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Change of Reference Frame for Tactile Localization during Child Development

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Title: Change of Reference Frame for Tactile Localization during Child Development
Language: English
Authors: Pagel, Birthe, Heed, Tobias, Roder, Brigitte
Source: Developmental Science. Nov 2009 12(6):929-937.
Availability: Wiley-Blackwell. 350 Main Street, Malden, MA 02148. Tel: 800-835-6770; Tel: 781-388-8598; Fax: 781-388-8232; e-mail: cs-journals@wiley.com; Web site: http://www.wiley.com/WileyCDA/
Peer Reviewed: Y
Physical Description: PDF
Page Count: 9
Publication Date: 2009
Document Type: Journal Articles
Reports - Research
Descriptors: Stimuli, Child Development, Blindness, Cognitive Processes, Task Analysis, Age Differences, Tactual Perception, Psychomotor Skills
DOI: 10.1111/j.1467-7687.2009.00845.x
ISSN: 1363-755X
Abstract: Temporal order judgements (TOJ) for two tactile stimuli, one presented to the left and one to the right hand, are less precise when the hands are crossed over the midline than when the hands are uncrossed. This "crossed hand" effect has been considered as evidence for a remapping of tactile input into an external reference frame. Since late, but not early, blind individuals show such remapping, it has been hypothesized that the use of an external reference frame develops during childhood. Five- to 10-year-old children were therefore tested with the tactile TOJ task, both with uncrossed and crossed hands. Overall performance in the TOJ task improved with age. While children older than 5 1/2 years displayed a crossed hand effect, younger children did not. Therefore the use of an external reference frame for tactile, and possibly multisensory, localization seems to be acquired at age 5.
Abstractor: As Provided
Entry Date: 2009
Accession Number: EJ858859
Database: ERIC
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  Value: <anid>AN0044628646;5g501nov.09;2019May28.14:05;v2.2.500</anid> <title id="AN0044628646-1">Change of reference frame for tactile localization during child development. </title> <p>Temporal order judgements (TOJ) for two tactile stimuli, one presented to the left and one to the right hand, are less precise when the hands are crossed over the midline than when the hands are uncrossed. This 'crossed hand' effect has been considered as evidence for a remapping of tactile input into an external reference frame. Since late, but not early, blind individuals show such remapping, it has been hypothesized that the use of an external reference frame develops during childhood. Five‐ to 10‐year‐old children were therefore tested with the tactile TOJ task, both with uncrossed and crossed hands. Overall performance in the TOJ task improved with age. While children older than 5½ years displayed a crossed hand effect, younger children did not. Therefore the use of an external reference frame for tactile, and possibly multisensory, localization seems to be acquired at age 5.</p> <p>In order to bind the input of the different senses into a coherent percept, the brain must use stimulus features encoded by all sensory systems, such as the spatial location of an event, and the time point of its occurrence. For example, to avoid a mosquito bite, both the tactually and the visually perceived location of the mosquito and the location of the hand selected to perform a defensive action have to be matched. Thus, tactile and visual information about the position of the action goal (the mosquito) and proprioceptive and visual information about the position of the hand have to be combined.</p> <p>Temporal order judgement (TOJ) tasks have been successfully employed to investigate the reference frames involved in sensory localization. In this task, participants decide which of two stimuli was presented first, while the stimulus onset asynchrony (SOA) between the stimuli is systematically varied. The just noticeable difference (JND) is defined as the SOA at which the response of a participant is correct in 75% of the trials. When two tactile stimuli are presented one to each hand, the JND increases when the hands are crossed over the midline compared to an uncrossed hand posture ([<reflink idref="bib29" id="ref1">29</reflink>]; [<reflink idref="bib39" id="ref2">39</reflink>]). This is surprising, given that, theoretically, posture would be irrelevant when asked to decide if the right or the left hand had been touched first. It has been postulated that the crossed hand effect reflects a default remapping of tactile stimuli, initially coded in a somatotopic or anatomical coordinate system, into an external reference frame. This remapping has been hypothesized to take longer when the anatomical and external coordinates are in conflict, as is the case with crossed hands ([<reflink idref="bib39" id="ref3">39</reflink>]).</p> <p>Congenitally blind people do not show such spatial remapping effects ([<reflink idref="bib26" id="ref4">26</reflink>]; [<reflink idref="bib25" id="ref5">25</reflink>]; [<reflink idref="bib28" id="ref6">28</reflink>]). It has, therefore, been suggested that the default remapping of sensory input into an external coordinate system is not innate, but is acquired during ontogeny and induced by developmental vision. In contrast to the early blind, late blind people (blindness acquired after the age of 12) showed a similar crossing effect as the sighted, indicating that vision induces a default remapping of tactile input during early life which remains in use even if vision is lost later on. Moreover, a change from proprioceptive to a visual dominance in spatial perception has been reported in children between 6 and 19 years (Warren & Pick, 1970). While the impact of proprioception on sensory localization decreased with age in sighted children, such a trend was not observed in congenitally blind children.