The what and why of perceptual asymmetries in the visual domain
Identifieur interne : 001D21 ( Pmc/Checkpoint ); précédent : 001D20; suivant : 001D22The what and why of perceptual asymmetries in the visual domain
Auteurs : A. K. M. Rezaul Karim [Bangladesh] ; Haruyuki Kojima [Japon]Source :
- Advances in Cognitive Psychology [ 1895-1171 ] ; 2010.
Abstract
Perceptual asymmetry is one of the most important characteristics of our visual
functioning. We carefully reviewed the scientific literature in order to examine
such asymmetries, separating them into two major categories: within-visual field
asymmetries and between-visual field asymmetries. We explain these asymmetries
in terms of perceptual aspects or tasks, the
Url:
DOI: 10.2478/v10053-008-0080-6
PubMed: 21228922
PubMed Central: 3019986
Affiliations:
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">The <italic>what</italic> and <italic>why</italic> of perceptual
asymmetries in the visual domain</title><author><name sortKey="Karim, A K M Rezaul" sort="Karim, A K M Rezaul" uniqKey="Karim A" first="A. K. M. Rezaul" last="Karim">A. K. M. Rezaul Karim</name><affiliation wicri:level="1"><nlm:aff id="AU1"><addr-line>Department of Psychology, University of Dhaka, Bangladesh</addr-line></nlm:aff><country xml:lang="fr">Bangladesh</country><wicri:regionArea>Department of Psychology, University of Dhaka</wicri:regionArea><wicri:noRegion>University of Dhaka</wicri:noRegion></affiliation></author><author><name sortKey="Kojima, Haruyuki" sort="Kojima, Haruyuki" uniqKey="Kojima H" first="Haruyuki" last="Kojima">Haruyuki Kojima</name><affiliation wicri:level="1"><nlm:aff id="AU2"><addr-line>Graduate School of Human and Socio-environment Studies, Kanazawa University, Japan</addr-line></nlm:aff><country xml:lang="fr">Japon</country><wicri:regionArea>Graduate School of Human and Socio-environment Studies, Kanazawa University</wicri:regionArea><wicri:noRegion>Kanazawa University</wicri:noRegion></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">21228922</idno><idno type="pmc">3019986</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3019986</idno><idno type="RBID">PMC:3019986</idno><idno type="doi">10.2478/v10053-008-0080-6</idno><date when="2010">2010</date><idno type="wicri:Area/Pmc/Corpus">001B63</idno><idno type="wicri:Area/Pmc/Curation">001B63</idno><idno type="wicri:Area/Pmc/Checkpoint">001D21</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">The <italic>what</italic> and <italic>why</italic> of perceptual
asymmetries in the visual domain</title><author><name sortKey="Karim, A K M Rezaul" sort="Karim, A K M Rezaul" uniqKey="Karim A" first="A. K. M. Rezaul" last="Karim">A. K. M. Rezaul Karim</name><affiliation wicri:level="1"><nlm:aff id="AU1"><addr-line>Department of Psychology, University of Dhaka, Bangladesh</addr-line></nlm:aff><country xml:lang="fr">Bangladesh</country><wicri:regionArea>Department of Psychology, University of Dhaka</wicri:regionArea><wicri:noRegion>University of Dhaka</wicri:noRegion></affiliation></author><author><name sortKey="Kojima, Haruyuki" sort="Kojima, Haruyuki" uniqKey="Kojima H" first="Haruyuki" last="Kojima">Haruyuki Kojima</name><affiliation wicri:level="1"><nlm:aff id="AU2"><addr-line>Graduate School of Human and Socio-environment Studies, Kanazawa University, Japan</addr-line></nlm:aff><country xml:lang="fr">Japon</country><wicri:regionArea>Graduate School of Human and Socio-environment Studies, Kanazawa University</wicri:regionArea><wicri:noRegion>Kanazawa University</wicri:noRegion></affiliation></author></analytic><series><title level="j">Advances in Cognitive Psychology</title><idno type="eISSN">1895-1171</idno><imprint><date when="2010">2010</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Perceptual asymmetry is one of the most important characteristics of our visual
functioning. We carefully reviewed the scientific literature in order to examine
such asymmetries, separating them into two major categories: within-visual field
asymmetries and between-visual field asymmetries. We explain these asymmetries
in terms of perceptual aspects or tasks, the <italic>what</italic> of the
asymmetries; and in terms of underlying mechanisms, the <italic>why</italic> of
the asymmetries. Tthe within-visual field asymmetries are fundamental to
orientation, motion direction, and spatial frequency processing. between-visual
field asymmetries have been reported for a wide range of perceptual phenomena.
foveal dominance over the periphery, in particular, has been prominent for
visual acuity, contrast sensitivity, and colour discrimination. Tthis also holds
true for object or face recognition and reading performance. upper-lower visual
field asymmetries in favour of the lower have been demonstrated for temporal and
contrast sensitivities, visual acuity, spatial resolution, orientation, hue and
motion processing. Iin contrast, the upper field advantages have been seen in
visual search, apparent size, and object recognition tasks. left-right visual
field asymmetries include the left field dominance in spatial (e.g.,
orientation) processing and the right field dominance in non-spatial (e.g.,
temporal) processing. left field is also better at low spatial frequency or
global and coordinate spatial processing, whereas the right field is better at
high spatial frequency or local and categorical spatial processing. All these
asymmetries have inborn neural/physiological origins, the <italic>primary
why</italic>, but can be also susceptible to visual experience, the
<italic>critical why</italic> (promotes or blocks the asymmetries by
altering neural functions).</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Adams, W J" uniqKey="Adams W">W. J. Adams</name></author><author><name sortKey="Graf, E W" uniqKey="Graf E">E. W. Graf</name></author><author><name sortKey="Ernst, M O" uniqKey="Ernst M">M. O. Ernst</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Albright, T D" uniqKey="Albright T">T. D. Albright</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Albright, T D" uniqKey="Albright T">T. D. Albright</name></author><author><name sortKey="Desimone, R" uniqKey="Desimone R">R. Desimone</name></author><author><name sortKey="Gross, C G" uniqKey="Gross C">C. G. Gross</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Anstis, S" uniqKey="Anstis S">S. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Adv Cogn Psychol</journal-id><journal-id journal-id-type="publisher-id">acp</journal-id><journal-title-group><journal-title>Advances in Cognitive Psychology</journal-title></journal-title-group><issn pub-type="epub">1895-1171</issn><publisher><publisher-name>University of Finance and Management in Warsaw</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">21228922</article-id><article-id pub-id-type="pmc">3019986</article-id><article-id pub-id-type="doi">10.2478/v10053-008-0080-6</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>The <italic>what</italic> and <italic>why</italic> of perceptual
asymmetries in the visual domain</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Karim</surname><given-names>A. K. M. Rezaul</given-names></name><xref ref-type="aff" rid="AU1"><sup>1</sup></xref></contrib><contrib contrib-type="author"><name><surname>Kojima</surname><given-names>Haruyuki</given-names></name><xref ref-type="aff" rid="AU2"><sup>2</sup></xref></contrib></contrib-group><aff id="AU1"><label>1</label><addr-line>Department of Psychology, University of Dhaka, Bangladesh</addr-line></aff><aff id="AU2"><label>2</label><addr-line>Graduate School of Human and Socio-environment Studies, Kanazawa University, Japan</addr-line></aff><author-notes><corresp id="Cor1">Corresponding authors: A. K. M. Rezaul Karim, e-mail:
<email>karim.akmr.monscho06@ gmail.com</email>, and Haruyuki Kojima, e-mail:
<email>hkojima@kenroku.kanazawa-u.ac.jp</email></corresp></author-notes><pub-date pub-type="epub"><day>15</day><month>12</month><year>2010</year></pub-date><pub-date pub-type="collection"><year>2010</year></pub-date><volume>6</volume><fpage>103</fpage><lpage>115</lpage><history><date date-type="received"><day>25</day><month>1</month><year>2010</year></date><date date-type="accepted"><day>19</day><month>6</month><year>2010</year></date></history><permissions><copyright-statement>Copyright: © 2009 University of Finance and
Management in Warsaw</copyright-statement><copyright-year>2009</copyright-year><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/2.5/"><license-p>This is an open-access article distributed under the terms of the Creative
Commons Attribution License, which permits unrestricted use, distribution,
and reproduction in any medium, provided the original work is properly
cited.</license-p></license></permissions><abstract><p>Perceptual asymmetry is one of the most important characteristics of our visual
functioning. We carefully reviewed the scientific literature in order to examine
such asymmetries, separating them into two major categories: within-visual field
asymmetries and between-visual field asymmetries. We explain these asymmetries
in terms of perceptual aspects or tasks, the <italic>what</italic> of the
asymmetries; and in terms of underlying mechanisms, the <italic>why</italic> of
the asymmetries. Tthe within-visual field asymmetries are fundamental to
orientation, motion direction, and spatial frequency processing. between-visual
field asymmetries have been reported for a wide range of perceptual phenomena.
