Learning by Observation: Insights from Williams

Syndrome
Francesca Foti1,2*., Deny Menghini3., Laura Mandolesi4,2, Francesca Federico1, Stefano Vicari3,
Laura Petrosini5,2
1 Department of Developmental and Social Psychology, University ‘‘Sapienza’’ of Rome, Rome, Italy, 2 IRCCS Santa Lucia Foundation, Rome, Rome, Italy, 3 Child
Neuropsychiatry Unit, Neuroscience Department, ‘‘Children’s Hospital Bambino Gesu’’’, Rome, Rome, Italy, 4 School of Movement Sciences (DiSIST), University of Naples
‘‘Parthenope’’, Naples, Italy, 5 Department of Psychology, University ‘‘Sapienza’’ of Rome, Rome, Italy

Abstract
Observing another person performing a complex action accelerates the observer’s acquisition of the same action and limits
the time-consuming process of learning by trial and error. Observational learning makes an interesting and potentially
important topic in the developmental domain, especially when disorders are considered. The implications of studies aimed
at clarifying whether and how this form of learning is spared by pathology are manifold. We focused on a specific
population with learning and intellectual disabilities, the individuals with Williams syndrome. The performance of twenty-
eight individuals with Williams syndrome was compared with that of mental age- and gender-matched thirty-two typically
developing children on tasks of learning of a visuo-motor sequence by observation or by trial and error. Regardless of the
learning modality, acquiring the correct sequence involved three main phases: a detection phase, in which participants
discovered the correct sequence and learned how to perform the task; an exercise phase, in which they reproduced the
sequence until performance was error-free; an automatization phase, in which by repeating the error-free sequence they
became accurate and speedy. Participants with Williams syndrome beneficiated of observational training (in which they
observed an actor detecting the visuo-motor sequence) in the detection phase, while they performed worse than typically
developing children in the exercise and automatization phases. Thus, by exploiting competencies learned by observation,
individuals with Williams syndrome detected the visuo-motor sequence, putting into action the appropriate procedural
strategies. Conversely, their impaired performances in the exercise phases appeared linked to impaired spatial working
memory, while their deficits in automatization phases to deficits in processes increasing efficiency and speed of the
response. Overall, observational experience was advantageous for acquiring competencies, since it primed subjects’ interest
in the actions to be performed and functioned as a catalyst for executed action.

Citation: Foti F, Menghini D, Mandolesi L, Federico F, Vicari S, et al. (2013) Learning by Observation: Insights from Williams Syndrome. PLoS ONE 8(1): e53782.
doi:10.1371/journal.pone.0053782
Editor: Tricia A. Thornton-Wells, Vanderbilt University, United States of America
Received July 12, 2012; Accepted December 4, 2012; Published January 10, 2013
Copyright: ß 2013 Foti et al. 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 author and source are credited.
Funding: This work was supported by the Fondation Jérôme Lejeune to L.M. The funders had no role in study design, data collection and analysis, decision to
publish, or preparation of the manuscript.
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: [email protected]
. These authors contributed equally to this work.

Introduction By using an innovative task based on learning to detect a visuo-

motor sequence, we demonstrated that in the presence of dyslexia
In humans and other animals new competencies may be learned the ability to learn by observation a previously observed visuo-
through active experience and through observation of others’ motor sequence was markedly impaired, while the ability to detect
experience [1,2]. Observing another person performing a complex a correct sequence by trial and error was preserved [16]. In the
action accelerates the observer’s acquisition of the same action and present research we focused on a population with learning as well
limits the time-consuming process of learning by trial and error as intellectual disability (ID), the Williams syndrome (WS) whose
[3–5]. Indeed, observational learning does not just involve copying well-known neuropsychological profile with specific points of
an action and requires that the observer transforms the strengths and weaknesses allowed singling out cognitive processes
observation into an action as similar as possible to the model in working as learning went by. WS individuals show severely
terms of the goal to be reached and the motor strategies to be impaired visuo-spatial processing, planning and implicit learning
applied [5–10]. [17–22], while they exhibit relatively preserved perception of the
Observational learning is already present at birth [5,11–13] and visual characteristics of objects and face recognition [23]. WS
it is crucial for developing complex abilities such as language, individuals have specific difficulty in maintaining visuo-spatial
social responsiveness, use of instruments to get things done [9,14]. information in working memory and in performing long-term
Thus, in children, learning new competencies by observing adults memory tasks [24,25], consistently with a deficit of dorsal stream.
or peers is a central process in cognitive development [15]. Considering that the visuo-motor task to be learned by observation
required to translate visual information into action, specific