</p> <p>Animal studies further affirm the important role of vision for the development of multisensory integration. In monkeys, multisensory cells have been demonstrated to exist in the superior colliculus immediately after birth. Although these neurons responded to more than one modality, their capability to integrate sensory input based on spatial features was found to develop only within the first months of life ([<reflink idref="bib37" id="ref7">37</reflink>]; for a review see [<reflink idref="bib34" id="ref8">34</reflink>]). Furthermore, rearing cats in the dark resulted in a lack of multisensory integration even for auditory‐somatosensory stimuli ([<reflink idref="bib34" id="ref9">34</reflink>]; [<reflink idref="bib35" id="ref10">35</reflink>]).</p> <p>It has been hypothesized ([<reflink idref="bib26" id="ref11">26</reflink>]) that selective pruning and possibly the growth of new connections during brain development after birth ([<reflink idref="bib15" id="ref12">15</reflink>]; [<reflink idref="bib16" id="ref13">16</reflink>]) result in specific multisensory, for example visual‐tactile, connections, leading to a change in the default reference system for sensory localization based on spatial features.</p> <p>In sum, a number of studies point towards an essential influence of vision on tactile localization, and this influence seems to be established during ontogenic development. However, from the existing literature it is not yet clear when vision starts to dominate tactile localization, i.e. at what age a default remapping of tactile stimuli into an external reference frame occurs. The fact that crossing effects in the blind did not differ from those of the sighted if blindness occurred after the age of 12 ([<reflink idref="bib26" id="ref14">26</reflink>]) suggests that at that age, an external reference frame is already in use. The present study therefore tested children between the ages of 5 and 10 years with a tactile TOJ task both in a parallel and in acrossed hand posture. An effect of hand posture is expected only when the brain remaps tactile input into an external reference frame. If the use of such a remapping develops between the ages of 5 and 10 years, we expect a crossed hand effect in the older, but not in the younger children.</p> <hd id="AN0044628646-2">Materials and methods</hd> <p></p> <hd id="AN0044628646-3">Participants</hd> <p>Children were recruited in day care institutions in the city of Hamburg. Prior to the study, children's parents were asked to complete a questionnaire assessing the overall development of their child, including possible past and present illnesses and developmental disorders. Children with severe ophthalmic problems, very low birth weight, or a strong suspicion of having an attention deficit hyperactive disorder (ADHD) (Conners' Abbreviated Parent‐Teacher Questionnaire index ≥ 15) were excluded. Sixty‐five out of 69 volunteering children qualified for the present study. They had normal tactile discrimination abilities and normal or corrected to normal vision and were all free of neurological problems. Fourteen children were excluded from the analyses, because they had not completed a sufficient number of trials (<emph>n </emph>=<emph> </emph>5) or had not been able to follow the task instructions (<emph>n </emph>=<emph> </emph>9).</p> <p>Four age groups were defined post hoc based on initial results (detailed in the Results section): (<reflink idref="bib1" id="ref15">1</reflink>) Group <emph>5 early</emph> (14 children, 9 female, mean age 5:03 years, range 4:10–5:05, one left handed); (<reflink idref="bib2" id="ref16">2</reflink>) group <emph>5 late</emph> (14 children, 9 female, mean age 5:09 years, range 5:06–5:11, one left handed); (<reflink idref="bib3" id="ref17">3</reflink>) group <emph>6–7</emph> (10 children, 6 female, mean age 6:09 years, range 6:00–7:06, two ambidextrous); (<reflink idref="bib4" id="ref18">4</reflink>) group <emph>8–10</emph> (13 children, 9 female, mean age 9:01 years, range 8:00–10:04, all right handed). Parents or legal guardians gave written informed consent prior to the experiment. The participants received a small toy at the end of the experiment. The experiment was approved by the ethics committee of the medical council of Hamburg and was conducted in accordance with the Declaration of Helsinki (2004).</p> <hd id="AN0044628646-4">Apparatus and stimuli</hd> <p>The tactile stimuli consisted of metallic pins with a diameter of 0.8 mm, which were lifted by 0.35 mm from their resting position. The pins were controlled using the software Presentation (Neurobehavioral Systems, Inc., Albany, CA, USA) and presented for 10 ms to the distal phalanxes of the left and right index fingers with SOAs of −1500, −900, −600, −400, −300, −200, −90, −55, −30, −15, 15, 30, 55, 90, 200, 300, 400, 600, 900 and 1500 ms (negative values indicate that the first stimulus was presented to a participant's left hand) according to the method of constant stimuli. The tactile stimulators were positioned at a distance of 25 cm in front of the participant, and separated by 20 cm, centered on the midline. The experiment was conducted in a quiet room in the day care center. Verbal responses from the participants were recorded by the experimenter (BP).</p> <hd id="AN0044628646-5">Procedure and design</hd> <p>All participants took part in two sessions, each lasting between 1 and 1.5 hours. In the first session, an intelligence test (Coloured Progressive Matrices, CPM), a visual perception test (Motor‐Free Visual Perception Test, MVPT‐3) and a test of tactile sensitivity (Semmes‐Weinstein‐Monofilaments) were run. For tactile sensitivity, we used the adult norms which have been shown to be valid for children ([<reflink idref="bib33" id="ref19">33</reflink>]).