foveal dominance over the periphery, in particular, has been prominent for
visual acuity, contrast sensitivity, and colour discrimination. Tthis also holds
true for object or face recognition and reading performance. upper-lower visual
field asymmetries in favour of the lower have been demonstrated for temporal and
contrast sensitivities, visual acuity, spatial resolution, orientation, hue and
motion processing. Iin contrast, the upper field advantages have been seen in
visual search, apparent size, and object recognition tasks. left-right visual
field asymmetries include the left field dominance in spatial (e.g.,
orientation) processing and the right field dominance in non-spatial (e.g.,
temporal) processing. left field is also better at low spatial frequency or
global and coordinate spatial processing, whereas the right field is better at
high spatial frequency or local and categorical spatial processing. All these
asymmetries have inborn neural/physiological origins, the <italic>primary
why</italic>, but can be also susceptible to visual experience, the
<italic>critical why</italic> (promotes or blocks the asymmetries by
altering neural functions).</p></abstract><kwd-group><kwd>visual perception</kwd><kwd>asymmetry</kwd><kwd>within-visual field</kwd><kwd>between-visual field</kwd><kwd>primary why</kwd><kwd>critical why</kwd><kwd>neural mechanisms</kwd><kwd>hemispheric specialization</kwd><kwd>visual experience</kwd></kwd-group></article-meta></front><body><sec sec-type="" id="S1"><title>INTRODUCTION</title><p>Visual perception is the process of interpreting and organizing visual information.
It involves our ability to recognize and identify the distinguishing features of
visual images such as shape, size, orientation, position, colour, etc. We have very
powerful vision and visual perception, with many surprising properties. One of the
most prominent properties is the perceptual variability or asymmetry resulting from
the brain’s preferential responses to some visual stimuli and/or to
stimuli at some specific retinal location. For example, vertical stimuli are
perceived and represented better than oblique stimuli (e.g., <xref ref-type="bibr" rid="R20">Campbell, Kulikowski, & Levinson, 1966</xref>; <xref ref-type="bibr" rid="R55">Furmanski & Engel, 2000</xref>; <xref ref-type="bibr" rid="R119">Mitchell, Freeman, & Westheimer,
1967</xref>), sensitivity to foveal stimuli is stronger than to peripheral stimuli
(e.g., <xref ref-type="bibr" rid="R47">Duncan & Boynton, 2003</xref>; <xref ref-type="bibr" rid="R65">Hansen, Pracejus, & Gegenfurtner, 2009</xref>;
<xref ref-type="bibr" rid="R172">Virsu & Rovamo, 1979</xref>), etc. Over
the last few decades, researchers have identified dozens of phenomena exhibiting
perceptual asymmetries that may not be apparent in our conscious awareness while we
are accomplishing everyday tasks. Yet we still do not have an account that gives a
comprehensive global picture of perceptual asymmetries and their emergence. This
review is, therefore, an in-depth analysis of the perceptual asymmetries in visual
psychophysics and visual neurology that have been documented thus far.</p><p>A careful look at previous research in these two areas indicates that, while some
information is processed quickly or efficiently, some information may be delayed or
processed less efficiently within the same location of the visual field. On the
contrary, when information in a particular visual field location, say, central, is
efficiently processed it may be poorly processed in the opposite location (here,
peripheral). Thus, perceptual asymmetries in the visual domain can be separated into
two major categories: within-visual field (WVF) asymmetries and between-visual field
(BVF) asymmetries. In this review, we term the perceptual aspects or tasks in which
asymmetries appear the <italic>what</italic> of the asymmetries and the underlying
mechanisms the <italic>why</italic> of the asymmetries. The <italic>why</italic> of
perceptual asymmetries can be further divided into the <italic>primary why</italic>and the <italic>critical why</italic>. The <italic>primary why</italic> refers to
the physiological mechanisms or neural organizations we are born with. The
<italic>critical why</italic>, on the other hand, refers to the experiential or
learning factor that interacts with the <italic>primary why</italic> and thereby
changes the organizational and functional features of cortical neurons. However, for
ease of comprehension the what and the <italic>primary why</italic> of the
asymmetries are explained together, and followed by an explanation of the
<italic>critical why</italic>.</p></sec><sec id="S2"><title>THE <italic>WHAT</italic> AND <italic>PRIMARY WHY</italic> OF THE WITHIN-VISUAL
FIELD ASYMMETRIES</title><sec id="S2A"><title>Asymmetry in orientation processing</title><p>Visual perception exhibits many examples of anisotropic behaviour, where the
percept’s relationship to the stimulus changes with the orientation
of the stimulus. Specifically, psychophysical studies have shown that
performance is better at the cardinal than the oblique orientation in contrast
sensitivity (<xref ref-type="bibr" rid="R19">Campbell & Kulikowski,
1966</xref>; <xref ref-type="bibr" rid="R119">Mitchell et al., 1967</xref>),
stereoacuity (<xref ref-type="bibr" rid="R122">Mustillo, Francis, Oross, Fox,
& Orban, 1988</xref>), grating acuity (<xref ref-type="bibr" rid="R10">Berkley, Kitterle, & Watkins, 1975</xref>; <xref ref-type="bibr" rid="R20">Campbell et al., 1966</xref>), and vernier acuity
(<xref ref-type="bibr" rid="R35">Corwin, Moskowitz-Cook, & Green,
1977</xref>; <xref ref-type="bibr" rid="R153">Saarinen & Levi,
1995</xref>; <xref ref-type="bibr" rid="R175">Westheimer & Beard,
1998</xref>). This fact, often referred to as the oblique effect (<xref ref-type="bibr" rid="R5">Appelle, 1972</xref>), is most prominent in central
vision (e.g., <xref ref-type="bibr" rid="R10">Berkley et al., 1975</xref>; <xref ref-type="bibr" rid="R109">Mansfield, 1974</xref>).</p><p>The oblique effect is functionally important as V1 (primary visual cortex/striate
cortex) neurons are organized into orientation columns (<xref ref-type="bibr" rid="R74">Hubel & Wiesel, 1968</xref>, <xref ref-type="bibr" rid="R75">1974a</xref>). This has also been confirmed in later single-neuron
electrophysiological (<xref ref-type="bibr" rid="R45">DeValois, Yund, &
Hepler, 1982</xref>; <xref ref-type="bibr" rid="R102">Li, Peterson,
& Freeman, 2003</xref>), optical imaging (<xref ref-type="bibr" rid="R31">Coppola, White, Fitzpatrick, & Purves, 1998</xref>), and
fMRI (<xref ref-type="bibr" rid="R55">Furmanski & Engel, 2000</xref>)
studies. For example, an fMRI study has demonstrated that grating acuity is
finer for cardinal (horizontal and vertical) than for oblique stimuli (<xref ref-type="bibr" rid="R55">Furmanski & Engel</xref>). This study also
reveals a corresponding asymmetry in neural populations in V1, that is neural
responses in V1 are greater for cardinal than oblique stimuli.</p><p>In addition to the oblique effect, scientists have demonstrated
“horizontal-vertical” asymmetry in a variety of visual
tasks. For example, our contrast sensitivity and spatial resolution are better
along the horizontal mid-line of the visual field than along the vertical
mid-line (<xref ref-type="bibr" rid="R21">Carrasco, Talgar, & Cameron,
2001</xref>; <xref ref-type="bibr" rid="R146">Rijsdijk, Kroon, & van
der Wildt, 1980</xref>). This is consistent with the fact that within the
cardinal more cells are devoted to horizontal than vertical orientation (<xref ref-type="bibr" rid="R102">Li et al., 2003</xref>). Thus, the orientation
asymmetries have a primarily physiological or neural basis.</p><p>Very recently Karim and Kojima (<xref ref-type="bibr" rid="R83">2010</xref>,
<xref ref-type="bibr" rid="R84">in press</xref>) have demonstrated that,
within a specific orientation performance in vernier offset detection may vary
as a function of vernier configuration (spatial arrangement of light bars). In
one study, they have claimed that vernier offset detection at the cardinal
orientation depends on the relative position of the vernier features (<xref ref-type="bibr" rid="R83">Karim & Kojima, 2010</xref>).
