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Re: Experimental Brain Research DOI:10.1007/s00221-007-1148-0
Shift of manual preference by lateralized practice generalizes to related motor tasks
Authors: LuisAugusto Teixeira · Victor Okazaki

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Journal: Experimental Brain Research
10.1007/s00221-007-1148-0

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LuisAugusto Teixeira LuisAugusto Teixeira
Department of Biodynamics Department of Biodynamics
University of Sao Paulo University of Sao Paulo
Sao Paulo, , Brazil Sao Paulo, , Brazil

q q
Metadata of the article that will be visualized in OnlineFirst
ArticleTitle Shift of manual preference by lateralized practice generalizes to related motor tasks
Article Sub-Title
Journal Name Experimental Brain Research
Corresponding Author Family Name Teixeira
Particle
Given Name Luis Augusto
Suffix
Division Department of Biodynamics
Organization University of Sao Paulo
Address Sao Paulo, Brazil
Email lateixei@usp.br

Author Family Name Okazaki

Particle
Given Name Victor
Suffix
Division Department of Biodynamics
Organization University of Sao Paulo
Address Sao Paulo, Brazil
Email

Received 10 July 2007

Schedule Revised
Accepted 14 September 2007
Abstract Previous investigation (Teixeira and Teixeira in Brain Cogn, in press, 2007) has evidenced a persistent shift
of manual preference for a particular motor task following lateralized practice. In the present study, we
assessed the extent to which shift of manual preference is generalizable to related motor tasks. Twenty right-
handers were assigned to an experimental or to a control group. The former were provided with practice on
a particular sequence of finger movements with their left hand only, while the latter remained inactive.
Participants were assessed on manual asymmetry, indexed by movement time, and manual preference for the
practiced and for other two sequences of finger movements (transfer tasks). Assessment was made before,
immediately after, and 30 days following (retention) practice sessions. Results showed that lateralized practice
led to significant bilateral reduction of movement time, maintaining the symmetric performance observed
before practice following task acquisition. Regarding manual preference, before task acquisition, all
participants in the experimental group were right-handed for the main task; immediately after practice their
predominant manual preference shifted to the left hand, a profile that was maintained in retention. This
persistent shift of manual preference was also observed for one of the transfer tasks requiring the same
sequence of transitions between finger movements. Indices of correlation between manual asymmetry and
manual preference were non-significant across tasks and phases, suggesting that manual preference was not
defined by lateral asymmetry of performance. We propose that manual preference is established by automatic
sensorimotor processing and/or increased confidence on a single hand from previous experiences.
Keywords (separated by '-') Handedness - Lateralization - Manual asymmetry - Generalization - Motor training
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 Exp Brain Res
DOI 10.1007/s00221-007-1148-0