</p> <p>Furthermore, to ensure that participants were able to perform the TOJ task, two pre‐tests were performed in the first session. (<reflink idref="bib1" id="ref20">1</reflink>) Tactile stimuli were randomly applied to just one hand either in an uncrossed or in a crossed hand posture (six targets each). Participants had to indicate verbally which hand was stimulated. In case of an error, the single‐target presentation was repeated up to three times. (<reflink idref="bib2" id="ref21">2</reflink>) Thereafter, participants performed 12 TOJ trials with uncrossed hands using the three longest SOAs only (<reflink idref="bib600" id="ref22">600</reflink>, 900, 1500 ms). The TOJ test was repeated up to three times when less than nine out of 12 answers were correct. All participants successfully completed at least one correct run of both pre‐tests.</p> <p>To avoid confusion in naming the right and left sides when indicating which hand had been stimulated first, the child's hands were labelled with a dog and a cat sticker, respectively. Participants gave a verbal response ('dog' or 'cat') according to the hand they had perceived as having been stimulated first.</p> <p>In the second session, one to 30 days after the first session, the children completed two blocks of 20 practice trials each for the TOJ task using 20 SOAs not used in the experiment proper (−3000 to 3000 ms). Finally, eight blocks of 40 test trials each were run with SOAs ranging between −1500 and 1500 ms (see above). Children performed a block of trials with the hands either crossed or uncrossed, with posture alternating every block. Half of the participants started with their hands in the uncrossed posture, the other half with their hands crossed. When the hands were crossed, the right arm was always crossed over the left arm to allow a comparison of our results to previous studies in adults (e.g. [<reflink idref="bib29" id="ref23">29</reflink>]; [<reflink idref="bib26" id="ref24">26</reflink>]; [<reflink idref="bib27" id="ref25">27</reflink>]; [<reflink idref="bib24" id="ref26">24</reflink>]). A towel was put between the arms for greater comfort. The faint clicking sounds produced by the tactile stimulators were masked with ambient white noise presented from a centrally located loudspeaker. In addition, participants wore noise protection headphones. Participants were instructed to maintain fixation throughout the trials on a centrally located small toy figure (7 cm high) positioned 25 cm straight ahead of them. Each trial started with a centrally presented warning sound (duration: 50 ms, pitch 900 Hz; 65 dB(A)). The first tactile stimulus was presented after 1300–1700 ms (average 1500 ms, rectangular distribution). Participants were informed that the task was unspeeded, and accuracy was stressed. They were encouraged to guess in case of uncertainty. A barking or meow sound was presented from a central loudspeaker according to the participant's verbal response. The next trial started with an inter‐trial interval of 1000 ms. The participant's response was entered by the experimenter into the computer.</p> <p>Thus, response times (RT) of the experimenter entering the responses of the participants in the computer were available indicating indirectly the response times of the children. These RTs were used to eliminate responses in which the children most likely were distracted or were not attending to the stimuli by eliminating trials with response times larger than three standard deviations of the participant's mean (about 2% of the trials for group <emph>5 early</emph>, 4% for group <emph>5 late</emph>, 2% for group <emph>6–7</emph> and 1% for group <emph>8–10</emph>).</p> <p>For each participant, the percentage of 'right first' responses was calculated separately for the two hand postures and for the 20 SOAs.</p> <p>To exclude children who gave random responses ([<reflink idref="bib4" id="ref27">4</reflink>]), we compared the performance in the 'left hand first' conditions (negative SOAs from 15 to 1500 ms) with the 'right hand first' conditions (positive SOAs from 15 to 1500 ms) for each participant in each posture with a paired‐sample <emph>t</emph>‐test. We assumed that responses were random when a child responded 'right first' and 'left first' equally often to all 'right hand first' stimuli, and/or responded 'right first' and 'left first' equally often to 'left hand first' stimuli. These tests were thus run separately for the 'right hand first' and the 'left hand first' trials. Children were excluded from further analysis when the percentage of 'right hand first responses' did not differ for the percentage 'left hand first responses' both when all SOAs (<emph>t</emph>(<reflink idref="bib9" id="ref28">9</reflink>) > −3, <emph>p </emph>> .1) were considered and when only the long SOAs (600–1500 ms; <emph>t</emph>(<reflink idref="bib2" id="ref29">2</reflink>) > −4, <emph>p </emph>> .1) were considered (this latter test excluded the short SOAs, because erroneous responses are expected for short SOAs by task design). Eight participants were excluded according to this criterion, six of them due to random performance in the crossed hand condition (<emph>5 early n </emph>=<emph> </emph>2; <emph>5 late n </emph>=<emph> </emph>2; <emph>6–7 n </emph>=<emph> </emph>1; <emph>8–10 n </emph>=<emph> </emph>1), and two due to random performance in both uncrossed and crossed conditions (<emph>5 early n </emph>=<emph> </emph>1; <emph>5 late n </emph>=<emph> </emph>1). Thus, only children who had understood the task instructions were included in the final sample. The reported results are based on the following group sizes: <emph>5 early</emph> (<emph>n </emph>=<emph> </emph>14), <emph>5 late</emph> (<emph>n </emph>=<emph> </emph>14), <emph>6–7</emph> (<emph>n </emph>=<emph> </emph>10) and <emph>8–10</emph> (<emph>n </emph>=<emph> </emph>13) (see Participants section, above).