Specifically, for a pair of horizontal light bars (vernier features) arranged
side-by-side with a large gap between them observers were, on average, better at
discriminating a vertical offset if the right-hand bar was below the left-hand
bar than vice versa. Similarly, for a pair of vertically oriented light bars,
one above the other, the horizontal offset detection was better if the lower bar
was on the left of the upper bar rather than on its right. In another study,
they have shown that this effect can be generalized to the oblique orientation
(<xref ref-type="bibr" rid="R84">Karim & Kojima, in press</xref>).
They concluded that these asymmetries might be due to neural preference or
selectivity for one particular vernier configuration rather than another and
that such preference possibly developed through early experience or through
evolution (<xref ref-type="bibr" rid="R84">Karim & Kojima, in
press</xref>).</p></sec><sec id="S2B"><title>Asymmetry in motion processing</title><p>Meridian-dependent effects (oblique effects) have also been found in our
perception of moving objects (e.g., <xref ref-type="bibr" rid="R28">Coletta,
Segu, & Tiana, 1993</xref>; <xref ref-type="bibr" rid="R104">Loffler
& Orbach, 2001</xref>; <xref ref-type="bibr" rid="R111">Matthews
& Qian, 1999</xref>). In general, psychophysical studies have
concentrated on the anisotropy of the precision in motion direction
discrimination. In contrast to motion detection thresholds, which have been
found to be isotropic (e.g., <xref ref-type="bibr" rid="R143">Raymond,
1994</xref>; <xref ref-type="bibr" rid="R169">Van de Grind, Koenderink, Van
Doorn, Milders, & Voerman, 1993</xref>), motion discrimination
thresholds depend on the absolute direction of motion. This meridian-dependent
anisotropy for direction of motion discrimination has been reported for random
dots (e.g., <xref ref-type="bibr" rid="R7">Ball & Sekuler, 1982</xref>;
<xref ref-type="bibr" rid="R61">Gros, Blake, & Hiris, 1998</xref>)
as well as for translating plaids composed of a couple of gratings (<xref ref-type="bibr" rid="R68">Heeley & Buchanan-Smith, 1992</xref>).
These anisotropies have been observed either in foveal vision or in a visual
space at specific eccentricity (i.e., within a specific location in the visual
field).</p><p>Electrophysiological studies show that within an orientation column of the V1,
cells share a similar preferred orientation but they have diverse physiological
properties, one of the most dramatic being direction selectivity (cf. <xref ref-type="bibr" rid="R62">Gur, Kagan, & Snodderly, 2005</xref>;
<xref ref-type="bibr" rid="R74">Hubel & Wiesel, 1968</xref>). That
is, at any preferred orientation neurons that are direction selective in V1
respond more strongly to one direction of motion than to the opposite direction
(<xref ref-type="bibr" rid="R129">Pasternak, Schumer, Gizzi, &
Movshon, 1985</xref>; <xref ref-type="bibr" rid="R144">Reid, Soodak,
& Shapley, 1991</xref>). The signal is then dispatched via the near
extrastriate V2 and V3 to the far extratriate V5/MT (middle temporal) for a
further analysis (e.g., <xref ref-type="bibr" rid="R2">Albright, 1984</xref>;
<xref ref-type="bibr" rid="R17">Britten, Shadlen, Newsome, &
Movshon, 1992</xref>). Numerous studies have shown that neurons of similar
orientation or direction-of-motion selectivity are clustered into functional
columns in the MT (<xref ref-type="bibr" rid="R3">Albright, Desimone, &
Gross, 1984</xref>; <xref ref-type="bibr" rid="R46">Diogo, Soares, Koulakov,
Albright, & Gattass, 2003</xref>; <xref ref-type="bibr" rid="R106">Malonek, Tootell, & Grinvald, 1994</xref>). In addition, a very
large proportion of MT neurons are selective for the direction of motion and the
orientation of moving gratings (<xref ref-type="bibr" rid="R14">Born &
Bradley, 2005</xref>). Using the stimuli of moving gratings, a recent
optical imaging study of owl monkeys has demonstrated that more of the MT
cortical space is devoted to representing cardinal than oblique orientation
(<xref ref-type="bibr" rid="R178">Xu, Collins, Khaytin, Kaas, &
Casagrande, 2006</xref>), the anisotropy being more prominent in central
vision (≤ 10°). Furthermore, neural responses to cardinal
orientation were greater than neural responses to oblique orientation. It has
been claimed that this data explains why there is greater sensitivity to motion
discrimination when stimuli are moved along the cardinal meridians (polar axes),
suggesting that the motion oblique effect either originates in the MT or is
enhanced at this level (<xref ref-type="bibr" rid="R178">Xu et al.</xref>).</p></sec><sec id="S2C"><title>Asymmetry in spatial frequency processing</title><p>The primate V1 is dominated by complex cells that respond preferentially not only
to orientation and motion direction but also to the spatial frequency (SF) of
the stimuli (<xref ref-type="bibr" rid="R44">DeValois & DeValois,
1988</xref>). Human psychophysical studies suggest that there is a
continuous distribution of the SF preference in the visual cortex. For example,
observations in SF-specific adaptation (<xref ref-type="bibr" rid="R11">Blakemore & Campbell, 1969</xref>) and SF discrimination (<xref ref-type="bibr" rid="R174">Watson & Robson, 1981</xref>) tasks
provide compelling evidence that the visual cortex has multiple processing
channels, each tuned into one of many different SF ranges (<xref ref-type="bibr" rid="R154">Sachs, Nachmias, & Robson, 1971</xref>; <xref ref-type="bibr" rid="R173">Watson, 1982</xref>). In accord with this,
electrophysiological studies have shown that V1 neurons have a wide range of SF
preferences (<xref ref-type="bibr" rid="R43">DeValois, Albrecht, &
Thorell, 1982</xref>; <xref ref-type="bibr" rid="R166">Tolhurst &
Thompson, 1981</xref>) and neighbouring neurons are more likely to prefer
similar SFs (<xref ref-type="bibr" rid="R41">DeAngelis, Ghose, Ohzawa, &
Freeman, 1999</xref>; <xref ref-type="bibr" rid="R105">Maffei &
Fiorentini, 1977</xref>; <xref ref-type="bibr" rid="R167">Tolhurst &
Thompson, 1982</xref>).</p><p>SF preference also appears in later stages of visual processing, the degree of
preference being varied across the visual areas. For example, animal studies
have shown that SF preference is higher in V1 than in extratriate V2 (<xref ref-type="bibr" rid="R53">Foster, Gaska, Nagler, & Pollen,
1985</xref>; <xref ref-type="bibr" rid="R80">Issa, Trepel, &
Stryker, 2000</xref>) and V3 (<xref ref-type="bibr" rid="R56">Gegenfurtner,
Kiper, & Levitt, 1997</xref>). This fact has also been confirmed in
a recent fMRI study of humans (<xref ref-type="bibr" rid="R72">Henriksson,
Nurminen, Hyvärinen, & Vanni, 2008</xref>).</p><p>These studies have demonstrated the SF preference either in central vision or in
a visual space at specific eccentricity (i.e., within a specific location of the
visual field). However, such preferences are more pronounced in the central
vision and decrease with eccentricity.</p></sec></sec><sec id="S3"><title>THE <italic>WHAT</italic> AND <italic>PRIMARY WHY</italic> OF THE BETWEEN-VISUAL
FIELD ASYMMETRIES</title><sec id="S3A"><title>Foveal versus peripheral asymmetries</title><p>Perceptual capacity depends on where stimuli are located in the visual field.
