda. Livingston. Gur. & Done. S. K... D. L.. A. Harrison. 14. R. (1991. Kauffmann-Muller. 593– 624. H. American Journal of Psychiatry. G.S. 19. movement. M. R. Towards a neuropsychology of schizophrenia. (2002). (2000). Milner. & McHaffie. M. D.. M. R.. 261– 276. M. (2003). Neuropsychology. M. 55. D. Backward masking in schizophrenia and mania: I—Specifying a mechanism. 526 –550. 7. (2003. Electrolytic lesions of the medial prefrontal cortex in rats disrupt performance on an analog of the Wisconsin Card Sorting Test but do not disrupt latent inhibition: Implications for animal models of schizophrenia. Stein. F. S. Lipska. Current Biology. T. M. R.. V. V. 20 –25. A.. A. D. A.. 96 –103. 131–144. Westwood. First. 59. (2001). Tremblay. & Grafton. (1993).. Meegan. A follow-up magnetic resonance imaging study of schizophrenia: Relationship of neuroanatomical changes to clinical and neurobehavioral measures. D. & Carey. (1994)... B.. 423– 431. colour. Reduced dorsal and orbital prefrontal gray matter volumes in schizophrenia. Nuechterlein. Amygdala volume in schizophrenia: Post-mortem study and review of magnetic resonance imaging findings. T.. Fahle. phenomenology. Bulthoff. A.. R. (1999). C. A. Goodale. (1999). Kapur. Spitzer. M.. Haffenden. M. Carnahan. G. H. DC: U. H. Gur. (1996). A.. 45. & Crow. M. Journal of Experimental Psychology: Human Perception and Performance. P. 180. 155–161. Cannon. C. Glover.. Y. & Milner. and pharmacology in schizophrenia. 221. Department of Health Education and Welfare. The neuroscience of grasping. M. Weiner. B. Journal of Comparative Neurology. Nature Reviews Neuroscience. Bilker. Cowell. (2002). Gibbon. E. W. 85. Gallacher. Grasping visual illusions: No evidence for a dissociation between perception and action. 331–338. et al. I. A. P. & Weinberger. 169 –184. S.. Levy. (1988). Franz. J. K.. C.). & Gur. 944 –948. F. (2000).. 57. & Goodale. (1976)... S. 10.. & Holzman. Elliott. Schizophrenia Bulletin.810 KING. 13. V. Motion perception in schizophrenia. Levy. E. N. G. The frontal lobes revisited (pp. Turetsky.. F.. L. Cowell. 939 –944.. Turetsky. & Westwood. (1995).. K. 30.. Nature. J. J. E. 99 –116. Journal of Cognitive Neuroscience... (1999). 153. I. Canadian Journal of Psychiatry.. L... Palafox. 144. Welsh. Doniger. 96. W. 149 –154. New York State Psychiatric Institute. Elliott.. Nature. 12. S. 349. S. J. . D. Elliott. J. S. Guy. 761–768. V. Matthysse. R. British Journal of Psychiatry.. The formation of finger grip during prehension: A cortically mediated visuomotor pattern. 426 – 445. 267–278. H. T. 273–306). P. V. What are the functional consequences of neurocognitive deficits in schizophrenia? American Journal of Psychiatry. U. M. K. Neurocognitive deficit in schizophrenia: A quantitative review of the evidence. (2004). M. Green. Castiello.. J. 177–181. J. Proceedings of the National Academy of Sciences USA. (1987). (1987). P. P. 726 –736. A. Manual performance in leukotomized and unleukotomized individuals with schizophrenia. B. P... Archives of General Psychiatry. (1998). L. Effects of visual illusions on grasping. An evolving view of duplex vision: Separate but interacting cortical pathways for perception and action. Research Version. S. G. K. Bulthoff.. R.. K. J. (1998). Giaccio. The Muller–Lyer illusion affects the planning and control of manual aiming movements.... New York: IBRN Press.. Haffenden. Kay.. (2000). British Journal of Psychiatry. & Holzman. Structured Clinical Interview for DSM–IV–TR Axis I Disorders. Canadian Journal of Experimental Psychology.. (1987). Arnold. Behavioral Brain Research. M.. 187–201. Current Opinion in Neurobiology. & Vermaat. G. W. 18.. The intrinsic architectonic and connectional organization of the superior temporal region of the rhesus monkey. & Goodale. F. CHRISTENSEN. S.. S. Matthysse.. 