</p> <hd id="AN0044628646-6">Results</hd> <p>Performance was assessed by calculating the slope of linear regression lines of probit‐transformed response probabilities for each participant and each hand condition ([<reflink idref="bib29" id="ref30">29</reflink>]; [<reflink idref="bib9" id="ref31">9</reflink>]). Only SOAs up to that SOA at which perfect performance was achieved were used for slope calculation. In addition, JNDs (the SOA at which participants responded 75% correct) were calculated based on these linear regression lines ([<reflink idref="bib31" id="ref32">31</reflink>]) in order to allow for comparison with previous TOJ studies ([<reflink idref="bib29" id="ref33">29</reflink>]; [<reflink idref="bib39" id="ref34">39</reflink>]) (see Table 1; note that better performance is indicated by steeper, i.e. larger slopes, but lower JNDs).</p> <p>1  Just noticeable differences (JNDs) for tactile temporal order judgements in the uncrossed and the crossed hand postures for the four age groups (mean, standard error of the mean and median, one outliner per group removed, see text for details)</p> <p> <ephtml> <table><thead valign="bottom"><tr><th valign="bottom">Age group</th><th valign="bottom"><italic>n</italic></th><th>JND (ms) 
Uncrossed posture</th><th>JND (ms) 
Crossed posture</th></tr><tr><th>Mean</th><th>± std. err</th><th>(Median)</th><th>Mean</th><th>± std. err</th><th>(Median)</th></tr></thead><tbody valign="top"><tr><td>5 early</td><td>(4:10 – 5:05)</td><td>13</td><td>1265</td><td>230</td><td>(1181)</td><td>1425</td><td>692</td><td>(949)</td></tr><tr><td>5 late</td><td>(5:06 – 5:11)</td><td>13</td><td>544</td><td>108</td><td>(405)</td><td>1129</td><td>257</td><td>(748)</td></tr><tr><td>6–7</td><td>(6:00 – 7:06)</td><td>9</td><td>58</td><td>13</td><td>(52)</td><td>362</td><td>71</td><td>(316)</td></tr><tr><td>8–10</td><td>(8:00 – 10:04)</td><td>12</td><td>48</td><td>5</td><td>(53)</td><td>147</td><td>27</td><td>(150)</td></tr></tbody></table> </ephtml> </p> <p>The crossed hand effect is the difference of the slope or JND of the uncrossed minus the crossed hand posture conditions. These crossed hand effects for each participant are plotted against age in Figure 1. Visual inspection of Figure 1 reveals that a crossing effect does not seem to be evident before the age of 5½ years. To statistically analyze if these effects are significant, children were split into four age groups (5 early (4:10 to 5:6 years), 5 late (5:7 to 5:11 years), 6–7 years, and 8–10 years; see Materials and methods for age and gender information of each subgroup).1 All the following analyses therefore compare these four age groups. The percentage of 'right first' responses for each SOA for each participant as well as the group means are displayed in Figure 2.</p> <p>Graph: 1 Crossed hand effect of each participant (slopes of the linear regression lines for the uncrossed hand posture minus slopes of the linear regression lines for the crossed hand posture), plotted against age.</p> <p>Graph: 2 Psychophysical functions for the temporal order judgement task. The percentage of 'right‐first' responses (ordinates) of each participant is plotted against stimulus onset asynchronies (SOAs) (abscissae) separately for the four age groups (a–d). Negative SOAs denote 'left‐first' stimulation; positive SOAs denote 'right‐first' stimulation. Grey and red thin lines depict data from uncrossed and crossed conditions, respectively, for single participants. Each dot represents the probability of 'right‐first' responses based on 4–8 judgements. Thick‐lined curves represent group average data.</p> <hd id="AN0044628646-7">Slope analysis</hd> <p>The slopes of the linear regressions (see Figure 3) were entered in a repeated measure ANOVA with the within‐participant factor Posture (uncrossed vs. crossed) and the between‐participants factor Age (<emph>5 early</emph>, <emph>5 late</emph>, <emph>6–7</emph>, <emph>8–10</emph>).2 The main effect of Posture was significant (<emph>F</emph>(<reflink idref="bib1" id="ref35">1</reflink>, 47) = 27.72, <emph>p </emph>< .001, h<subs>p</subs><sups>2</sups> = .371), indicating that, overall, performance was better with uncrossed hands. Furthermore, TOJs improved with age, indicated by increasing slopes (see Figure 3) with increasing age (main effect of Age, <emph>F</emph>(<reflink idref="bib3" id="ref36">3</reflink>, 47) = 18.70 <emph>p </emph>< .001, h<subs>p</subs><sups>2</sups> = .544). Most importantly, however, the influence of posture, and thus the crossed hand effect, differed with age, as indicated by the highly significant interaction of Posture and Age (<emph>F</emph>(<reflink idref="bib3" id="ref37">3</reflink>, 47) = 7.97, <emph>p </emph>< .001, h<subs>p</subs><sups>2</sups> = .337).</p> <p>Graph: 3 Group mean slopes of the linear regression lines for the uncrossed (II, white bars) and the crossed (X, grey filled bars) hand posture depicted separately for the four age groups. Open and filled dots indicate the performance of single participants in the uncrossed and crossed hand conditions.</p> <p>A separate ANOVA for the uncrossed hand condition revealed a significant main effect of Age (uncrossed: <emph>F</emph>(<reflink idref="bib3" id="ref38">3</reflink>, 47) = 16.70, <emph>p </emph>< .001, h<subs>p</subs><sups>2</sups> = .516). All pair‐wise comparisons between groups (Games‐Howell procedure) were significant (<emph>p </emph>< .03), except those for adjacent age groups; the differences between <emph>5 early</emph> vs. <emph>5 late</emph> (<emph>p </emph>= .099) and <emph>5 late</emph> vs. group <emph>6–7</emph> (<emph>p </emph>= .054) were marginally significant; only the two oldest groups did not differ (group <emph>6–7</emph> vs. group <emph>8–10</emph>, <emph>p </emph>= .999).