Something we see out of the corner of our eye is blurred until we turn our eyes
to look directly at it. This is partly due to the sparse distribution of cones
in the periphery and partly due to the neural structures of the visual cortices.
That is, the density of the receptors in our visual system decreases as distance
from the fovea increases (e.g., <xref ref-type="bibr" rid="R37">Curcio, Sloan,
Kalina, & Hendrickson, 1990</xref>; <xref ref-type="bibr" rid="R38">Curcio, Sloan, Packer, Hendrickson, & Kalina, 1987</xref>). This
lack of uniformity carries through to lateral geniculate nucleus (<xref ref-type="bibr" rid="R29">Connolly & Van Essen, 1984</xref>) and
into visual cortices in both human (e.g., <xref ref-type="bibr" rid="R4">Anstis,
1998</xref>; <xref ref-type="bibr" rid="R140">Qiu et al., 2006</xref>; <xref ref-type="bibr" rid="R160">Sjöstrand, Olsson, Popovic, &
Conradi, 1999</xref>) and non-human primates (e.g., <xref ref-type="bibr" rid="R76">Hubel & Wiesel, 1974b</xref>; <xref ref-type="bibr" rid="R113">Maunsell & Van Essen, 1987</xref>; <xref ref-type="bibr" rid="R170">Van Essen, Newsome, & Maunsell, 1984</xref>).
Consequently, visual acuity (<xref ref-type="bibr" rid="R44">DeValois &
DeValois, 1988</xref>; <xref ref-type="bibr" rid="R47">Duncan &
Boynton, 2003</xref>), contrast sensitivity (<xref ref-type="bibr" rid="R172">Virsu & Rovamo, 1979</xref>), and colour
detection/discrimination (e.g., <xref ref-type="bibr" rid="R65">Hansen et al.,
2009</xref>; <xref ref-type="bibr" rid="R120">Mullen, 1991</xref>; <xref ref-type="bibr" rid="R121">Mullen & Kingdom, 1996</xref>; <xref ref-type="bibr" rid="R124">Newton & Eskew, 2003</xref>) fall
significantly towards the periphery. Many other visual functions such as object
and face identification (<xref ref-type="bibr" rid="R151">Rousselet, Thorpe,
& Fabre-Thorpe, 2004</xref>), stereopsis and reading are also
essentially limited to the central visual field (<xref ref-type="bibr" rid="R9">Battista, Kalloniatis, & Metha, 2005</xref>; <xref ref-type="bibr" rid="R180">Zegarra-Moran & Geiger 1993</xref>). Consequently, the
visual cortex’s early selective response towards stimuli such as
faces declines dramatically if presented a few degrees away from the fovea or
central fixation (<xref ref-type="bibr" rid="R49">Eimer, 2000</xref>; <xref ref-type="bibr" rid="R82">Jeffreys, Tukmachi, & Rockley,
1992</xref>).</p></sec><sec id="S3B"><title>Upper versus lower visual field asymmetries</title><p>Visual performance degrades in the periphery of the visual field, but not
proportionately in the lower and upper fields. Typically, the lower visual field
supports better performance than the upper visual field, even at the same
eccentricity (<xref ref-type="bibr" rid="R40">Danckert & Goodale,
2001</xref>; <xref ref-type="bibr" rid="R99">Levine & McAnany,
2005</xref>; <xref ref-type="bibr" rid="R114">McAnany & Levine,
2007</xref>). Psychophysical studies have demonstrated the dominance of the
lower field in temporal and contrast sensitivities (<xref ref-type="bibr" rid="R161">Skrandies, 1987</xref>), visual acuity (<xref ref-type="bibr" rid="R161">Skrandies, 1987</xref>), spatial resolution (<xref ref-type="bibr" rid="R145">Rezec & Dobkins, 2004</xref>), and in hue
(<xref ref-type="bibr" rid="R99">Levine & McAnany, 2005</xref>) and
motion (<xref ref-type="bibr" rid="R48">Edwards & Badcock, 1993</xref>;
<xref ref-type="bibr" rid="R95">Lakha & Humphreys, 2005</xref>;
<xref ref-type="bibr" rid="R99">Levine & McAnany, 2005</xref>; <xref ref-type="bibr" rid="R143">Raymond, 1994</xref>) processing. This phenomenon
is known as the vertical meridian asymmetry, which becomes more pronounced with
eccentricity (<xref ref-type="bibr" rid="R21">Carrasco et al., 2001</xref>) and
with increased spatial frequency. It is barely present for low spatial-frequency
Gabor stimuli, and gradually becomes more pronounced for intermediate and high
spatial frequencies (<xref ref-type="bibr" rid="R21">Carrasco et al.,
2001</xref>; <xref ref-type="bibr" rid="R103">Liu, Heeger, &
Carrasco, 2006</xref>; <xref ref-type="bibr" rid="R161">Skrandies,
1987</xref>). Many studies have also reported that the lower field
advantages may be restricted to the vertical meridian (polar axis that runs from
above the observer’s line of sight, through the fixation point, and
to below the observer’s line of sight) and may not be observed in
non-meridian locations (<xref ref-type="bibr" rid="R21">Carrasco et al.,
2001</xref>; <xref ref-type="bibr" rid="R103">Liu et al., 2006</xref>; <xref ref-type="bibr" rid="R165">Talgar & Carrasco, 2002</xref>).</p><p>Neurophysiological studies have confirmed the advantages of the lower visual
field’s sensitivity to contrast patterns (<xref ref-type="bibr" rid="R135">Portin, Vanni, Virsu, & Hari, 1999</xref>), high contrast
checkerboards (<xref ref-type="bibr" rid="R52">Fioretto et al., 1995</xref>),
non-attended colour changes (<xref ref-type="bibr" rid="R39">Czigler, Balazs,
& Pato, 2004</xref>), and motion (<xref ref-type="bibr" rid="R93">Kremláček, Kuba, Chlubnová, &
Kubová, 2004</xref>). In addition, this sort of measure has
revealed the advantages of the lower hemi-field over the upper hemi-field,
indicating that the asymmetry is not specific to the vertical meridian as
opposed to the psychophysical reports described above. Specifically, non-human
primate studies have shown that the cone and ganglion cell densities in the
retina are greater for the lower than for the upper visual field (<xref ref-type="bibr" rid="R130">Perry & Cowey, 1985</xref>). Slightly
more neural tissue is devoted to the lower than the upper visual field
representations in LGN (lateral geniculate nucleus; <xref ref-type="bibr" rid="R29">Connolly & Van Essen, 1984</xref>), V1 (<xref ref-type="bibr" rid="R170">Van Essen et al., 1984</xref>), and MT (<xref ref-type="bibr" rid="R113">Maunsell & Van Essen, 1987</xref>). Human
electrophysiological studies have also indicated functional specialization for
the lower and upper visual fields. For example, visual 100 ms evoked potential
peaks 11 to 12 ms earlier for lower visual field stimuli than for upper visual
field stimuli (<xref ref-type="bibr" rid="R98">Lehmann & Skrandies,
1979</xref>; <xref ref-type="bibr" rid="R161">Skrandies, 1987</xref>).
