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Psychological Research (2012) 76:542–560

DOI 10.1007/s00426-011-0383-y

R EV IE W

Long- and short-term plastic modeling of action prediction


abilities in volleyball
Cosimo Urgesi · Maria Maddalena Savonitto ·
Franco Fabbro · Salvatore M. Aglioti

Received: 29 June 2011 / Accepted: 29 September 2011 / Published online: 2 November 2011
© Springer-Verlag 2011

Abstract Athletes show superior abilities not only in exe- Introduction


cuting complex actions, but also in anticipating others’
moves. Here, we explored how visual and motor experi- Common coding hypotheses suggest a shared representa-
ences contribute to forge elite action prediction abilities in tion of perceived and executed actions (Hommel, Musseler,
volleyball players. Both adult athletes and supporters were Aschersleben, & Prinz, 2001; Prinz, 1997) and a bidirec-
more accurate than novices in predicting the fate of volley- tional link between perceptual and motor codes. Behavioral
ball Xoating services by viewing the initial ball trajectory, studies have shown that perceiving static or dynamic dis-
while only athletes could base their predictions on body plays of moving body parts aVects the execution of congru-
kinematics. Importantly, adolescents assigned to physical ent as compared with incongruent body movements (Brass,
practice training improved their ability to predict the fate of Bekkering, Wohlschläger, & Prinz, 2000; Kilner, Paulig-
the actions by reading body kinematics, while those nan, & Blakemore, 2003; Stürmer, Aschersleben, & Prinz,
assigned to the observational practice training improved 2000). Moreover, contextual active execution of a motor
only in understanding the ball trajectory. The results sug- task strongly inXuences action perception (Springer et al.,
gest that physical and observational practice might provide 2011). Accessing the motor representations used during
complementary and mutually reinforcing contributions to planning and execution of actions might be a deceivingly
the superior perceptual abilities of elite athletes. Moreover, simple mechanism for the striking ability to create an antic-
direct motor experience is required to establish novel per- ipatory representation of others’ actions (Perrett, Xiao, Bar-
ceptuo-motor representations that are used to predict oth- raclough, Keysers, & Oram, 2009; Schütz-Bosbach &
ers’ actions ahead of realization. Prinz, 2007; Wilson & Knoblich, 2005). Indeed, the full
sequence of motion is rarely visible during interactions
with a dynamic world and missing information needs com-
C. Urgesi (&) · M. M. Savonitto · F. Fabbro pletion via top-down modulation (Komatsu, 2006; Pessoa,
Dipartimento di Scienze Umane, Thompson, & Noë, 1998). Furthermore, perceiving an
Università di Udine, via Margreth, 3, 33100 Udine, Italy external stimulus as well as planning the appropriate motor
e-mail: cosimo.urgesi@uniud.it
response takes time, inducing an intrinsic delay of brain
C. Urgesi · F. Fabbro reactions to external stimuli, which is incompatible with the
Istituto di Ricovero e Cura a Carattere ScientiWco dynamics of moving stimuli. Thus, to interact optimally
‘E. Medea’, Polo Friuli Venezia Giulia, with moving objects or creatures, the perceptual system
San Vito al Tagliamento, Pordenone, Italy
needs to build up an anticipatory representation of the
S. M. Aglioti motion sequence using internal models of the rules that
Dipartimento di Psicologia, govern the motion of objects in the environment (Hubbard,
Università Sapienza di Roma, Rome, Italy 2005; Motes, Hubbard, Courtney, & Rypma, 2008; Zago &
Lacquaniti, 2005). While visual experience with moving
S. M. Aglioti
Istituto di Ricovero e Cura a Carattere ScientiWco objects or creatures may contribute to the creation of pre-
Fondazione Santa Lucia, Rome, Italy dictive models of motion, we may additionally use our