</p> <p>A separate ANOVA for the crossed hand posture condition revealed a significant main effect of Age (<emph>F</emph>(<reflink idref="bib3" id="ref39">3</reflink>, 47) = 6.99, <emph>p </emph>= . 001, h<subs>p</subs><sups>2</sups> = .308) as well, but only one of the pair‐wise comparisons reached significance (<emph>5 late</emph> vs. group <emph>8–10</emph>, <emph>p </emph>= .040, marginal significant <emph>5 early</emph> vs. group <emph>8–10</emph>, <emph>p </emph>= .067, all other comparisons <emph>p </emph>> .140).</p> <p>All groups, with the exception of group <emph>5 early,</emph> performed worse in the crossed than uncrossed hand condition (group <emph>5 early</emph>, <emph>t</emph>(<reflink idref="bib13" id="ref40">13</reflink>)= −1.09, <emph>p </emph>= .297; group <emph>5 late</emph>, <emph>t</emph>(<reflink idref="bib13" id="ref41">13</reflink>) = 2.67, <emph>p </emph>= .019; group <emph>6–7</emph>, <emph>t</emph>(<reflink idref="bib9" id="ref42">9</reflink>) = 3.33, <emph>p </emph>= .009; group <emph>8–10</emph>, <emph>t</emph>(<reflink idref="bib12" id="ref43">12</reflink>) = 2.98, <emph>p </emph>= .012).</p> <p>Inspection of Figure 2 suggests that children in the youngest group did not reach ceiling performance even at the largest SOAs. This may suggest that their responses are sometimes random (for example, due to lapses of attention). The difference between the 5 early and 5 late groups could then simply be due to the 5 late group responding randomly less often, and the absence of a crossed hand effect in the 5 early group may be due to the data being masked by random responses of those trials which were not correctly performed by the young children. We therefore added four random responses (amounting to 33% of noise) to the data of each 5 late child and repeated the analyses for this age group. The crossing effect was still significant (<emph>t</emph>(<reflink idref="bib13" id="ref44">13</reflink>) = 2.348, <emph>p </emph>= .035), indicating that the 5 early and 5 late groups performed differently even when one assumes that the 5 early group's responses are contaminated with noise.</p> <hd id="AN0044628646-8">JND analysis</hd> <p>JNDs are listed in Table 1. Because JNDs are inversely and non‐linearly related to slopes, the transformation of slopes into JNDs bears the risk of generating outliers ([<reflink idref="bib32" id="ref45">32</reflink>]; [<reflink idref="bib27" id="ref46">27</reflink>]). After eliminating outliers, a crossed hand effect was confirmed by separate <emph>t</emph>‐tests for all but the youngest age group (<emph>5 early, t</emph>(<reflink idref="bib12" id="ref47">12</reflink>) = −0.228, <emph>p </emph>= .823; all other groups, <emph>t </emph>> −3,09, <emph>p </emph>< .009).</p> <hd id="AN0044628646-9">Intelligence and visual development</hd> <p>The CPM and MVPT raw scores were converted into percentile ranks and normalized scores, respectively. Neither the slopes in the uncrossed (<emph>p </emph>= .962) and crossed (<emph>p </emph>= .576) conditions, nor the performance difference between the two hand conditions (slope difference = slope uncrossed minus slope crossed) (<emph>p </emph>= .721) were significantly correlated with general fluid intelligence, as measured with the CPM.</p> <p>Visual development, measured with the MVPT, and chronological age (in months) correlated significantly with slopes in the uncrossed and crossed hand conditions (uncrossed: MVPT <emph>r</emph>(<reflink idref="bib51" id="ref48">51</reflink>) = .58, <emph>p</emph> <.001; chronological age <emph>r</emph>(<reflink idref="bib51" id="ref49">51</reflink>) = .65, <emph>p</emph> =.001; crossed: MVPT <emph>r</emph>(<reflink idref="bib51" id="ref50">51</reflink>) = .40, <emph>p </emph>= .003; chronological age <emph>r</emph>(<reflink idref="bib51" id="ref51">51</reflink>) = .45, <emph>p</emph> =.001). However, when controlling for the effect of chronological age using a partial correlation analysis, slopes in the uncrossed and crossed postures did not correlate significantly with visual development (uncrossed: <emph>p </emph>= .319; crossed: <emph>p </emph>= .590).</p> <hd id="AN0044628646-10">Discussion</hd> <p>The aim of the present study was to determine if the use of an external reference frame for tactile localization is acquired during ontogeny in humans, and if so, then at what age. Children aged 5 to 10 years were tested in a TOJ task, in which they judged the order of two tactile stimuli, one presented to either hand, while adopting either an uncrossed or a crossed hand posture. Whereas children older than 5½ years performed more accurately with uncrossed than with crossed hands, hand posture did not affect task performance in children younger than 5½ years.</p> <p>As seen in Figure 1, the default use of an external reference frame for tactile localization is not a sudden step during development since not all older children show a crossing effect. However, for none of the younger children was a crossed hand effect observed. Thus external reference frames for tactile localization come into play after the age of 5½ years. Furthermore, each child was tested only once. Thus, no conclusions are possible about how the transition between the use of an anatomical and external reference frame, quickly or more gradually, takes place.