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 Learning by Observation in Williams Syndrome

function of dorsal stream network [26,27], WS individuals appear The clinical diagnosis of WS was confirmed by fluorescence in
to be the ideal participants to investigate the cognitive processes situ hybridization (FISH) genetic investigation, which showed the
involved in the observational learning. Performances of a group of characteristic deletion on chromosome band 7q11.23. WS
WS individuals were compared with those of a mental age- and participants were part of a larger pool of individuals with learning
gender-matched group of typically developing (TD) children on a disabilities attending the Children’s Hospital Bambino Gesù of
task requiring the learning of a visuo-motor sequence. The Rome for clinical and rehabilitative follow-up. All of them lived at
participants learned the sequence either by performing the task home with their families. The parents of all individuals who
after observing an actor detect the sequence of correct items by participated in the study provided written informed consent. This
trial and error (observational training) or by actually performing study was approved by the Ethic Committee of the Children’s
the task by trial and error (Fig. 1). Hospital Bambino Gesù of Rome and conducted according to the
Helsinki declaration.
Materials and Methods WS individuals were tested in a quiet room at the Children’s
Hospital Bambino Gesù. TD children were individually tested in a
Participants quiet room at their schools.
Twenty-eight WS participants and 32 TD children (used as
controls) matching the WS individuals for mental age (MA) have Intelligence Evaluation and Neuropsychological
been examined in the present study constituted by two experi- Assessment
mental conditions: Learning by Trial and Error followed by In the present study, the brief version of the Leiter International
Observational Learning (Condition 1); Observational Learning Performance Scale–Revised [28] was employed (four out of 10
followed by Learning by Trial and Error (Condition 2) (Table 1). subtests: Figure Ground, Form Completion, Sequential Order and
Only WS individuals with mental age (MA) of at least 5 years were Repeated Patterns). The brief IQ and the corresponding mental
included in the present research because participants with inferior ages were computed. Visuo-motor integration [29], visuo-spatial
MA did not succeed in completing the task. No significant perception [30] and memory [31] were assessed (Table 3).
differences in chronological age (CA), MA and IQ (P always .0.2)
among participants performing Conditions 1 and 2 were found Experimental Procedure
(Table 2). Each participant was sat in front of a computer touch screen
(distance 60 cm). In both Conditions, the experimenter acting as

Figure 1. Schematic diagrams of the two experimental conditions. Condition 1: Learning by Trial and Error followed by Observational
Learning: participants detected a sequence by trial and error (TE1), then they observed an actor detecting a sequence different from the one they had
previously detected (observational training) and, finally, they reproduced the observed sequence (OBS2). Condition 2: Observational Learning
followed by Learning by Trial and Error: participants were submitted to an observational training, then they reproduced the observed sequence
(OBS1) and, finally, detected by trial and error a different sequence they had never observed (TE2). The incorrect positions touched by the actor
during the observational training are evidenced in grey. S: starting point; F: final point.
doi:10.1371/journal.pone.0053782.g001

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 Learning by Observation in Williams Syndrome

Table 1. Description of WS groups (WS1 and WS2) and TD groups (TD1 and TD2) performing the two different experimental
conditions.

Condition 1: Learning by Trial and Error followed by Observational Learning

CA MA IQ
Group Number Gender Mean ± SEM Mean ± SEM Mean ± SEM

WS1 14 9M 19.8361.42 6.5260.16 54.8761.69

TD1 16 11 M 6.7860.15 7.0260.28 106.1262.51

Condition 2: Learning by Observation followed by Learning by Trial and Error

WS2 14 8M 17.6461.37 6.6060.19 53.6861.36

TD2 16 8M 6.7660.11 7.4060.28 111.6262.06

CA: Chronological Age.