145–152.. R. H.. Behavioral Brain Research. & Pandya. Glazebrook. Progress in Brain Research. J. Jeannerod. G. M. 982–986. G. Lyons. 423. Archives in General Psychiatry. D. D. (1998). J. A. Abnormal Involuntary Movement Scale (AIMS). (1986).. Archives in General Psychiatry. (2001). Heinrichs. D. 321–330. Chen. N. I. B. Levy. 9. The effect of pictorial illusion on prehension and perception. P.. & Feldon. A.. In ECDEU assessment manual for psychopharmacology (pp. AND WESTWOOD Goodale.. 3416 –3468. Psychophysical isolation of a motion-processing deficit in schizophrenics and their relatives and its association with impaired smooth pursuit. D. 534 –537). (1997). M. Jakobson. H.. The role of vision in the on-line correction of illusion effects on action. 11. C. January 10). 56. 27. Schizophrenia Research.. Green. (2005). Postpubertal emergence of hyperresponsiveness to stress and to amphetamine after neonatal excitotoxic hippocampal damage: A potential animal model of schizophrenia. D. (2001). Nakayama. Chen. V. D. L. Opposing basal ganglia processes shape midbrain visuomotor activity bilaterally. 203–211. Chance. & Elliott. A.. D. R. American Journal of Psychiatry. A. 4. S. Intact implicit learning in schizophrenia. H. B. & Opler. & Welsh.. Journal of Clinical and Experimental Neuropsychology. Meulemans. 122–136. 160. Nakayama. (1999).. H. L. M. A. Y. Grossman. P.. D. K. Mendoza. Archives in General Psychiatry. T. H. 11. E. M.. M.

. Tilikete. (1995). Visual scanpaths in schizophrenia: Is there a deficit in face recognition? Schizophrenia Research. O’Donnell. 189 –199. Journal of Experimental Psychology: Human Perception and Performance. B. D. L. W. Spencer... Selemon. F. 52. A.and negative-symptom schizophrenia. 145. B. 55. Arousal. Psychiatry Research. Wide Range Achievement Test (3rd ed. (2001). & Mishkin. Visual magnocellular impairment in adult developmental dyslexics. E.. Regional specificity in the neuropathologic substrates of schizophrenia: A morphometric analysis of Broca’s area 44 and area 9. N. F.. C. A. W. Vighetto. Onitsuka. D. Loughland.. 116 –149. L. P. Archives of General Psychiatry.. 455– 462. Neuro-Ophthamology. A. Selemon. Lucia. K. J. 59. G. (2006). P. In A... (1998). L. M. Milner. G. A. 60.. Tek. (1969). 404 – 423. 2006 Revision received November 14. (1980). & Braff.. J. L. Milgram.. Advances in the analysis of visual behavior (pp. Saccuzzo. 9. Oldfield. (1996). Psychological Review. Herman. Wilkinson... S. E. Grasping the past: Delay can improve visuomotor performance.). New York: Plenum. D. (1982). G. A. Wechsler.. CA: Jerome M. Sattler. Pandya. DE: Wide Range. A spectacle-mounted liquid-crystal tachistoscope. The visual brain in action. 238 –245. H. N... 805– 820. Place. Comparative architectonics of the neocortex of mammals and their evolutionary interpretation. Selective deficits in visual perception and recognition in schizophrenia. Schizophrenia Research. Shenton. E. Behavior Research Methods. 549 –586). & 811 Javitt. Schizophrenia Research. Inc. & Barnes. E. The assessment and analysis of handedness: The Edinburgh Inventory.. Journal of Abnormal Psychology. Smith. Williams. (1985).. Blaxton. Pandya.... E. Schwarzkopf. Frumin. Barber. Prefrontal cortex in relation to other cortical areas in rhesus monkey: Architecture and connections. 