1 R ES EA R C H N O T E

2 Shift of manual preference by lateralized practice generalizes

3 to related motor tasks
4 Luis Augusto Teixeira · Victor Okazaki

F
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Author Proof

5 Received: 10 July 2007 / Accepted: 14 September 2007

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6  Springer-Verlag 2007

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7 Abstract Previous investigation (Teixeira and Teixeira in preference is established by automatic sensorimotor pro- 36
8 Brain Cogn, in press, 2007) has evidenced a persistent shift cessing and/or increased conWdence on a single hand from 37
9 of manual preference for a particular motor task following previous experiences. 38
10 lateralized practice. In the present study, we assessed the
11
12
extent to which shift of manual preference is generalizable
to related motor tasks. Twenty right-handers were assigned D
Keywords Handedness · Lateralization ·
Manual asymmetry · Generalization · Motor training
39
40
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13 to an experimental or to a control group. The former were
14 provided with practice on a particular sequence of Wnger
15 movements with their left hand only, while the latter Introduction 41
16 remained inactive. Participants were assessed on manual
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17 asymmetry, indexed by movement time, and manual prefer- Lateralization of human motor function has been proposed 42
18 ence for the practiced and for other two sequences of Wnger as a dynamic and multifaceted process contingent upon 43
19 movements (transfer tasks). Assessment was made before, individual related determinants (Serrien et al. 2006). One 44
20 immediately after, and 30 days following (retention) prac- such individual determinant of human lateralization has 45
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21 tice sessions. Results showed that lateralized practice led to been shown to be systematic motor experiences using a sin- 46
22 signiWcant bilateral reduction of movement time, maintain- gle limb, heretofore named lateralized practice (LP). 47
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23 ing the symmetric performance observed before practice Research has indicated that LP has a role in shifting manual 48
24 following task acquisition. Regarding manual preference, preference (preference of a single hand to perform motor 49
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25 before task acquisition, all participants in the experimental tasks). Evidence has been presented that as right-handed 50
26 group were right-handed for the main task; immediately children (Singh et al. 2001) and adults (Teixeira 2007) 51
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27 after practice their predominant manual preference shifted advance in age, their preference for the right hand becomes 52
28 to the left hand, a proWle that was maintained in retention. stronger. Furthermore, left-handed children are frequently 53
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29 This persistent shift of manual preference was also enforced to shift their manual preference to the right hand 54
30 observed for one of the transfer tasks requiring the same (Zverev 2006), and in many cases this environmental pres- 55
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31 sequence of transitions between Wnger movements. Indices sure is successful. Meng (2007) reported that in a large 56
32 of correlation between manual asymmetry and manual group of left-handed children, social eVort to shift manual 57
33 preference were non-signiWcant across tasks and phases, preference produced the desired eVect in approximately 58
34 suggesting that manual preference was not deWned by lat- 60% of the cases (see also Porac et al. 1986). Both strength- 59
35 eral asymmetry of performance. We propose that manual ening of manual preference in right-handers and shift of 60
manual preference in left-handers seem to be due to exten- 61
sive lateralized motor experiences in diVerent activities, 62
modulating previous tendencies of use of the hands. In 63
A1 L. A. Teixeira (&) · V. Okazaki
agreement with this interpretation, Mikheev and colleagues 64
A2 Department of Biodynamics,
A3 University of Sao Paulo, Sao Paulo, Brazil (2002) showed that judo athletes prefer more frequently 65
A4 e-mail: lateixei@usp.br than other individuals to perform certain movements with 66

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67 their left hand, although overall right-handed. This observa- enhance its chance of been chosen to perform that task in 115
68 tion suggests that learning is able to shift manual preference future situations. If this proposition is correct, shift of man- 116
69 for particular motor tasks in right-handers also, who usually ual preference following LP for a particular motor task 117
70 are more consistent in the use of the preferred hand than should generalize to motor tasks requiring similar move- 118
71 left-handers (Healey et al. 1986). ments, independent of performance asymmetries between 119
72 Direct evidence from controlled experimental investiga- the hands. This hypothesis was scrutinized in the present 120
73 tion of the eVect of LP on the establishment of lateral pref- study by assessing immediate and enduring eVects of prac- 121
74 erence in right-handers has been presented recently. tice with the nonpreferred left hand of sequential Wnger 122
75 Teixeira and Teixeira (2007) evaluated the eVect of LP of movements on manual asymmetry and manual preference. 123
76 sequential Wnger movements of the nonpreferred left hand This analysis was conducted for the practiced task and for 124
77 on manual asymmetry and preference. The results revealed variations of it, focusing on the relationship between prefer- 125
78 a noticeable shift of manual preference for the experimental ence and performance. 126

F
79 task following practice, with some participants shifting

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80 from the original consistent right hand preference to consis-
Author Proof

81 tent left hand preference. That was an enduring eVect, since Materials and methods 127

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82 1 month following the end of practice, shift of manual pref-
83 erence for the experimental task was still observable (cf. Participants 128
84 McGonigle and Flook 1978, for analogous Wndings in ani-

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85 mals). Additional aspects in these results deserve further Thirteen male and seven female university students (age 129
86 attention. First, following practice of the nonpreferred left range 17–46 years, M = 23.45, SD = 7.92) participated in 130
87 hand, no signiWcant diVerence between performance of the the study. All participants were right-handers, as indicated 131
88 right and the left hand (manual asymmetry) was observed. by the Edinburgh manual dominance inventory (OldWeld 132
89
90
Thus, it was shown that the symmetric performance
between the hands, as usually observed in complex sequen- D 1971), with median score equal to 4.5 on a 5-point scale.
Ethical approval for this study was granted by the Institu-
133
134
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91 tial Wnger movements1 in adults (Teixeira and Paroli 2000; tional Review Board and all procedures were in accordance 135
92 Hausmann et al. 2004) and children (Denckla 1974; Fagard with the standards established in the Declaration of Hel- 136
93 1987), is not signiWcantly changed by the amount of LP sinki. 137
94 employed in the study. Second, the results pointed out
EC