MA: Mental Age.
IQ: Intelligence Quotient.
doi:10.1371/journal.pone.0053782.t001

the actor (F.F.) was sat near the participant. A 868 black matrix that all participants observed the same pattern of correct and
appeared on the touch screen. The subject was asked to find a incorrect touches. Two minutes after the end of the observational
hidden sequence of ‘‘correct’’ squares prepared in advance by the training the participants were required to actually reproduce the
experimenters. The sequence was composed of 10 adjacent spatial observed correct sequence.
positions in the matrix, which formed a ‘‘snake-like’’ pattern The tasks involved three phases: the Detection Phase (DP) that
(Fig.1). To explain the task to each participant the experimenter ended once the participants found the tenth correct position; the
used the same verbal instructions: ‘‘You have to find a route Exercise Phase (EP) in which they had to repeat the 10-item
formed by ten squares. When you touch a correct square it will be sequence until their performance was error-free; the Automatiza-
turned grey and you will hear a sound; conversely, if you touch a tion Phase (AP) that ended when the correct sequence was
wrong square, it will be turned red. In this case, you have to find a repeated three consecutive times without errors.
new grey square. You have to start the route each time you find a
new correct square. After finding the whole route, you have to re- Parameters
touch it three times without making lighted red squares’’. The Error parameters: DP errors, calculated as the number of
participants started touching a grey square, which was the first incorrect items touched in detecting the ten correct positions; EP
element of the sequence and was always lit up. In the search for repetitions, calculated as the number of replications needed to reach
the second correct square, the participants had to touch one of the the error-free performance. Time parameters: AP times (in msec),
four squares bordering the grey square by moving in the matrix calculated as the time spent carrying out each of the three
vertically or horizontally, but never diagonally. Each touched repetitions of the sequence.
square (correct or incorrect) was lit up for 500 ms and then lighted
off again; thus, no trace of the performed sequence remained on
the screen. In learning the sequence by trial and error, the Analysis of Error
participants tried to find the correct sequence immediately after To assess the kind of error further parameters were taken into
the verbal instructions. Conversely, in the observational learning account considering the two phases DP and EP together: the
task after the verbal instructions the participants observed the number of sequence errors, as touching a ‘‘correct’’ square in
experimenter while she detected a 10-item sequence by trial and ‘‘wrong’’ moment (e.g. touching E7 before than F7); side-by-side
error (observational training). The experimenter performed the errors, as touching the squares bordering the ‘‘correct’’ sequence
task by always making the same errors in the same positions, so (e.g. E8); illogical errors, as touching any other square (e.g. B5);

Table 2. Comparisons of chronological age (CA), mental age (MA) and Intelligence Quotient (IQ) between WS groups (WS1 and
WS2) and TD groups (TD1 and TD2) that performed the two different experimental conditions.

CA MA IQ
Group Mean ± SEM F (freedom degrees) Mean ± SEM F (freedom degrees) Mean ± SEM F (freedom degrees)

WS1 19.8361.42 (1,26) 6.5260.16 (1,26) 54.8761.69 (1,26)

0.61 0.04 0.15
P = 0.43 P = 0.83 P = 0.70
WS2 17.6461.37 6.6060.19 53.6861.36
TD1 6.7860.15 (1,30) 7.0260.28 (1,30) 106.1262.51 (1,30)
0.005 0.45 1.43
P = 0.94 P = 0.50 P = 0.23
TD2 6.7660.11 7.4060.28 111.6262.06

doi:10.1371/journal.pone.0053782.t002

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 Learning by Observation in Williams Syndrome

Table 3. Statistical comparison of visuo-spatial performances of WS and TD participants.

WS TD
Mean ± SEM Mean ± SEM Effect F(1, 58) value P

Visuo-motor integration 13.3960.50 15.2560.26 Group 11.37 0.0013

Visuo-spatial short-term memory (VSS) 2.7960.20 3.5360.14 Group 8.29 0.0055
Task 9.63 0.0029
Interaction 4.82 0.032
Newman–Keuls test
VSS: 0.00021
VOS: 0.36
Visuo-object short-term memory (VOS) 2.6460.12 2.9160.11
Visuo-perception test – Spatial (VPT-S) 15.1861.07 18.036.77 Group 7.06 0.010
Task 65.75 ,0.0001
Interaction 4.06 0.048
Newman–Keuls test
VPT-S: 0.00038
VPT-F: 0.31
Visuo-perception test – Form (VPT-F) 11.3260.36 12.1660.26

doi:10.1371/journal.pone.0053782.t003

perseverations, as consecutively touching the same item or a fixed Attentional Task