</p> <p>Our study replicated an overall improvement in the TOJ task with age, as reported previously for auditory stimuli ([<reflink idref="bib22" id="ref52">22</reflink>]; [<reflink idref="bib4" id="ref53">4</reflink>]). Interestingly, performance in our study improved more in the uncrossed than in the crossed hand condition. The crossed hand effect in children older than 5½ years was mainly due to increasing accuracy in the uncrossed hand condition, whereas performance in the crossed hand condition did not improve to the same degree with age. Assuming that the crossed hand deficit in TOJ tasks reflects a conflict between an anatomical and an external reference frame, which seem to be both activated for tactile localization ([<reflink idref="bib17" id="ref54">17</reflink>]; [<reflink idref="bib39" id="ref55">39</reflink>]), the present results suggest that children under the age of 5½ years do not automatically remap tactile stimuli into an external reference frame. It may furthermore be speculated that the acquisition of an external reference frame primarily <emph>facilitates</emph> tactile localization under normal body postures, rather than causing confusion in a crossed posture, as has frequently been suggested (e.g. [<reflink idref="bib29" id="ref56">29</reflink>]; [<reflink idref="bib26" id="ref57">26</reflink>]).</p> <p>The transformation of spatial information from an anatomical (somatotopic) frame of reference into external coordinates requires continuous updating of spatial coordinates with every body movement. Studies in monkeys have demonstrated that neurons in the intraparietal sulcus encode stimuli from the different modalities in a number of reference frames ([<reflink idref="bib10" id="ref58">10</reflink>]; [<reflink idref="bib1" id="ref59">1</reflink>]; [<reflink idref="bib2" id="ref60">2</reflink>]).</p> <p>It has been hypothesized that these neurons are important for transforming modality specific coordinate systems into a reference frame that can be accessed by all sensory systems ([<reflink idref="bib1" id="ref61">1</reflink>]).</p> <p>In accordance with these reports in monkeys, both parietal and frontal brain regions have been shown to be involved in coordinate transformations between different sensory systems in healthy human adults ([<reflink idref="bib6" id="ref62">6</reflink>]). In addition, patients with parietal or frontal lobe damage often display deficits in spatial perception across modalities (e.g. neglect), suggesting that multisensory integration, and thus coordinate transformations, are mediated in homologous brain areas as in monkeys ([<reflink idref="bib18" id="ref63">18</reflink>]).</p> <p>Importantly, the maturation of parietal and frontal cortical areas continues until adulthood ([<reflink idref="bib3" id="ref64">3</reflink>]; [<reflink idref="bib30" id="ref65">30</reflink>]). It might therefore be hypothesized that selective pruning and growth of neural structures results in specific multisensory connections, which underlie the improvement of sensory integration from different modalities during ontogeny. A default use of an external (visual) reference frame for sensory localization, irrespective of stimulus modality, might emerge in the brain during this process.</p> <p>There is evidence that the cortex is also important for the development of multisensory integration in subcortical brain areas. It has been demonstrated that, although multisensory neurons exist at birth in the primate's superior colliculus (SC), specific integrational properties based on spatial features emerge only during the first months of life ([<reflink idref="bib37" id="ref66">37</reflink>]), and that this development is gated by the development of cortical influences ([<reflink idref="bib36" id="ref67">36</reflink>]). Therefore, brain regions capable of integrating input across sensory systems in adult individuals do not seem to become fully functional before higher cortical regions have reached a sufficient level of maturation. In accordance with such findings in animals, it has been reported that, compared to adults, multisensory capabilities in infants are limited and only partially developed ([<reflink idref="bib20" id="ref68">20</reflink>]).</p> <p>The brain preferentially represents visual spatial information in the hemisphere contralateral to the visual hemifield; a change of eye position or a change of stimulus position to another hemifield then implies a transfer of spatial information across the hemispheres ([<reflink idref="bib13" id="ref69">13</reflink>]; [<reflink idref="bib21" id="ref70">21</reflink>]). In the same way, multisensory integration seems to involve interhemispheric transfer. A split brain patient, whose brain cannot relay information between the hemispheres, showed normal interference from visual stimuli in a tactile localization task when the hands were uncrossed (i.e. visual and tactile information was relayed to the same hemisphere of the brain), but (in contrast to healthy participants) not when the hands were crossed (i.e. when visual and tactile information initially arrived in different hemispheres), indicating that a hemispheric transfer is required for multisensory interference to occur ([<reflink idref="bib31" id="ref71">31</reflink>]). Interestingly, the main phase of corpus callosum maturation is between the ages of 4 and 18 years ([<reflink idref="bib19" id="ref72">19</reflink>]). In line bisection tasks, children younger than 6–7 years bisect horizontal lines slightly to the left of their midpoint when using the left hand, and to the right when using the right hand ([<reflink idref="bib7" id="ref73">7</reflink>]). Adults, in contrast, bisect lines with a slight shift to the left of their midpoint, independent of the hand used. The general leftward bias in adults has been attributed to the dominance of the right hemisphere in spatial attention. When the right hand is used, communication is required between the motor cortex in the left hemisphere and the attention system in the right hemisphere. Thus, the symmetrical bisection bias shown by young children might be related to the incomplete functional maturation of callosal transfer ([<reflink idref="bib11" id="ref74">11</reflink>]). The adults' general leftward bias has been reported to emerge around the age of 7 or 8 years ([<reflink idref="bib5" id="ref75">5</reflink>]) or even later, at the age of 13 years ([<reflink idref="bib12" id="ref76">12</reflink>]).