95 incongruence between manual asymmetry and manual pref- Task and equipment 138
96 erence. This point was detected in the group analysis, with
97 no signiWcant correlation between magnitude of manual The main experimental motor task consisted of sequentially 139
98 asymmetry and strength of manual preference, and in the touching the thumb with the other four Wngers in the fol- 140
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99 large number of individual cases of divergence between lowing order: index, ring, middle, and little Wnger. One trial 141
100 manual preference and manual asymmetry. These results consisted of performing this sequence (cycle) of Wnger 142
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101 are contradictory to the assumption that manual preference movements three times without interruption. The aim of the 143
102 for a given task is a function of the relative proWciency task was to complete a trial in the shortest period of time. 144
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103 between the right and the left hand (Bishop 1989), and sug- Two other sequences of Wnger movements were used in a 145
104 gest that manual preference and manual asymmetry are transfer situation. One task (serial) consisted of modiWca- 146
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105 aVected diVerently by LP. tion of the sequence of the main task by touching the thumb 147
106 Incongruence between manual preference and manual with the other Wngers in the following order: little, ring, 148
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107 asymmetry in Teixeira and Teixeira’s (2007) study has middle, and index Wnger. As in the main task, one trial con- 149
108 been explained by proposing that an important component sisted of performing three cycles of between-Wnger touch- 150
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109 of manual preference arises from increased conWdence on a ing movements. The other task (repetitive) consisted of 151
110 single limb as a result of LP, rather than from performance keeping the overall sequence of movements as for the main 152
111 advantage of one hand. On the basis of this conceptualiza- task, but performing three repetitive touches with each 153
112 tion, an individual becomes more conWdent on that hand Wnger before starting another triplet of touches. Thus, each 154
113 used most in previous opportunities to perform a motor trial in this task was performed by making consecutive trip- 155
114 task, and this increased conWdence on a single hand would lets of touches to the thumb with the index, then the ring, 156
middle, and last the little Wnger. 157
1 Participants performed the main and transfer tasks while 158
F1 See the eVect of task complexity on manual asymmetry in Hausmann
F2 et al.’s (2004) results, with increased symmetry of performance in sitting on a chair, having the elbow of the active hand 159
F3 more complex sequential Wnger movements in contrast to advantage of upheld on a table. The forearm was kept stable by the 160
F4 the preferred hand in repetitive Wnger tapping. participants without physical constraints in a predominant 161

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162 vertical orientation, slightly bent forward, with the active following the end of practice, manual preference for the 213
163 hand pronated. Movements were Wlmed using a digital main and transfer tasks, and manual asymmetry for the 214
164 camera (SONY, DV-500), and images were analyzed at practiced task were evaluated by repeating the procedures 215
165 60 Hz in order to measure movement time (MT) on each employed in the pretest. In the sequence, participants were 216
166 trial. assessed on manual asymmetry in the serial and repetitive 217
transfer tasks, with the same procedures as for the main 218
167 Experimental design and procedures task. Sequence of evaluation of transfer tasks was counter- 219
balanced across participants. During pretest, posttest, reten- 220
168 Experimental procedures were initiated by assessing over- tion and transfer trials no feedback was provided about 221
169 all handedness and then speciWc manual preference for the movement time. Trials with any sequencing errors or inter- 222
170 experimental tasks. Manual preferences for the main and ruption of Wnger movements (approximately 7% of the tri- 223
171 for the two transfer tasks were assessed by asking partici- als) were aborted and repeated immediately. 224

F
172 pants about their manual preference for those particular Measurement of movement time was made as a function 225

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173 tasks. Overall handedness and speciWc manual preferences of number of frames (17 ms each) in the video analysis. 226
Author Proof

174 were assessed on a Wve-point continuous rating scale: Movement time in a trial corresponds to the interval 227