sequence of items. Furthermore, in the task of observational The sustained attentional abilities of all participants were tested.
learning we calculated the number of imitative errors, as touching the Participants sat in front of a computer monitor and were required
same squares wrongly touched by the actor during the observa- to put their left index fingers on the A key of the keyboard and to
tional training (e.g. F4) (Fig.1). put their right index fingers on the L key. The visual stimulus was
a grey circle presented on monitor center for a duration varying
Condition 1: Learning by Trial and Error Followed by from 1400 (short) to 2600 (long) msec in steps of 200 msec in a
Observational Learning randomized order. Participants were submitted to a brief training
Fourteen WS and 16 TD individuals (Table 1) firstly detected a in which they were instructed to judge 20 stimuli as short or long
sequence by Trial and Error (TE1) and, after ten minutes from and to press the A or L keys, respectively. In the testing phase the
task end, they were submitted to the observational training. After participants had to judge the duration of 70 stimuli (10 stimuli of
two minutes, participants were required to actually reproduce the each of the 7 durations) and to press the A or L keys as quickly as
observed sequence (OBS2). There was no fixed time limit for possible after the stimulus appeared. The computer program
executing the task. recorded reaction times (with 1-ms resolution) and accuracy of the
A pilot study was conducted to verify if the two sequences response. The responses were then analyzed by clustering them in
blocks of ten (regardless of stimulus duration) (i.e. 1–10, 11–20,
arranged to be used as ‘‘TE’’ and ‘‘OBS’’ sequences did not differ
21–30….61–70).
as to degree of difficulty. Six TD children [3 M] of mental age
6.1060.3 detected the two different sequences by trial and error;
presentation order was randomized among participants. Errors Statistical Analyses
made in detecting each sequence were calculated by one-way The data were first tested for normality (Shapiro-Wilk normality
ANOVA with repeated measures. The analysis failed to reveal any test) and homoscedasticity (Levene test) and then compared by
significant difference between sequences (F(1,5) = 0.63, P = 0.46), using two-way, three-way or four-way analyses of variance
confirming sequences of the same difficulty. (ANOVAs). The two-way ANOVAs were performed by applying
the mixed model for independent variable (group) and repeated
measures (error, square or block). Three-way ANOVAs (group6
Condition 2: Learning by Observation Followed by condition6task) were performed on most parameters, while the
Learning by Trial and Error four-way ANOVA on the three AP times was performed by
Fourteen WS and 16 TD individuals (Table 1) first observed the applying the mixed model for independent variables (group,
experimenter detect a sequence (OBS1) and then actually condition and task) and repeated measures (time). These analyses
reproduce it. After ten minutes from task end, they detected a were followed by post-hoc multiple comparisons using Newman–
different sequence by trial and error (TE2). Thus, the difference of Keuls test. In evaluating mapping abilities the error categories
the two conditions was that participants reproduced a sequence were analyzed by Chi-Square.
learned by observation after (Condition 1) or before (Condition 2) the Because the 28 WS participants were differently aged (N = 9 age
detection of a different sequence by trial and error. range: 8;9–14;1; N = 10 age range: 14;9–19;9; N = 9 age range:
To evaluate mental representative mapping abilities, at the end 22;9–35;3), we verified the sample homogeneity by comparing the
of the reproduction of the sequence participants were required to performances of three differently aged WS sub-groups on three
draw the arrangement of the sequence on a 868 matrix sketched main parameters of the learning tasks they performed (DP errors;
on a paper sheet. Thus, any participant drew the arrangement of EP repetitions and AP times) by using MANOVAs. These analyses
two sequences, one learned by observation and the other one by revealed no significant difference among WS sub-groups’ perfor-
trial and error. Mapping abilities were evaluated by tabulating the mances. Namely, in the tasks of learning by trial and error (TE1–
variable ‘‘errors’’ into three categories: ‘‘no error’’, ‘‘one error’’ TE2), the MANOVA revealed a not significant sub-group effect
and ‘‘more than one error’’. (F(2,25) = 0.12, P = 0.88) and a significant parameter effect

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 Learning by Observation in Williams Syndrome