Similarly, MEG response amplitude has been reported to be greater for the lower
than the upper visual field in human observers (<xref ref-type="bibr" rid="R135">Portin et al., 1999</xref>). All this evidence for processing differences
and functional effects concerns eccentricities greater than around 5°
(<xref ref-type="bibr" rid="R135">Portin et al.</xref>). Thus, it is unclear
whether the processing of visual information differs between the lower and upper
visual fields near the fovea.</p></sec><sec id="S3C"><title>Left versus right visual field asymmetries</title><p>Perceptual processing in the left and right visual fields depends on the
spatiality of stimulus. Typically, spatial information is processed more
precisely in the left visual field and non-spatial information in the right
visual field (<xref ref-type="bibr" rid="R15">Boulinguez, Ferrois, &
Graumer, 2003</xref>; <xref ref-type="bibr" rid="R32">Corballis,
2003</xref>; <xref ref-type="bibr" rid="R33">Corballis, Funnell, &
Gazzaniga, 2002</xref>; <xref ref-type="bibr" rid="R127">Okubo &
Nicholls, 2008</xref>). Specifically, the left visual field is better at
simple line orientation, vernier offset and size discriminations (<xref ref-type="bibr" rid="R33">Corballis et al., 2002</xref>), and the right
visual field at temporal (<xref ref-type="bibr" rid="R127">Okubo &
Nicholls, 2008</xref>), linguistic and cognitive processing (<xref ref-type="bibr" rid="R32">Corballis, 2003</xref>). The superiority of the
left visual field is explained by the right hemisphere (RH) dominance over the
left hemisphere (LH) in spatial attention, as demonstrated in studies with
healthy individuals (<xref ref-type="bibr" rid="R69">Heilman & Van Den
Abell, 1979</xref>; <xref ref-type="bibr" rid="R163">Sturm, Reul, &
Willmes, 1989</xref>) as well as in unilateral lesion (<xref ref-type="bibr" rid="R112">Mattingley, Bradshaw, Nettleton, & Bradshaw,
1994</xref>), and neuroimaging studies (<xref ref-type="bibr" rid="R34">Corbetta, Miezin, Shulman, & Peterson, 1993</xref>). On the other
hand, presentation of verbal (i.e., non-spatial) stimuli produces
left-hemisphere activation, which triggers a rightward attentional bias and
results in a right visual field advantage (<xref ref-type="bibr" rid="R27">Cohen, 1982</xref>; <xref ref-type="bibr" rid="R87">Kinsbourne,
1970</xref>). Furthermore, an in-depth analysis of previous research reveals
that spatial processing in the left and right visual fields can be different in
many ways, as illustrated below.</p><sec id="S3C1"><title>High versus low spatial frequency processing</title><p>Visual analytic skill is critically determined by the stimulus spatial
frequency (SF), depending on its location in the visual field. The stimuli
with low SF are processed more efficiently in the left visual field and
those with high SF are processed more efficiently in the right visual field.
This asymmetric processing is directly associated with the functional
specialization of the RH and LH, which correspond to the left and right
visual fields, respectively. That is, the RH is dominant for low SF
processing whereas the LH is dominant for high SF processing. Evidence of
this hemispheric specificity has been provided by psychophysical and
behavioural studies using gratings of different spatial frequencies (<xref ref-type="bibr" rid="R26">Christman, Kitterle, & Hellige,
1991</xref>; <xref ref-type="bibr" rid="R88">Kitterle, Hellige,
& Christman, 1992</xref>; <xref ref-type="bibr" rid="R89">Kitterle & Selig, 1991</xref>) and natural pictures of low and
high spatial frequencies (<xref ref-type="bibr" rid="R132">Peyrin, Chauvin,
Chokron, & Marendaz, 2003</xref>; <xref ref-type="bibr" rid="R133">Peyrin et al., 2006</xref>). Recent functional brain imaging
studies conducted on healthy participants also support this pattern of
functional cerebral organization (<xref ref-type="bibr" rid="R131">Peyrin,
Baciu, Segebarth, & Marendaz, 2004</xref>; <xref ref-type="bibr" rid="R134">Peyrin et al., 2005</xref>).</p></sec><sec id="S3C2"><title>Global versus local processing</title><p>As global and local stimuli are typically conveyed by low SF and high SF,
respectively, global information is processed more efficiently in the left
visual field and local information in the right visual field (cf. <xref ref-type="bibr" rid="R59">Grabowska & Nowicka, 1996</xref>;
<xref ref-type="bibr" rid="R81">Ivry & Robertson, 1998</xref>;
<xref ref-type="bibr" rid="R157">Sergent, 1982</xref>). For example,
reaction time to a global target is faster than to a local target when
stimuli are presented in the left visual field, and vice versa when they are
presented in the right visual field (<xref ref-type="bibr" rid="R77">Hübner, 1997</xref>; <xref ref-type="bibr" rid="R86">Kimchi
& Merhav, 1991</xref>; <xref ref-type="bibr" rid="R157">Sergent,
1982</xref>). This global versus local processing asymmetry has been
confirmed in neuropsychological (lesion) studies (e.g., <xref ref-type="bibr" rid="R42">Delis, Robertson, & Efron,
1986</xref>; <xref ref-type="bibr" rid="R73">Hickok, Kirk, &
Bellugi, 1998</xref>; <xref ref-type="bibr" rid="R96">Lamb, Robertson,
& Knight, 1989</xref>), and in other behavioural studies with
healthy humans (e.g., <xref ref-type="bibr" rid="R13">Blanca, Zalabardo,
Gari-Criado, & Siles, 1994</xref>; <xref ref-type="bibr" rid="R78">Hübner, 1998</xref>; <xref ref-type="bibr" rid="R171">Versace & Tiberghien, 1988</xref>; <xref ref-type="bibr" rid="R179">Yovel, Yovel, & Levy, 2001</xref>).</p><p>In line with this, neuroimaging studies have shown that global and local
perception is mediated by separate subsystems in the RH and LH,
respectively. For example, ERPs (Event-related potentials) to compound
stimuli presented at the central fixation induce a larger occipito-temporal
negativity (<xref ref-type="bibr" rid="R70">Heinze, Johannes,
Münte, & Magun, 1994</xref>; <xref ref-type="bibr" rid="R155">Schatz & Erlandson, 2003</xref>) or target-specific
difference waves (<xref ref-type="bibr" rid="R63">Han, Liu, Yund, &
Woods, 2000</xref>; <xref ref-type="bibr" rid="R139">Proverbio, Minniti,
& Zani, 1998</xref>) over the RH in global stimulus condition,
but over the LH in local stimulus condition. Similarly, PET and fMRI studies
have shown increased regional cerebral blood flow or hemodynamic responses
in the right lateral occipital cortex when attending to the global structure
of compound stimuli, but in the left occipital cortex when attending to the
composing local elements (<xref ref-type="bibr" rid="R51">Fink et al.,
1996</xref>; <xref ref-type="bibr" rid="R64">Han et al., 2002</xref>;
<xref ref-type="bibr" rid="R110">Martinez et al., 1997</xref>).</p></sec><sec id="S3C3"><title>Coordinate versus categorical processing</title><p> Visual processing depends on how the stimulus elements are spatially related
in the visual display. Kosslyn (<xref ref-type="bibr" rid="R91">1987</xref>)
theorized that the visual system uses two types of spatial relations:
coordinate relations and categorical relations. Coordinate relations specify
precise spatial locations of objects or object parts in terms of metric
units and give exact distances. Categorical relations, in contrast, assign a
spatial configuration or a range of positions to an equivalence class (e.g.,
above/below, left/right, inside of/outside of) without defining the exact
metric properties. In the last couple of decades, scientists have reported
visual field asymmetry in processing these kinds of dual spatial relations.