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175 1 = left always, 2 = left usually, 3 = indiVerent, 4 = right between the moment of visually detectable initiation of clo- 228
176 usually, and 5 = right always. In the sequel, participants sure between the index Wnger and the thumb in the Wrst 229
177 were provided with instructions about the task, and then cycle of movements and the moment at which no further 230

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178 with familiarization trials. Familiarization consisted of displacement was detected in the closure between the little 231
179 three trials performed slowly, followed by another set of Wnger and the thumb in the last cycle of movements. Mea- 232
180 three trials performed at a fast rate. This procedure was surement of movement time across experimental phases 233
181 employed for both hands immediately before initial evalua- and participants were made by a single rater. In order to 234
182
183
tion of manual asymmetry.
For experimental treatment, participants were pseudor- D estimate reliability of movement time measurement, a sam-
ple of 30 trials (10 from each experimental phase) were
235
236
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184 andomly assigned to one of two groups: experimental evaluated twice by the rater with a period of 1 week 237
185 (n = 10; 6 males, 4 females), or control (n = 10; 7 males, 3 between measurements. The rate of perfect coincidence 238
186 females). The experiment was divided into four phases: (proportion of agreements) between the two measurements 239
187 pretest, practice (or rest), posttest, and retention. In the pre- was 67%, while in 27% of the trials it a discrepancy of one 240
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188 test, performance of both hands was assessed on the experi- frame was found, and in 6% of the trials there was a dis- 241
189 mental task only. Participants performed three trials for crepancy of two frames. 242
190 each hand in sequence, having order of hands counterbal-
191 anced across participants. There were regular intervals of
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192 approximately 10 s between trials, and the interval between Results 243
193 assessments of each hand was 1 min approximately.
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194 Following pretest, the experimental group were provided Manual asymmetry 244
195 with practice for the left hand on the main task, while the
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196 control group had no activities other than those usually per- Assessment of manual asymmetry for the main task was 245
197 formed on their daily living duties. Practice trials were conducted through a three-way 2 (group) £ 2 (hand) £ 3 246
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198 divided into three sessions, conducted on diVerent days, (phase) analysis of variance with repeated measures on the 247
199 completed within a period of 1 week. In each session, the last two factors. The results showed signiWcant main eVects 248
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200 experimental group performed Wve blocks of 20 trials, with of group, F(1,18) = 5.63, P < 0.05, and phase, F(2,36) = 39.94, 249
201 rest intervals of a few seconds self determined by the par- P < 0.0001. The main eVect of group is due to shorter MTs 250
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202 ticipants. Thus, at the end of this phase participants had per- for the experimental group (M = 2.45 s) in comparison with 251
203 formed 300 trials, corresponding to 900 cycles of Wnger controls (M = 3.07 s), while the main eVect of phase is due 252
204 movements. In order to increase motivation to improve per- to longer MTs in the pretest (M = 3.32 s) in comparison 253
205 formance, participants trained the task in couples. During with posttest (M = 2.46 s) and retention (M = 2.50 s), which 254
206 each session of practice, while one participant performed did not diVer from each other. No other main eVect or inter- 255
207 the task, the other in a couple registered the time spent to action was detected. This result indicates a similar reduc- 256
208 complete each trial with a stopwatch. Movement time was tion of MT for the right and the left hand in the posttest and 257
209 informed to participants after every trial, and they were retention (Fig. 1a). As an advantage was detected favoring 258
210 asked to improve their performance across training ses- the experimental group across all phases, an additional 259
211 sions. Practice trials were performed under supervision of analysis taking values of pretest as a covariate was made. 260
212 the laboratory staV. One (posttest) and 30 (retention) days Given that no signiWcant main eVect of hand or associate 261

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(M = 2.80 and 2.55 s, respectively) group, while the main 282

eVects of phase are due to increment of MT from posttest 283
(M = 2.50 and 2.32 s, respectively) to retention (M = 2.77 284
and 2.57 s, respectively). 285

Manual preference 286

Statistical analysis of manual preference for the main and 287

transfer tasks was made by comparing the preference scores 288
across experimental phases separately for each group. This 289
analysis was initially conducted through Friedman’s rank 290
test, followed by Wilcoxon’s matched pairs signed-ranks 291