</p> <p>Changes in hemispheric transfer with age have also been demonstrated in tactile processing as well. [<reflink idref="bib23" id="ref77">23</reflink>] studied children of ages 5, 7, 9, and 11 years. When asked to indicate with the thumb which finger had been touched, children of all age groups performed better when they used the thumb of the same hand than when they used the thumb of the opposite hand. However, performance depended on hand posture, and the effect of hand posture differed with age. Five‐year‐old children performed better when both the tested and the responding hand were held palm down than when the stimulated hand was palm down, while the responding hand was turned palm up. In contrast, children of age 9–11 years performed better with the stimulated hand down and the responding palm up. That is, only the older children performed better when the position of the fingers of both hands matched in external space. Importantly, when the 5‐year‐old children were tested again 4 months later, their performance had improved in the second condition, i.e. the one that required the use of external coordinates and most likely callosal transfer. This study, therefore, is in agreement with our results suggesting the increasing importance of external reference frames for sensory localization in the second half of age 5.</p> <p>These findings together suggest that the maturation of the corpus callosum and the increased efficiency of callosal transfer may contribute to the improvement of the spatial remapping capabilities required for the use of an external reference frame even in the TOJ task used in the present study.</p> <p>In sum, the present study provides evidence that children start using an external reference frame for the localization of tactile stimuli after about 5½ years of age. The use of this default remapping seems to facilitate tactile localization and might contribute to the increase of the capacities for multisensory action control during child development.</p> <hd id="AN0044628646-11">Acknowledgements</hd> <p>For their support and cooperation, we thank the directors, parents, and children of the kindergardens Akademie für Kinder, Albertinen‐Kindertagesstätte, Kita Antje, Kita Christianskirche, Betriebskindergarten Hamburg‐Eppendorf, Kita Flughafenstrasse, Kita Hummelsbüttel, Kita Monetastraße, Praxis Ausbildungs‐ und Kindertagesstätte Niendorf and Kita Wernigeroder Weg. BP received a graduate student fellowship of the University of Hamburg. The study was funded by the German Research Foundation (DFG).</p> <ref id="AN0044628646-12"> <title> Footnotes </title> <blist> <bibl id="bib1" idref="ref15" type="bt">1</bibl> <bibtext>  The result pattern of a crossing effect becoming evident only after the age of 5½ was already visible after an initial analysis of 16 children for the whole group of 5‐year‐olds. We therefore recruited more children between 5 and 6 years to arrive at the now reported group sizes of 14 for both the 5 early and the 5 late groups to substantiate this observation.</bibtext> </blist> <blist> <bibl id="bib2" idref="ref16" type="bt">2</bibl> <bibtext>  Because the statistical assumptions of normal distribution and variance homogeneity cannot be assumed for small sample sizes, we additionally ran analogous analyses with non‐parametric tests. The overall result pattern obtained was similar to the parametric analyses. Age groups were compared with a Kruskal‐Wallis test for the uncrossed posture (chi‐square = 35.3 with 3 <emph>df</emph>, <emph>p</emph><.001) and crossed posture (chi‐square = 23.7 with 3 <emph>df</emph>, <emph>p</emph><.001). The differences between each pair of age groups were assessed with Mann‐Whitney U‐tests: in the uncrossed hand posture, significant differences were found between all groups (<emph>p</emph><.01) except between groups 6–7 and 8–10 (<emph>p</emph> = .5435). In the crossed posture, significant differences were revealed between all groups (<emph>p</emph><.05) except between 5 early vs. 5 late (<emph>p</emph> = .383). The difference between the uncrossed and the crossed condition for each age group was assessed with a Wilcoxon Signed Ranks test: performance was significantly better in the uncrossed than in the crossed hand condition (all <emph>p</emph>s<.02) for all groups except for group 5 early (<emph>p</emph> = .650).</bibtext> </blist> <blist> <bibl id="bib3" idref="ref17" type="bt">3</bibl> <bibtext> Tobias Heed has previously published under the name Tobias Schicke.</bibtext> </blist> </ref> <ref id="AN0044628646-13"> <title> References </title> <blist> <bibtext> Avillac, M., Deneve, S., Olivier, E., Pouget, A., & Duhamel, J.R. (2005). Reference frames for representing visual and tactile locations in parietal cortex. Nature Neuroscience, 8, 941 – 949.</bibtext> </blist> <blist> <bibtext> Avillac, M., Olivier, E., Denève, S., Hamed, S.B., & Duhamel, J.‐R. (2004). Multisensory integration in multiple reference frames in the posterior parietal cortex. 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Nature Neuroscience, 4, 759 – 765.