1564 – 1570. D.. J. & Goldman-Rakic. McKinnell. W. Morey. M. Abnormally high neuronal density in the schizophrenic cortex: A morphometric analysis of prefrontal area 9 and occipital area 17.). Jones (Eds. G. Sattler. F... D. (1990). Oxford. Gordon... Reading impairment and visual processing deficits in schizophrenia. Journal of Abnormal Psychology. J. Architecture and connection of cortical association areas. Gold. Pandya. Wechsler Adult Intelligence Scale (3rd ed. 2007 Accepted January 11.. Milner.. mediated? American Journal of Psychiatry. 410 – 420. Saykin. 620 – 625. D. M. Silipo. 89. S. B. C. Phillips. W. A. Odessa. M. not peripherally. C.. & Silverstein. C.. Rajkowska. M.. Pisella.. 167. Swearer. 146 –153. R. Archives of General Psychiatry.. 251–270. Mitchell.. J. 63–94. 409 – 418.. C. 19. P. S. Niznikiewicz. 97–113. B. 163. J. M.. H. Progress in Brain Research.. C. L. D.. N. Nelson. 1896 –1901. S.. & McGuinness. M. 37. Architecture and connections of the frontal lobe.. Visual perceptual and working memory impairments in schizophrenia.. C. D.. & Yeterian.. J. & GoldmanRakic. Current Biology. (1996). In E. & Robbins... H. N. 153. R. 449 – 456. C.. American Journal of Psychiatry. P. (2002). 433– 440. M. Perecman (Ed.. (2002). 687– 692. G. Schechter. & Riordan. Nestor. T.. 11. Slaghuis... G. (1975). A. 32. L. Mansfield (Ed.. H. E. 49 – 62. M. S. 40. F. R. G. Assessment of children: WAIS-III supplement. (1998). M. S. MA: MIT Press. T. (1998). Journal of Abnormal Psychology. (1996). (1999)... 82. Z. L. J. E. D. L. T.. 187–201. Jalbrzikowski. C. 105. Spatial frequency discrimination in schizophrenia. J. P. T. R. M. J. Willis-Owen. Perceptual grouping in disorganized schizophrenia. Knight. F. C. (1999). K. Backward versus forward visual masking deficits in schizophrenic patients: Centrally. Shenton.. R. P. and effort in the control of attention. Talcott. A. Kleinman. M. Contrast sensitivity for stationary and drifting spatial frequency gratings in positive. J. R. Sanides. Mrzljak. Y.. In R.. 3– 61).. T. Instruments and Computers. The effect of the Muller–Lyer illusion on the planning and control of manual aiming movements. R. D. Dijkerman. & Yeterian. 153. C. Neuropsychologia. Butler.. D. D. Cadenhead. Two cortical visual systems. L. C. 87. & Rossetti. C.. Stimulus configuration and context effects in perceptual organization in schizophrenia. C. Annals of the New York Academy of Science. H. Wilmington..). Received August 30. Osborn. 85. Nelson. 41–72). Wilk. M. S.. D. West. M. Barnes. Potts. K. M. pp. Uhlhaas. P. Journal of Abnormal Psychology. 2008 . L. Archives of General Psychiatry. (1987). & Kamin.. L.. 413– 422. M.. M. Flashman.. Hippocampal volume reduction in schizophrenia as assessed by magnetic resonance imaging: A meta-analytic study. & McCarley. McIntosh. San Diego. Richardson. W. L. & Stein. Perceptual organization in schizophrenia. Cambridge. M. Cerebral cortex: Association and auditory cortices (Vol. D. E. (1999). M. D. L. L. (1995). 69 –77. P. (1997). A. W. J. 20. American Journal of Psychiatry. The frontal lobes revisited (pp. W. G. McMahon. Nestor. Stylianopoulos. Peters & E. K. A... W.. A. (1971). D. (2003). New York: IBRN Press. C. Hansen. (1987). G. Functional and structural deficits in brain regions subserving face perception in schizophrenia. 4. Pantelis. (2006). Kuroki. & Ryan. Ungerleider. 111. TX: The Psychological Corporation.). & Davidson. S. & Buchanan. activation. J. England: Oxford University Press. Archives of General Psychiatry. G. H. Silverstein.GRASPING BEHAVIOR IN SCHIZOPHRENIA (2006). R. N. & Gilmore.. (1991). J. F. Comparison of set-shifting ability in patients with chronic schizophrenia and frontal lobe damage. I. P. O’Donnell. J. & Goodale. & McCarley. (1993). L.). Pribram. FL: Psychological Assessment Resources. (2006). Owen. San Antonio.. Personality Assessment Inventory. et al. D. Revheim. 105–117. 107. L.

Sign up to vote on this title
UsefulNot useful