F
test to make follow-up contrasts. The results indicated a 292

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signiWcant phase eVect for the experimental,
F2 =14.77, 293
Author Proof

P < 0.001, but not for the control,
F2 =2.33, P > 0.3, 294

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group. Paired comparisons between phases for the results of 295
the experimental group showed signiWcant reduction of the 296
manual preference score in the posttest, Z = 2.66, P < 0.01, 297

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and retention, Z = 2.52, P < 0.02, in comparison with the 298
pretest, while the diVerence between the two latter experi- 299
mental phases did not reach signiWcance. 300
Analysis of the serial transfer task did not detect a sig- 301

D niWcant phase eVect either for the experimental or for the

control group (Ps > 0.05), while analysis of the repetitive
302
303
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transfer task revealed similar eVects in comparison with the 304
main experimental task. Friedman’s rank test indicated sig- 305
niWcant diVerences between phases for the experimental, 306

F2 =10.69, P < 0.005, but not for the control,
F2 =2.00, 307
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Fig. 1 Movement time (s) of the right and left hands for the experi- P > 0.3, group. Paired comparisons between phases for the 308
mental and the control group across phases for the main (a), and for results of the experimental group showed a signiWcant 309
the serial (b) and repetitive (c) transfer tasks; SD represented by
reduction of the manual preference score in the posttest, 310
vertical bars
Z = 2.37, P < 0.05, and retention, Z = 2.20, P < 0.05, in 311
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comparison with the pretest, with no signiWcant diVerence 312

262 interaction were detected in the primary analysis, the statis- between the latter two phases. 313
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263 tical model employed was a two-way 2 (group) £ 2 (phase: Complementary analysis was made by comparing scores 314
264 posttest £ retention) analysis of covariance with repeated of manual preference between groups across tasks and 315
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265 measures on the second factor. Results indicated a signiW- phases through Mann–Whitney U test. For the main task, 316
266 cant main eVect of group, F(1,17) = 9.34, P < 0.01, indicat- the results indicated signiWcantly lower scores in the exper- 317
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267 ing that practice per se accounts for a signiWcant part of the imental group as compared with the control group in the 318
268 diVerence of performance observed between groups in the posttest, Z = 3.33, P < 0.001, and retention, Z = 2.14, 319
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269 latter experimental phases. P < 0.05. For transfer tasks, analysis indicated signiWcantly 320
270 Assessment of manual asymmetry in the transfer tasks lower scores in the experimental group only for the serial 321
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271 were performed through three-way 2 (group) £ 2 task in the posttest, Z = 2.43, P < 0.05. Figure 2 presents 322
272 (hand) £ 2 (phase: posttest vs. retention) analyses of vari- the median scores of manual preference observed at each 323
273 ance with repeated measures on the last two factors. For the phase for the experimental (panel a) and the control group 324
274 serial transfer task, analysis showed signiWcant main eVects (panel b). Ranges of variation (extreme scores) are repre- 325
275 of group, F(1,18) = 4.40, P = 0.05, and phase, F(1,18) = 14.15, sented by vertical bars. 326
276 P < 0.005 (Fig. 1b). The same proWle was found for the Correlation between the scores of manual asymmetry and 327
277 repetitive transfer task, with signiWcant main eVects of manual preference was estimated by applying Spearman 328
278 group, F(1,18) = 4.24, P = 0.05, and phase, F(1,18) = 8.96, rank order analysis. Manual asymmetry was calculated by 329
279 P < 0.01 (Fig. 1c). In both analyses, the main eVects of subtracting MT of the left hand from MT of the right hand, 330
280 group are due to shorter MTs in the experimental (M = 2.47 and then dividing the product by their sum. The higher the 331
281 and 2.34 s, respectively) as compared with the control value, the larger was the diVerence of performance between 332

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in addition, suggests that the left hemisphere has a role not 360
only in planning but also in the execution of complex 361
movements by either hand. As a corollary, it is apparent 362
that both the right and the left hand share an important part 363
of the neural ensemble employed to control sequential 364
movements, leading to symmetric performance between the 365
hands and to equivalent improvement as a result of uniman- 366
ual practice. 367
Manual preference for a given motor task has been pro- 368
posed to be established as a function of superior perfor- 369
mance of one hand in comparison with the other (Bishop 370
1989). From this proposition, larger manual asymmetries 371