(F(2,50) = 154.54, P,0.0001). The interaction was not significant Attentional Task
(F(4,50) = 0.13, P = 0.96). In the tasks of observational learning Two-way ANOVAs (group6block) on reaction times or
(OBS1–OBS2) the MANOVA also revealed a not significant sub- response accuracy of the WS and TD groups revealed no
group effect (F(2,25) = 0.47, P = 0.62) and a significant parameter attentional decay in both groups, as indicated by not significant
effect (F(2,50) = 85.46, P,0.0001). The interaction was not signif- difference in the reaction times in the seven blocks (F(6,348) = 1.55,
icant (F(4,50) = 0.47, P = 0.75). Thus, we pooled together the 28 P = 0.15). A similar result was obtained when response accuracy
differently aged WS individuals. was analyzed (F(6,348) = 1.80, P = 0.10). Notably, a significant
All statistical analyses were performed by using Statistica 8.0 for difference was found between WS and TD groups on reaction
Windows and the significance level was established at P,0.05. times (F(1,58) = 13.52, P = 0.00051), given WS participants pressed
the keys at the appearance of the stimulus more quickly than TD
Results children (Fig. 5).

Learning Tasks Discussion

WS participants performed a number of DP errors not
significantly different from TD children after the observational Our study adopted a matched-group design to determine
trainings (OBS1–OBS2) and were significantly impaired in whether the learning performance of WS individuals was above or
detecting the sequence by trial and error in TE1 compared with below that expected given their general level of intellectual
any other intra- or inter-group condition (Fig. 2A), as revealed by functioning indexed as MA. However, although this design is one
post-hoc comparisons (always P,0.001) on the second-order the most commonly employed measures of matching in ID
interaction (F(1,56) = 8.37, P = 0.0054) of the three-way ANOVA research, we are aware that it has limitations in respect to ID-
(group6condition6task). matched control group design that takes into account the cognitive
In EP, when individuals repeated the sequence until their profile of the specific pathology. Nevertheless, even the ID-
performance was error-free, WS participants needed a significant- matched control group cannot be taken as a guarantee of
ly higher number of repetitions in comparison to TD children normative group, due cognitive profiles among different etiological
regardless of condition (1 or 2) and trial (OBS or TE), as revealed groups with ID exhibit different peaks and troughs [32]. In an
by the group effect (F(1,56) = 9.58, P = 0.0030) of the three-way attempt to overcome the difficulties in matching individuals of
ANOVA (Fig. 2B). The analysis of the three AP times revealed different groups on any one particular measure it has been
that although all participants exhibited significantly reduced times proposed the use of regression techniques that take the factors
as the task went by (F(2,112) = 27.62, P,0.00001), WS individuals related to task performance into account [32]. However, this
were significantly slower than TD children (F(1,56) = 10.37, measure requires specific statistical properties of the data (as
P = 0.0021), revealing a difficulty in automatizing the sequence homogeneity of regression slopes or sample size), hardly available
(Fig. 2C). in studies on population affected by rare genetic conditions as WS.
The present study documented as WS participants significantly
beneficiated of observational training as TD MA-matched
Analysis of Error
children. This was true specifically in the DPs of learning tasks,
In TE1, although WS and TD participants did not differ in
while as for EPs and APs, in all tasks regardless of presentation
the number of illogical errors, WS individuals exhibited values
order (1 or 2) or learning modalities (OBS or TE), WS participants
of sequence, side-by-side and perseverative errors higher than
performed significantly worse than TD children. The powerfully
TD children, as revealed by post-hoc comparisons made on the
positive effect of observational training was present not only in
interaction (F(3,84) = 3.14, P = 0.029) of the two-way ANOVA
reproducing the previously observed sequences (OBS1 and OBS2)
(group6kind of error) (Fig. 3). The highest number of sequence but also affected the subsequent detection of a sequence by trial
errors of WS individuals was found in E7 and F7 squares when and error (TE2). However, the practice effect, inevitably present in
a change of strategy was required (i.e. after an error re-starting any second task, potentially could affect performances.
the sequence from the first item rather than continuing along Since WS individuals exhibit difficulties in maintaining visuo-
on the ‘‘snake’’) (Fig. 1), as revealed by post-hoc comparisons spatial information in working memory and in performing spatial
made on the interaction (F(9,252) = 1.96, P = 0.044) of the two- long-term memory tasks (Table 3) [20,24,25], their heavily
way ANOVA (group6square) (Fig. 4). As for side-by-side errors, impaired performances in all EPs appear linked to spatial working
the high number of errors of WS individuals was due to their memory deficits and difficulties in bringing together the short
significantly more frequent touching of a wrong square when a sequences detected during DP, in maintaining them in working
change of direction was required (squares: D7, F6, E1) (Fig. 1), memory to recall the whole sequence trace and in monitoring the
as revealed by post-hoc comparisons made on the interaction correct execution of the sequence. These findings indicate that the
(F(27,756) = 2.42, P,0.0001) of the two-way ANOVA (group6 observational training exerts beneficial effects mainly on the
square) (Fig. 4). acquisition of strategies to be applied.
The analysis of error in the remaining tasks (OBS2, OBS1 and In both Conditions, WS participants displayed AP times longer
TE2) revealed no significant difference between groups, even if than TD children, even if progressively diminishing as the task
significant difference among kind of errors was found (always went by. This finding was not a consequence of the fine motor
P,0.00001) (Fig. 3). deficits usually reported in WS individuals. In fact, consistently
with Vicari et al. (2007) [22], the reaction times exhibited by WS
Mapping Abilities group in the Attentional Task were even shorter than those of TD
Mental representative mapping abilities of the participants were group. Thus, the longer WS times were related to deficits in
evaluated by having them draw the arrangement of sequences they automatization processes increasing efficiency and speed of the
had just performed. No significant difference among categories of response to reach highest levels of performance [33]. Automatizing
errors and between groups was found in any sequence (P at least skills is mainly linked to the functions of sub-cortical structures, as
.0.4). the cerebellum and basal ganglia and to their bidirectional