In particular, categorical spatial processing is better in the right visual
field and coordinate spatial processing is better in the left visual field
(<xref ref-type="bibr" rid="R71">Hellige & Michimata,
1989</xref>; <xref ref-type="bibr" rid="R92">Kosslyn et al.,
1989</xref>). For example, Hellige and Michimata (<xref ref-type="bibr" rid="R71">1989</xref>) had participants judge if a dot was above or
below a line (a categorical task) or judge if the dot was within 2 cm of the
line (a coordinate task). A right visual field advantage was present for the
categorical task and a left visual field advantage was present for the
coordinate task. Kosslyn (<xref ref-type="bibr" rid="R91">1987</xref>) has
attributed this kind of fact directly to the functional specialization of
the two hemispheres. He proposed that the LH is preferentially associated
with the between-item categorical processing (e.g., one item is
“above” or “below” the other, a
discrete judgment) and the RH is preferentially associated with the
between-item coordinate processing (e.g., one item is
“near” to or “far” from the
other, an analog judgment). These distinct types of spatial processing may
also occur for the relative positions of parts or features of a single
stimulus item (<xref ref-type="bibr" rid="R162">Slotnick & Moo,
2006</xref>). </p><p> Kosslyn’s model, that there are two types of spatial
representations each with a specific lateralization pattern, has received
some support in different lines of behavioural research (<xref ref-type="bibr" rid="R8">Banich & Federmeier, 1999</xref>; <xref ref-type="bibr" rid="R94">Laeng & Peters, 1995</xref>; <xref ref-type="bibr" rid="R118">Michimata, 1997</xref>; <xref ref-type="bibr" rid="R125">Niebauer, 2001</xref>; <xref ref-type="bibr" rid="R126">Okubo
& Michimata, 2002</xref>; <xref ref-type="bibr" rid="R152">Rybash & Hoyer, 1992</xref>; <xref ref-type="bibr" rid="R177">Wilkinson & Donnelly, 1999</xref>). However, some attempts to
replicate the findings have failed (<xref ref-type="bibr" rid="R18">Bruyer,
Scailquin, & Coibion, 1997</xref>; <xref ref-type="bibr" rid="R158">Sergent, 1991a</xref>, <xref ref-type="bibr" rid="R159">1991b</xref>), though no studies have yet found the reverse pattern of
hemispheric dissociation. Procedural differences might explain this failure
of replication. For example, a study conducted by Bruyer et al. (<xref ref-type="bibr" rid="R18">1997</xref>) suggests that the hemispheric
dissociation for categorical and coordinate processing is highly unstable
and sensitive to subtle methodological factors, which could preclude its
general application. In that study, Kosslyn’s hypothesized
dissociation was observed in the manual requirement but not in the oral
response requirement, in the feedback but not in the no-feedback condition,
and in younger but not in elderly observers. Kosslyn’s hypothesis
should, therefore, be carefully tested employing similar stimuli and
procedures so that a firm conclusion about the existence of hemispheric
dissociation for coordinate and categorical processing can be arrived at. </p><p> Besides the two categories of perceptual asymmetries, there is evidence of
top-left lighting prior in 3D shape discrimination task that does not fall
in either of the categories. Half a century earlier, Gestalt psychologist
Metzger (<xref ref-type="bibr" rid="R117">1936</xref>) noticed that left-lit
scenes have greater perceptual value than right-lit ones. His observations
gave rise to an intriguing possibility: The visual system assumes that light
is coming from the left-above rather than straight-above. Sun and Perona
(<xref ref-type="bibr" rid="R164">1998</xref>) have tested this
proposition by asking observers to look for a convex or concave object lit
from one direction among similar objects lit from the opposite direction. In
this study, the shaded targets are detected more quickly when the
illumination position is between 30° and 60° to the left
of vertical. Both left- and right-handed participants show this tendency,
but it is more pronounced among the right-handed. This preference also
occurs in artists, participants across schools and periods of art history,
indicating its ecological significance. The top-left lighting preference has
been supported in a number of studies (e.g., <xref ref-type="bibr" rid="R57">Gerardin, de Montalembert, & Mamassian, 2007</xref>; <xref ref-type="bibr" rid="R108">Mamassian, Jentzsch, Bacon, &
Schweinberger, 2003</xref>), with the difference that it may not be
associated with handedness (<xref ref-type="bibr" rid="R107">Mamassian
& Goutcher, 2001</xref>). For example, in a recent study of
localization of an odd part of the Polo Mint stimulus Gerardin et al. (<xref ref-type="bibr" rid="R57">2007</xref>) found better performance for the
stimuli lit from the left than from the right. In another study of shape
from shading, Mamassian et al. (<xref ref-type="bibr" rid="R108">2003</xref>) detected a stronger top-left preference when the stimulus is
presented foveally rather than para-foveally. These authors have also
claimed that the N2 and P1 components in the visual occipital and temporal
areas might be responsible for the preference towards the leftward lighting
position, thus indicating a neural basis for the phenomenon. However, there
is still no evidence that this preference can be associated with hemispheric
specialization. Hence, the phenomenon is unspecified in this review. </p></sec></sec></sec><sec id="S4"><title>VISUAL EXPERIENCE: THE CRITICAL WHY OF PERCEPTUAL ASYMMETRIES</title><p>As discussed above, the asymmetric processing of visual information has either a
physiological or a neural basis. One of the most conspicuous functional properties
of neurons in the visual cortex is orientation selectivity, as more cortical
circuitry represents cardinal orientations rather than oblique orientations. The
development of this feature is primarily under endogenous control (e.g., <xref ref-type="bibr" rid="R25">Chapman, Stryker, & Bonhoeffer, 1996</xref>;
<xref ref-type="bibr" rid="R30">Coppola & White, 2004</xref>; <xref ref-type="bibr" rid="R176">Wiesel & Hubel, 1974</xref>), but can also be
altered by visual experience, sometimes with dramatic effects (e.g., <xref ref-type="bibr" rid="R12">Blakemore & Van Sluyters, 1975</xref>; <xref ref-type="bibr" rid="R36">Crair, Gillespie, & Stryker, 1998</xref>;
<xref ref-type="bibr" rid="R156">Sengpiel, Stawinski, & Bonhoeffer,
1999</xref>). Specifically, an early electrophysiological study of monkeys
(<xref ref-type="bibr" rid="R176">Wiesel & Hubel, 1974</xref>) and a
recent optical imaging study of ferrets (<xref ref-type="bibr" rid="R30">Coppola
& White, 2004</xref>) have demonstrated that overrepresentation of
cardinal orientations in the visual cortex does not require experience of an
anisotropic visual environment. Visual experience is not necessary for the initial
development of cortical orientation maps. Early maps are seen in ferrets’
visual cortices before natural eye opening (<xref ref-type="bibr" rid="R25">Chapman