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would lead to stronger manual preference for the advan- 372

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taged hand, while symmetric performance between the 373
Author Proof

hands would result in indiVerent manual preference. Our 374

O
results discredit such a proposition. Conversely, the present 375
Wndings suggest that manual preference and manual asym- 376
metry of performance are independent dimensions of 377

PR
behavior. Before LP of the nonpreferred left hand, all par- 378
Fig. 2 Median scores of manual preference for the experimental (a) ticipants declared right hand preference for the practiced 379
and for the control (b) groups across phases for the main, and for the
serial and repetitive transfer tasks; range of scores across participants and transfer tasks, which is in agreement with a more gen- 380
are represented by vertical bars eral trend toward consistent right hand preference, as indi- 381

333 the hands. This analysis was applied separately for each D cated by the Edinburgh inventory of manual dominance.
Following LP, a signiWcant shift of manual preference for
382
383
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334 group by task by phase. The analysis did not detect any sig- the main experimental task was found, with a number of 384
335 niWcant correlation between scores of manual asymmetry cases of conversion from right to left hand preference. Of 385
336 and manual preference either for the main (rs range: ¡0.27 particular interest for the purposes of the present study was 386
337 to 0.54, P values > 0.05) or for transfer (rs range: ¡0.44 to the Wnding that such shift of manual preference to the left 387
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338 0.61, P values > 0.05) tasks. hand was observed to be generalizable to a transfer task. 388
Interestingly, generalization of manual preference was 389
not equivalent between the two transfer tasks, although 390
339 Discussion both the serial and repetitive tasks required fast sequences 391
R

of Wnger movements. This diVerential eVect between tasks 392

340 Consistent with previous Wndings (Teixeira and Teixeira is apparently related to the order of the movements required 393
R

341 2007), the results showed that practice of the left hand on by each task. While for the serial task a simpler sequence of 394
342 the main task did not modify the symmetric performance movements from the little to the index Wnger was required, 395
O

343 observed before practice, with similar improvement of per- in the repetitive task, triplets of Wnger movements had to be 396
344 formance between the right and the left hand. This Wnding performed in the same order of the practiced task. As LP of 397
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345 is conceived to be related to the neural network engaged to sequential motor actions has been shown to lead to changed 398
346 control actions of this nature. Previous studies have demon- activation of cerebral areas associated with planning of the 399
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347 strated that performance of complex sequential Wnger sequential order of movements (Grafton et al. 1998, 2002; 400
348 movements, by either hand, is featured by cerebral activa- HlustÂk et al. 2002; Haaland et al. 2004; Parsons et al. 401
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349 tion of bilateral cortical areas (Solodkin et al. 2001), with 2005), neural adaptability at that neural component by 402
350 prominence of the premotor (Hlustik et al. 2002), primary learning a particular sequence of transitions between Wnger 403
351 motor (Verstynen et al. 2005), and parietal (Haaland et al. movements might be related to conversion from right to left 404
352 2004) cortices of the left hemisphere. Haaland et al. (2004) manual preference in the repetitive transfer task. Support- 405
353 have proposed that the left dorsal premotor and parietal ing this proposition, Parsons and colleagues (2005) have 406
354 areas are engaged when advance planning is necessary to shown that increased cortical activity in transfer tasks fol- 407
355 perform complex sequences of movements requiring selec- lowing extensive practice of sequential Wnger movements 408
356 tion of diVerent eVectors and abstract organization of the correlated with increased reaction time, but not with 409
357 sequence, regardless of the performing hand. More intense increased movement time. This result suggests that the 410
358 activity of the left primary motor cortex speciWcally in the transfer tasks interfered with the neural representation of 411
359 execution of complex movements (Verstynen et al. 2005), plans for the sequential movements, but not with processes 412