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 Learning by Observation in Williams Syndrome

Figure 2. Performances exhibited by WS and TD participants in the two experimental conditions. DP: Detection Phase; EP: Exercise
Phase; AP: Automatization Phase. Data are expressed as mean 6 SEM. The asterisks indicate the significance level of post hoc comparisons between
groups (***P,0.001).
doi:10.1371/journal.pone.0053782.g002

interconnections with cortical structures [34,35,36,37]. The cause By analyzing the kind of errors, some remarks can be made.
of automatization and procedural deficits of WS individuals could First of all, both groups made a very low number of illogical errors,
be their remarkable hypoplasia of the basal ganglia [38] and the thus suggesting all participants similarly managed task fundamen-
disproportionate enlargement of the cerebellum [39,40,41,42,43]. tals (Fig. 3 and 4). As for imitative errors, no difference between
Indeed, in WS individuals skill-learning abilities are impaired, as groups was found, thus suggesting participants did imitate but did
revealed by their performance in Tower of London test [21], not hyperimitate [47]. Conversely, WS individuals made more
Serial Reaction Time task [22] and Radial or Multiple Reward sequence and side-by-side errors than TD children in TE1,
Mazes [44,45,46]. particularly when a change of direction was required. Errors in

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 Learning by Observation in Williams Syndrome

Figure 3. Errors exhibited by WS and TD participants in the two experimental conditions. Data are expressed as mean 6 SEM. The
asterisks indicate the significance level of post hoc comparisons between groups (*P,0.05; ** P,0.005; *** P,0.001).
doi:10.1371/journal.pone.0053782.g003

stopping the more easy ‘‘keep-straight’’ response and performing The prevalently frontal processes require a modulation in
the more demanding ‘‘turn-left’’ response resulted by the WS more posterior brain systems, via the attention networks. Basic
difficulty in suppressing a previously correct but then inappropri- aspects of attentional processing are selective spatial attention
ate response. Correctly responding requires executive control that allows maintaining the focus of processing between spatial
processes based on frontal lobe function, as response inhibition, locations, and the attentional processing that allows a kind of
cognitive flexibility and attentional shifting [48,49,50,51]. WS ‘‘selection for action’’ [58,59,60]. Namely, the action of
individuals are impaired in spatial planning, working memory, reaching the right square required attentional modulation to
cognitive flexibility and inhibiting well-learned responses become plan, select and initiate the appropriate behavior, to direct it
inappropriate to the situation [52]. Indeed, the executive function toward the selected goal, and to inhibit actions inappropriate for
deficits that impaired WS performance dramatically reduced after the current goal. Because many brain structures that are part of
the observational training, once more indicating the teaching the attention networks are included within the dorsal stream
power of the observation. WS participants made perseverative network [27], it is not surprising that WS participants
errors that could result from difficulties in withholding the performed more errors when behavioral inhibition and attentive
inappropriate repetition of a response despite knowing that it shifting were required but no help from observing the actor was
was not the correct one. This is an important component of top- provided [61]. The ‘‘dorsal-stream vulnerability’’ in WS [62] is
down executive control. Notably, perseverations may be symptom manifested not only in the spatial and visual processing
not only of prefrontal dysfunction but also of cerebellar and basal occurring within the occipital and parietal areas but also in
ganglia damage provoking ‘‘frontal-like’’ cognitive deficits the processing of spatial information by frontal control systems,
[34,53,54,55,56,57]. as reported in an fMRI study [63].