et al., 1996</xref>), and normal orientation maps develop in kittens that have
been binocularly deprived for the first three weeks of their lives (<xref ref-type="bibr" rid="R36">Crair et al., 1998</xref>). However, longer periods of
binocular deprivation cause degradation of orientation preference maps (<xref ref-type="bibr" rid="R36">Crair et al., 1998</xref>), indicating that visual
experience is necessary for their maintenance. This is consistent with prior
electrophysiological (<xref ref-type="bibr" rid="R12">Blakemore & Van
Sluyters, 1975</xref>) and later optical imaging results (<xref ref-type="bibr" rid="R156">Sengpiel et al., 1999</xref>). The development of orientation
selectivity does not require visual experience, but is critically dependent on
spontaneous neuronal activity (<xref ref-type="bibr" rid="R24">Chapman,
Gödecke, & Bonhoeffer, 1999</xref>; <xref ref-type="bibr" rid="R30">Coppola & White, 2004</xref>). Absence of normal visual
experience can block spontaneous neural activity and hence orientation
selectivity.</p><p> Unlike orientation selectivity, the development of direction selectivity requires
visual experience. Li, Fitzpatrick, and White (<xref ref-type="bibr" rid="R100">2006</xref>) investigated the development of direction selectivity in
ferrets’ visual cortices using optical imaging and electrophysiological
techniques. In their study, direction selectivity was detected several days after
eye opening, this strengthened to adult levels over the following 2 weeks. Visual
experience was essential for this process, as shown by the absence of direction
selectivity in dark-reared ferrets. The impairment persisted in dark-reared ferrets
that were given experience of light after this period, despite the recovery of
orientation preference. Similarly, a recent study has shown that the visually
naïve ferrets’ visual cortices exhibited a well defined system
of orientation columns, but lacked the columnar pattern of direction selective
responses (<xref ref-type="bibr" rid="R101">Li, Hooser, Mazurek, White, &
Fitzpatrick, 2008</xref>). These results provide strong evidence that visual
experience increases the magnitude of direction selectivity, but there was no change
in orientation selectivity after training. The researchers concluded that early
experience with moving visual stimuli drives the rapid emergence of direction
selective responses in the visual cortex. Thus, visual experience is necessary for
the development of direction selectivity, as opposed to the development of
orientation selectivity. We suggest that the development of direction selectivity
and orientation selectivity are two independent processes of the visual system, the
former being experience-bound while the latter is not. In addition, direction
selective cortical neurons may be more susceptible or vulnerable to early visual
experience than orientation selective neurons. </p><p>Visual experience, or learning, also has a crucial role in modifying the response
properties of higher cortical neurons. Numerous neurophysiological studies provide
evidence for after-training enhanced stimulus selectivity. In particular, neurons in
monkey IT (inferior temporal) cortices show enhanced selectivity after training for
novel objects (<xref ref-type="bibr" rid="R90">Kobatake, Wang, & Tanaka,
1998</xref>; <xref ref-type="bibr" rid="R147">Rolls, 1995</xref>), holistic
multiple-part configurations (<xref ref-type="bibr" rid="R6">Baker, Behrmann,
& Olson, 2002</xref>), and even physically unrelated pairs of shapes
(<xref ref-type="bibr" rid="R116">Messinger, Squire, Zola, & Albright,
2005</xref>). The time required for response changes in some of these neurons
parallel the time required for learning (<xref ref-type="bibr" rid="R115">Messinger,
Squire, Zola, & Albright, 2001</xref>), suggesting a strong link between
underlying neuronal plasticity and behavioural improvement. Furthermore, learning
can shape the assignment of novel objects into classes (<xref ref-type="bibr" rid="R149">Rosenthal, Fusi, & Hochstein, 2001</xref>). This shaping is
done by modulating the selectivity of neurons in the inferior temporal and frontal
cortex for features crucial for the categorization process (<xref ref-type="bibr" rid="R54">Freedman, Riesenhuber, Poggio, & Miller, 2006</xref>). A
couple of studies have, however, reported that more PF (prefrontal) neurons (<xref ref-type="bibr" rid="R142">Rainer & Miller, 2000</xref>) or more V4
neurons (<xref ref-type="bibr" rid="R141">Rainer, Lee, & Logothetis,
2004</xref>) are selective towards repeated rather than novel stimuli at a
moderate level of image degradation, and at undegradation the effect was reverse or
equivalent. This indicates that stimulus selectivity is not a stable property of
cortical neurons; rather it is sensitive to the context of stimulation. We suggest
that response preference varies not only across the stimulated areas of the visual
cortex, but also within a particular area depending on the context of the
stimulation. However, in order to reach a firm conclusion regarding this trend the
hypothesis should be experimentally addressed in all other visual areas.</p><p>Psychophysical studies have also shown that visual experience modifies visual
response. For example, a 3D shape discrimination study has demonstrated that the
visual system’s prior knowledge or assumption about the direction of
lighting (i.e., “light-from-above” prior) can be modified by
visual-haptic training in humans (<xref ref-type="bibr" rid="R1">Adams, Graf,
& Ernst, 2004</xref>). This study has shown that training affects not
only subsequent shape perception of trained stimuli but also generalizes to affect
the perceived reflectance of novel stimuli. The effect has been successfully
replicated in a recent study on shape discrimination (<xref ref-type="bibr" rid="R23">Champion & Adams, 2007</xref>). In addition, Champion and
Adams have shown that convexity prior in visual search tasks (<xref ref-type="bibr" rid="R97">Langer & Bülthoff, 2001</xref>) can be reduced by
training. These findings suggest that cortical neurons learn where light-sources are
usually located, as well as the actual shape of the object, from interactions with
the environment, and use this information to interpret subsequent visual
stimuli.</p><p>In their recent studies Karim and Kojima (<xref ref-type="bibr" rid="R83">2010</xref>, <xref ref-type="bibr" rid="R84">in press</xref>) have shown that
vernier acuity improves as a function of training in both the cardinal and oblique
orientations. In addition, configurational asymmetry in vernier acuity reduces more
or less with training in the cardinal (<xref ref-type="bibr" rid="R83">Karim
& Kojima, 2010</xref>) but not in the oblique orientation (<xref ref-type="bibr" rid="R84">Karim & Kojima, in press</xref>). This
indicates a cardinal versus oblique orientation difference in sensitivity to
training. They interpreted this fact by the same mechanism of the oblique effect.