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413 controlling their implementation. Thus, it is apparent that been presented by Teixeira and Teixeira (2007). According 466
414 changing an extensively practiced sequence of movements to that proposition, an important component deWning man- 467
415 requires resources from higher order levels of processing, ual preference is conceived to be an increased conWdence 468
416 in order to specify a new sequence and inhibit the learned on a single hand developed on the basis of the recent his- 469
417 ones, making movement control less automatic. Require- tory of diVerential use of the limbs on a motor task. From 470
418 ment of a more controlled execution of the sequence of this conceptualization, an individual becomes more conW- 471
419 Wnger movements leads to a context similar to performing a dent on that hand used more frequently in previous oppor- 472
420 new motor task, potentially reducing the preference of the tunities to perform a motor task. This increased conWdence 473
421 practiced hand in the transfer task. From this interpretation, would inXuence the choice of the hand to perform the prac- 474
422 we hypothesize that sequence of transitions between ticed and related motor tasks in future instances. Findings 475
423 Wngers, rather than dexterity in moving individual Wngers, of generalizable shift of manual preference, thus, are in 476
424 was the main factor determining a signiWcant shift of man- agreement with expectations from the proposition of estab- 477

F
425 ual preference for repetitive but not for the serial task. lishment of manual preference from increased conWdence 478

O
426 Persistence of a shifted manual preference for the main on the practiced limb. 479
Author Proof

427 and for the repetitive transfer task after 30 days of rest dem- In conclusion, our Wndings represent direct evidence for 480

O
428 onstrates that LP had an enduring eVect (cf. Teixeira and the eVect of learning on human laterality, supporting the 481
429 Teixeira 2007). As indicated by non-signiWcant values of notion of handedness as a dynamic aspect of motor behav- 482
430 correlation between scores of manual asymmetry and man- ior (cf. Provins 1997; Serrien et al. 2006). Worth noticing 483

PR
431 ual preference, such shift of manual preference was found in this regard is the fact that the reported modiWcations of 484
432 not to be due to asymmetrization of performance favoring manual preference took place in right-handers, some of 485
433 the left hand (cf. Dassonville et al. 1997). Even though whom declared to have become consistent left-handers for 486
434 manual asymmetry can not be disregarded as a factor con- those experimental tasks after practice. Further, demonstra- 487
435
436
tributing to deWnition of manual preference in motor tasks
featured by expressive manual asymmetry favoring one D tion that manual preference was not related with manual
asymmetry leads to the notion that general manual prefer-
488
489
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437 hand, lack of association between manual asymmetry and ence might be the precursor of manual asymmetries in a 490
438 manual preference in the main and transfer tasks in this number of motor tasks rather than the prevalent point of 491
439 study requires alternative explanations other than perfor- view that manual preference is determined by manual 492
440 mance asymmetry for the observed variation of manual asymmetry. Generalization of manual preference forged by 493
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441 preference. unimanual experiences on a single task to related motor 494

442 Investigating functional brain activation following prac- actions may be a key point to understand why one hand is 495
443 tice of the preferred right hand on a visuomotor task, Flo- consistently preferred for execution of voluntary move- 496
444 yer-Lea and Matthews (2004) observed that the initial, ments even when that hand is not superior in performance. 497
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445 attentionally demanding stage of learning was associated

446 with greater activity in predominantly cortical regions,
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447 including prefrontal, sensorimotor, and parietal cortices. As References 498