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 Learning by Observation in Williams Syndrome

Figure 4. Incorrect items touched on the screen by WS and TD participants in performing the tasks. On the right, the chromatic scale
indicates the sum of incorrectly touched items (brown and blue denote maximal and minimal values, respectively). S: starting point; F: final point.
doi:10.1371/journal.pone.0053782.g004

At the end of testing, mental representative mapping abilities performed to detect rules and sequence. In fact, the present
were evaluated by drawing the arrangement of squares just results indicate that the observation of action has a strong
discovered. WS and TD participants were similarly able to impact on action memory. The influence of action perception
represent the shape of the ‘‘snake’’. This finding is consistent with on action production requires cross-modal information be
the observation that WS individuals exhibit no difficulty in coordinated. Action and perception share the distal reference
mentally visualizing objects without spatially manipulating them and are coded in a common representational medium [65], so
[25,64] and supports the indication that the present learning that perceiving an action activates the corresponding motor
protocols encompassed requests of visual imagery. representation within the observer automatically and without
If observative experience functions as a catalyst for executed conscious effort [66,67,68].
action, it can be advanced that observing a sequence prior to The close interplay between observation and execution of
experiencing it primes subjects’ interest in the actions to be actions found in the present study is supported by studies
providing evidence of a striking overlap in the brain systems
recruited for one’s own action, observation of others’ action and
imitation of action [69]. In particular, when imitation is aimed at
learning novel actions, the activation of the ‘‘core circuit for
imitation’’ [70] involving the inferior frontal gyrus, the inferior
parietal lobule and the superior temporal sulcus seems to be
integrated with activation of the dorso-lateral and ventro-medial
prefrontal cortex, for selection of motor acts and error prediction
[71], of the premotor areas, for motor preparation [70,72,73] as
well as of cerebellar areas, whether or not it is accompanied by
actual motor acts [74,75,76].
The existence of direct feed-forward connections from
perceptual to motor processes allows observation sculpting
motor abilities by exploiting the functional overlapping between
perception and action systems. It has been suggested that
observation of actions engages motor-related processes similar to
those of actual execution, promoting the development of an
efference copy of the descending motor commands, which in
combination with a forward model provides a prediction of
sensory consequences [77,78,79,80]. Thus, action observation,
efficiently translated into the matching motor representation,
powerfully activates the feed-forward predictive processes, so
that learning does occur. Notably, even in WS individuals the
Figure 5. Performances exhibited by WS and TD participants in beneficial effect of observation was evident although linked only
the Attentional Task. Data are expressed as mean 6 SEM. to the DP. Action observation seems to result in an amelioration
doi:10.1371/journal.pone.0053782.g005 of frontal functions, as motor strategy planning, decision-making

PLOS ONE | www.plosone.org 8 January 2013 | Volume 8 | Issue 1 | e53782

 Learning by Observation in Williams Syndrome

processes or response inhibition needed to guide planned Acknowledgments

sequential actions. Thus, observational training allows the
acquisition of the strategies to be applied to identify and learn We would like to thank the individuals with Williams syndrome and their
parents for making this study possible. Moreover, we wish to thank Dr. P.
the visuo-spatial sequence. Notably, when the observation did
Pani for his kind and competent help in analyzing data.
not play any role (as in the DP of TE1), frontal deficits
markedly affected WS performance. However, it has to be
underlined that as far as the observational training was Author Contributions
beneficial, in WS individuals it did not succeed in smoothing Conceived and designed the experiments: F. Foti DM LM F. Frederico SV
out the deficits in processing visuo-spatial information mainly LP. Performed the experiments: F. Foti. Analyzed the data: F. Foti DM LP.
linked to their repeatedly described dorsal stream vulnerability Contributed reagents/materials/analysis tools: DM LM SV LP. Wrote the
[62]. paper: F. Foti DM LP.

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