That is, a much lower percentage of V1 neurons are tuned to the oblique than to the
cardinal orientation (<xref ref-type="bibr" rid="R31">Coppola et al., 1998</xref>;
<xref ref-type="bibr" rid="R43">DeValois et al., 1982</xref>; <xref ref-type="bibr" rid="R55">Furmanski & Engel, 2000</xref>; <xref ref-type="bibr" rid="R102">Li et al., 2003</xref>). In addition, neurons with
the oblique preferences exhibit wider orientation tuning widths than neurons with
the cardinal preferences (<xref ref-type="bibr" rid="R85">Kennedy & Orban,
1979</xref>; <xref ref-type="bibr" rid="R123">Nelson, Kato, & Bishop,
1977</xref>; <xref ref-type="bibr" rid="R128">Orban & Kennedy,
1981</xref>; <xref ref-type="bibr" rid="R148">Rose & Blakemore,
1974</xref>). Thus, the asymmetry might reduce with training at a slower rate in
the oblique than in the cardinal orientation.</p><p>Taking all these results together, visual experience can modify or shape the response
properties of cortical neurons that contribute to perceptual asymmetries. Thus,
visual experience (or learning) can play a critical role (promoting or hindering) in
perceptual asymmetry.</p></sec><sec id="S5"><title>SOME UNRESOLVED AND CONTRADICTORY ISSUES</title><p>In spite of the successful demonstration of visual perceptual asymmetries in various
dimensions, there remain a number of contradictory and unresolved issues in the
scientific literature. As part of the above review, the cardinal superiority of
visual performance over the oblique orientation directly corresponds to the
asymmetry of neural organization in the primary visual cortex. But, it is still
unclear why visual acuity is finer for vertical stimuli than it is for horizontal
stimuli (<xref ref-type="bibr" rid="R153">Saarinen & Levi, 1995</xref>), a
demonstration opposed to the fact that more cells are devoted to horizontal than to
vertical orientation (<xref ref-type="bibr" rid="R102">Li et al., 2003</xref>). Some
scientists have tried to associate this demonstration with the everyday fact that we
experience more vertical than horizontal stimuli (<xref ref-type="bibr" rid="R60">Gregory, 1997</xref>). However, we wonder to what extent such normal visual
experience can alter the physiological preponderance and why such inconsistency does
not occur for some other visual tasks such as contrast sensitivity and spatial
resolution (<xref ref-type="bibr" rid="R21">Carrasco et al., 2001</xref>; <xref ref-type="bibr" rid="R146">Rijsdijk et al., 1980</xref>). That is, our daily
experience may not be responsible for altering typical orientation asymmetry. In
fact, the orientation detectors and many other feature detectors in the two
hemispheres are neither functionally equivalent nor are they absolutely exclusive or
independent. So, we propose that any consistency/inconsistency between our
orientation perception and cell representation can be determined by the
balanced/imbalanced functional interaction of the two hemispheres rather than by
normal visual experience.</p><p> A more important topic that requires scientific attention is that all the previous
studies of visual experience (see above) are concerned with how experience or
learning shapes the WVF asymmetries. No attempt has been made to explore whether
this factor can also accelerate or hinder the BVF asymmetries. For example, the
left-right visual field asymmetries have been associated with the functional
specializations of the corresponding hemispheres, but no attention has been paid to
whether experience can alter their specialized functions. In the case of the
upper-lower visual field asymmetries the lower field advantages have been
interpreted by finer attentional resolution in the lower than in the upper visual
field (<xref ref-type="bibr" rid="R66">He, Cavanagh, & Intriligator,
1996</xref>, <xref ref-type="bibr" rid="R67">1997</xref>; <xref ref-type="bibr" rid="R79">Intriligator & Cavanagh, 2001</xref>), but we still do not
know whether this increased attentional resolution is learned or innate. We suggest
that it might be learned, at least partly, because we usually look downward rather
than upward in our daily activities. However, past scientific studies cannot give a
satisfactory explanation of the superiority of the upper field over the lower field
in visual search (<xref ref-type="bibr" rid="R138">Previc & Blume,
1993</xref>), apparent size (<xref ref-type="bibr" rid="R150">Ross,
1997</xref>), and object recognition tasks (<xref ref-type="bibr" rid="R22">Chambers, McBeath, Schiano, & Metz, 1999</xref>) tasks. In fact, these
observations are contradictory to the demonstration that attentional resolution is
finer (<xref ref-type="bibr" rid="R66">He et al., 1996</xref>, <xref ref-type="bibr" rid="R67">1997</xref>; <xref ref-type="bibr" rid="R79">Intriligator &
Cavanagh, 2001</xref>) and neural representation is larger in the lower than in
the upper visual field (<xref ref-type="bibr" rid="R29">Connolly & Van
Essen, 1984</xref>; <xref ref-type="bibr" rid="R98">Lehmann & Skrandies,
1979</xref>; <xref ref-type="bibr" rid="R113">Maunsell & Van Essen,
1987</xref>; <xref ref-type="bibr" rid="R135">Portin et al., 1999</xref>; <xref ref-type="bibr" rid="R161">Skrandies, 1987</xref>; <xref ref-type="bibr" rid="R170">Van Essen et al., 1984</xref>). However, a theoretical account of the
disparity between upper and lower field dominance was proposed by Previc (<xref ref-type="bibr" rid="R136">1990</xref>), referring to the differences between
the two major streams of primate’s visual processing: the subcortical
(magnocellular/parvocellular) level (<xref ref-type="bibr" rid="R16">Breitmeyer,
1992</xref>) and cortical (dorsal/ventral organization) level (<xref ref-type="bibr" rid="R50">Ettlinger, 1990</xref>; <xref ref-type="bibr" rid="R58">Goodale & Milner, 1992</xref>; <xref ref-type="bibr" rid="R168">Ungerleider & Mishkin, 1982</xref>). Previc (<xref ref-type="bibr" rid="R137">1998</xref>) posited that the processing of stimuli
from lower and upper visual fields are promoted by these two neural systems, and
that they are related to the near (peripersonal) and far (extrapersonal) spaces,
respectively. According to this perspective, the specialization of the lower and
upper visual fields and their neural systems depends on the segregation of the near
and far spaces, which occurred during primate evolution. The lower visual field was
mainly involved in the perceptual processes required for visuomotor coordination in
the peripersonal space, largely performed by the dorsal pathways of the
primate’s visual system. And the upper visual field was linked to the
visual search and recognition mechanisms directed towards the extrapersonal space,
primarily controlled by the ventral system. However, Previc’s ideas are
based on his assumption about primate evolution and lacks empirical support. </p></sec><sec id="S6"><title>CONCLUDING REMARKS</title><p>This review demonstrates the wide range of perceptual variability or asymmetry both
within- and between-visual fields. The within-visual field asymmetries are typically
caused by neural preferences or asymmetric neural distribution in visual cortices.
However, silencing cortical activity or preventing visual experience may block the
typical development of the asymmetries. The foveal-peripheral asymmetries have been
attributed to the biased distribution of retinal cones and cortical neurons. The
upper-lower visual field asymmetries have been explained by asymmetric neural
distribution and attentional resolution, whereas the left-right visual field
asymmetries have been explained by stimulus driven attentional bias or by the
functional specialization of the two hemispheres. However, it remains unknown
whether visual experience can change the BVF asymmetries. Also, it has yet to be
investigated whether either hemisphere can independently determine the WVF
asymmetries. This would be very challenging because such an attempt would require
physiological isolation of the cerebral hemispheres. The two hemispheres of the
brain are actually designed to constantly communicate with one another and their
separation for experimental purpose may lead to functional abnormality or at least
some discrepancies between pre- and post-separation. However, we suggest that as
human brains are plastic, like the WVF asymmetries the BVF asymmetries can be
modified by visual experience. Empirical confirmation of this hypothesis would
enable scientists to alter the functional properties of the hemispheres in the
expected direction. This knowledge could be used for human welfare, particularly for
the brain-damaged patient population.</p></sec></body><back><ack><title>ACKNOWLEDGEMENTS</title><p>We sincerely thank the anonymous reviewers whose constructive comments helped us to
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