448 performance improved with extensive practice, cortical
O

449 activity decreased (cf. Morgen 2004), giving place to Bishop DVM (1989) Does hand proWciency determine hand prefer- 499
450 enhanced activity in subcortical motor regions (cf. Floyer- ence? Br J Psychol 80:191–199 500
Dassonville P, Zhu XH, Ugurbil K, Kim SG, Ashe J (1997) Functional 501
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451 Lea and Matthews 2005; Puttemans et al. 2005). These activation in motor cortex reXects the direction and the degree of 502
452 Wndings suggest that as a task is progressively automated handedness. Proc Natl Acad Sci USA 94:14015–14018 503
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453 by means of learning, subcortical circuits become domi- Denckla MB (1974) Development of motor co-ordination in normal 504
454 nant, making movement control attentionally less eVortful. children. Dev Med Child Neurol 16:720–742 505
Fagard J (1987) Does manual asymmetry of right-handers change be- 506
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455 In line with these results, we propose that shift of manual tween six and nine years of age? Hum Mov Sci 6:321–332 507
456 preference might be a consequence of a more automatic Floyer-Lea A, Matthews PM (2004) Changing brain networks for vis- 508
457 processing of sensorimotor information required to main- uomotor control with increased movement automaticity. J Neuro- 509
458 tain accurate motor output. Lateralized practice would lead physiol 92:2405–2412 510
Floyer-Lea A, Matthews PM (2005) Distinguishable brain activation 511
459 then to a less eVortful processing when the task is per- networks for short- and long-term motor skill learning. J Neuro- 512
460 formed by the practiced hand, with a more comfortable exe- physiol 94:512–518 513
461 cution of the motor act, a situation potentially leading to its Grafton ST, Hazeltine E, Ivry RB (1998) Abstract and eVector speciWc 514
462 selection. representations of motor sequences identiWed with PET. J Neuro- 515
sci 18:9420–9428 516
463 An alternative explanation for shift of manual preference Grafton ST, Hazeltine E, Ivry RB (2002) Motor sequence learning with 517
464 as a result of LP, although not incompatible with the the nondominant left hand: a PET functional imaging study. Exp 518
465 automatization of sensorimotor processing hypothesis, has Brain Res 146:369–378 519

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520 Haaland KY, Elsinger CL, Mayer AR, Durgerian S, Rao SM (2004) Provins KA (1997) Handedness and speech: a critical reappraisal of the 550
521 Motor sequence complexity and performing hand produce diVer- role of genetic and environmental factors in the cerebral laterali- 551
522 ential patterns of hemispheric lateralization. J Cogn Neurosci zation of function. Psychol Rev 104:554–571 552
523 16:621–636 Puttemans V, Wenderoth N, Swinnen SP (2005) Changes in brain acti- 553
524 Hausmann M, Kirk IJ, Corballis MC (2004) InXuence of task complex- vation during the acquisition of a multifrequency bimanual coor- 554
525 ity on manual asymmetries. Cortex 40:103–110 dination task: from the cognitive stage to advanced levels of 555
526 Healey JM, Liederman J, Geschwind N (1986) Handedness is not a automaticity. J Neurosci 25:4270–4278 556
527 unidimensional trait. Cortex 22:33–53 Serrien DJ, Ivry RB, Swinnen SP (2006) Dynamics of hemispheric 557
528 Hlustik P, Solodkin A, Gullapalli RP, Noll DC, Small SL (2002) Func- specialization and integration in the context of motor control. Nat 558
529 tional lateralization of the human premotor cortex during sequen- Neurosci 7:160–167 559
530 tial movements. Brain Cogn 49:54–62 Singh M, Manjary M, Dellatolas G (2001) Lateral preference among 560
531 McGonigle BO, Flook J (1978) The learning of hand preferences by Indian school children. Cortex 37:231–241 561
532 squirrel monkey. Psychol Res 40:93–98 Solodkin A, Hlustik P, Noll DC, Small SL (2001) Lateralization of mo- 562
533 Meng LF (2007) The rate of handedness conversion and related factors tor circuits and handedness during Wnger movements. Eur J Neu- 563

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534 in left-handed children. Laterality 12:131–138 rol 8:425–434 564
535 Mikheev M, Mohrb C, Afanasiev S, Landis T, Thut G (2002) Motor Teixeira LA (2007) Categories of manual asymmetry and their varia- 565

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539 McFarland H, Martin R, Cohen LG (2004) Kinematic speciWcity actions: preference versus performance]. Motriz 6:1–8 569
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542 OldWeld RC (1971) The assessment and analysis of handedness: the Verstynen T, Diedrichsen J, Albert N, Aparicio P, Ivry RB (2005) Ipsi- 572
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544 Parsons MW, Harrington DL, Rao SM (2005) Distinct neural systems relates to task complexity. J Neurophysiol 93:1209–1222 574
545 underlie learning visuomotor and spatial representations of motor Zverev YP (2006) Cultural and environmental pressure against left- 575
546 skills. Hum Brain Mapp 24:229–247 hand preference in urban and semi-urban Malawi. Brain Cogn 576
547 Porac C, Coren S, Searleman A (1986) Environmental factors in hand 60:295–303 577
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