</bibtext> </blist> </ref> <aug> <p>By Birthe Pagel; Tobias Heed and Brigitte Röder</p> <p>Reported by Author; Author; Author</p> </aug> <nolink nlid="nl1" bibid="bib29" firstref="ref1"></nolink> <nolink nlid="nl2" bibid="bib39" firstref="ref2"></nolink> <nolink nlid="nl3" bibid="bib26" firstref="ref4"></nolink> <nolink nlid="nl4" bibid="bib25" firstref="ref5"></nolink> <nolink nlid="nl5" bibid="bib28" firstref="ref6"></nolink> <nolink nlid="nl6" bibid="bib37" firstref="ref7"></nolink> <nolink nlid="nl7" bibid="bib34" firstref="ref8"></nolink> <nolink nlid="nl8" bibid="bib35" firstref="ref10"></nolink> <nolink nlid="nl9" bibid="bib15" firstref="ref12"></nolink> <nolink nlid="nl10" bibid="bib16" firstref="ref13"></nolink> <nolink nlid="nl11" bibid="bib33" firstref="ref19"></nolink> <nolink nlid="nl12" bibid="bib600" firstref="ref22"></nolink> <nolink nlid="nl13" bibid="bib27" firstref="ref25"></nolink> <nolink nlid="nl14" bibid="bib24" firstref="ref26"></nolink> <nolink nlid="nl15" bibid="bib31" firstref="ref32"></nolink> <nolink nlid="nl16" bibid="bib13" firstref="ref40"></nolink> <nolink nlid="nl17" bibid="bib12" firstref="ref43"></nolink> <nolink nlid="nl18" bibid="bib32" firstref="ref45"></nolink> <nolink nlid="nl19" bibid="bib51" firstref="ref48"></nolink> <nolink nlid="nl20" bibid="bib22" firstref="ref52"></nolink> <nolink nlid="nl21" bibid="bib17" firstref="ref54"></nolink> <nolink nlid="nl22" bibid="bib10" firstref="ref58"></nolink> <nolink nlid="nl23" bibid="bib18" firstref="ref63"></nolink> <nolink nlid="nl24" bibid="bib30" firstref="ref65"></nolink> <nolink nlid="nl25" bibid="bib36" firstref="ref67"></nolink> <nolink nlid="nl26" bibid="bib20" firstref="ref68"></nolink> <nolink nlid="nl27" bibid="bib21" firstref="ref70"></nolink> <nolink nlid="nl28" bibid="bib19" firstref="ref72"></nolink> <nolink nlid="nl29" bibid="bib11" firstref="ref74"></nolink> <nolink nlid="nl30" bibid="bib23" firstref="ref77"></nolink>
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  Data: Change of Reference Frame for Tactile Localization during Child Development
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  Data: <searchLink fieldCode="AR" term="%22Pagel%2C+Birthe%22">Pagel, Birthe</searchLink><br /><searchLink fieldCode="AR" term="%22Heed%2C+Tobias%22">Heed, Tobias</searchLink><br /><searchLink fieldCode="AR" term="%22Roder%2C+Brigitte%22">Roder, Brigitte</searchLink>
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  Data: <searchLink fieldCode="SO" term="%22Developmental+Science%22"><i>Developmental Science</i></searchLink>. Nov 2009 12(6):929-937.
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  Data: Wiley-Blackwell. 350 Main Street, Malden, MA 02148. Tel: 800-835-6770; Tel: 781-388-8598; Fax: 781-388-8232; e-mail: cs-journals@wiley.com; Web site: http://www.wiley.com/WileyCDA/
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  Data: 9
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  Data: <searchLink fieldCode="DE" term="%22Stimuli%22">Stimuli</searchLink><br /><searchLink fieldCode="DE" term="%22Child+Development%22">Child Development</searchLink><br /><searchLink fieldCode="DE" term="%22Blindness%22">Blindness</searchLink><br /><searchLink fieldCode="DE" term="%22Cognitive+Processes%22">Cognitive Processes</searchLink><br /><searchLink fieldCode="DE" term="%22Task+Analysis%22">Task Analysis</searchLink><br /><searchLink fieldCode="DE" term="%22Age+Differences%22">Age Differences</searchLink><br /><searchLink fieldCode="DE" term="%22Tactual+Perception%22">Tactual Perception</searchLink><br /><searchLink fieldCode="DE" term="%22Psychomotor+Skills%22">Psychomotor Skills</searchLink>
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  Data: 10.1111/j.1467-7687.2009.00845.x
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  Data: 1363-755X
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  Data: Temporal order judgements (TOJ) for two tactile stimuli, one presented to the left and one to the right hand, are less precise when the hands are crossed over the midline than when the hands are uncrossed. This "crossed hand" effect has been considered as evidence for a remapping of tactile input into an external reference frame. Since late, but not early, blind individuals show such remapping, it has been hypothesized that the use of an external reference frame develops during childhood. Five- to 10-year-old children were therefore tested with the tactile TOJ task, both with uncrossed and crossed hands. Overall performance in the TOJ task improved with age. While children older than 5 1/2 years displayed a crossed hand effect, younger children did not. Therefore the use of an external reference frame for tactile, and possibly multisensory, localization seems to be acquired at age 5.
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        Value: 10.1111/j.1467-7687.2009.00845.x
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      – SubjectFull: Stimuli
        Type: general
      – SubjectFull: Child Development
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      – SubjectFull: Blindness
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      – SubjectFull: Cognitive Processes
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      – SubjectFull: Task Analysis
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      – SubjectFull: Age Differences
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      – SubjectFull: Tactual Perception
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      – SubjectFull: Psychomotor Skills
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    Titles:
      – TitleFull: Change of Reference Frame for Tactile Localization during Child Development
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  BibRelationships:
    HasContributorRelationships:
      – PersonEntity:
          Name:
            NameFull: Pagel, Birthe
      – PersonEntity:
          Name:
            NameFull: Heed, Tobias
      – PersonEntity:
          Name:
            NameFull: Roder, Brigitte
    IsPartOfRelationships:
      – BibEntity:
          Dates:
            – D: 01
              M: 11
              Type: published
              Y: 2009
          Identifiers:
            – Type: issn-print
              Value: 1363-755X
          Numbering:
            – Type: volume
              Value: 12
            – Type: issue
              Value: 6
          Titles:
            – TitleFull: Developmental Science
              Type: main
ResultId 1