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previous motor experience and knowledge to create an Overwalle & Baetens, 2009). Furthermore, recent studies
anticipatory representation of human actions (Flach, have reported action-speciWc unimodal visual and motor
Knoblich, & Prinz, 2004; Ramnani & Miall, 2004; Verfaillie (Chong, Cunnington, Williams, Kanwisher, & Mattingley,
& Daems, 2002). The fact that diVerent neurocognitive pro- 2008; Dinstein et al., 2010; Kilner, Neal, Weiskopf, Fris-
cesses are involved in the representation of predictive mod- ton, & Frith, 2009; Lingnau, Gesierich, & Caramazza,
els of human and non-human motion patterns is suggested 2009) as well as cross-modal adaptation of neural activity
by studies showing a compression of time perception in these fronto-parietal areas (Chong et al., 2008; Dinstein
and an increased precision of time discrimination during et al., 2010; Kilner et al., 2009), suggesting the same neural
observation of biological versus non-biological motion population may be involved in the representation of speciWc
(Moscatelli, Polito, & Lacquaniti, 2011; Carrozzo, actions that are either observed or executed. Importantly,
Moscatelli, & Lacquaniti, 2010; Orgs, Bestmann, Schuur, repeated execution of pulling or pushing actions may
& Haggard, 2011). Such eVects are reminiscent of the com- induce a bias to categorize ambiguous actions as opposite
pression of subjective time during voluntary actions (e.g., to the one that had been trained, likely through a motor-to-
Wenke & Haggard, 2009) and suggest that similar time per- visual adaptation of visuo-motor neurons (Cattaneo et al.,
ception processes are called into play in action execution 2010). Transcranial magnetic stimulation (TMS) over the
and observation. inferior frontal gyrus disrupted this visual aftereVect, sug-
The ability to create an anticipatory representation of gesting that this area contains mirror-like populations of
ongoing actions is crucial in sport, where, due to the dra- neurons that are recruited in action execution and observa-
matically fast times required by motor performance, ath- tion and may directly inXuence the visual perception of
letes must base their prediction on the initial action cues actions (Cattaneo et al., 2010). Accordingly, dysfunctional
provided by the opponent players. Indeed, waiting to fully activity of the inferior frontal cortex provoked by repetitive
perceive the consequences of their moves would be highly TMS or brain lesion is associated to impairments in diVer-
ineVective. It is well established that achieving excellence ent types of action perception tasks, including: discriminat-
in sport involves not only the ability to execute complex ing static hands with diVerent implied actions (Candidi,
actions but also to understand early the moves of the other Urgesi, Ionta, & Aglioti, 2008; Urgesi, Candidi, Ionta, &
players (Abernethy & Zawi, 2007; Aglioti, Cesari, Romani, Aglioti, 2007); estimating the weight of objects from the
& Urgesi, 2008). Much less is known on how superior per- observation of lifting actions (Pobric & Hamilton, 2006);
ceptuo-motor ability is established during sport learning. In recognizing the correct motor performance of transitive or
the present paper, we review previous evidence on the rela- intransitive gestures (Pazzaglia, Smania, Corato, & Aglioti,
tive role of visual and motor representations of actions and 2008a); matching viewed gestures to their typical sound
on the eVects of visual and motor learning on action execu- (Pazzaglia, Pizzamiglio, Pes & Aglioti, 2008b); and order-
tion and perception. Then, we present the results of two ing in the correct temporal sequence a series of snapshots
behavioral experiments on the role of observational and depicting the diVerent phases of an action (Fazio et al.,
physical practice on action prediction in volleyball. 2009). Thus, previous studies have shown not only that
observed and executed actions share a common neural rep-
Overlapping representations for action execution resentation, but also that the activity of the action represen-
and observation tations in the motor system aVects action perception
(Avenanti & Urgesi, 2011).
Neurophysiological and neuroimaging studies have pro-
vided evidence that action execution and action observation Visual representation of implied actions
share at least partially overlapping neural structures. The
direct demonstration in monkeys (Kraskov, Dancause, Information on the neural underpinnings of predictive
Quallo, Shepherd, & Lemon, 2009; Rizzolatti & Craighero, motion perception derives from studies of the brain activa-
2004; di Pellegrino, Fadiga, Fogassi, Gallese, & Rizzolatti, tions associated to the extrapolation of motion information
1992) birds (Prather, Peters, Nowicki, Mooney, 2008) and from static snapshots that depict objects or creatures in the
humans (Mukamel, Ekstrom, Kaplan, Iacoboni, & Fried, middle of motion. Viewing these implied motion stimuli
2010) of mirror neurons discharging both during action activates the same medial temporal cortex areas that are
execution and during observation of the very same actions involved in processing real motion (Kourtzi & Kanwisher,
has provided a strong evidence for the neural substrate of 2000; Proverbio, Riva, & Zani, 2009; Senior et al., 2000).
the action-perception common coding. Neuroimaging stud- Furthermore, studies in humans (Krekelberg et al., 2005;
ies have shown that action observation in humans engages a Lorteije et al., 2007) and monkeys (Krekelberg et al., 2003)
network of fronto-parietal areas which are also involved in suggest that the same populations of cells in extrastriate
action execution (Rizzolatti & Craighero, 2004; Van visual areas code for both implied and real motion.

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Importantly, TMS interference with medial temporal area from static images that imply body actions. Furthermore, in
abolishes the representational momentum eVect for objects a further single-pulse TMS study (Urgesi et al., 2010), we
(Senior, Ward, & David, 2002), i.e. the distortion of the showed that mirror motor facilitation may be selectively
recognition memory for the Wnal position or conWguration linked to the extraction of dynamic information about for-
of a moving object forward along its path of motion (Freyd, ward but not backward action paths. Indeed, observation of
1983). Thus, there is evidence that creating a full dynamic start and middle phases of hand actions engendered a sig-
representation of motion from viewing only single snap- niWcantly higher mirror motor facilitation than observing
shots may rely on the activation of the visual motion repre- their Wnal postures, independently from the Wnger conWgu-
sentations likely established during our visual experiences ration at diVerent hand apertures. These results are in keep-
with moving objects. ing with the Wndings that mirror motor facilitation is
Besides medial temporal cortex activation (Kourtzi & suppressed by the artiWcial introduction of delayed aper-
Kanwisher, 2000; Peigneux et al., 2000), observation of tures or sudden closure of Wngers during action observation
static body postures implying actions engenders activation (Gangitano, Mottaghy, & Pascual-Leone, 2004) and that
also in the parietal (Hermsdörfer et al., 2001) and superior motor activation has been found in response to symbolic
temporal sulcus (STS) areas (Peuskens, Vanrie, Verfaillie, cues signaling an upcoming movement (Kilner, Vargas,
& Orban, 2005). Furthermore, neurons in the monkey’s Duval, Blakemore, & Sirigu, 2004). Thus, mirror mapping
STS cortex respond to the presentation of both moving bod- seems to be inXuenced by the predictability of the temporal
ies and static images of body postures implying actions sequence of observed movements.
(Barraclough, Xiao, Oram, & Perrett, 2006; Jellema & A similar temporal modulation has been reported for the
Perrett, 2003a; Jellema & Perrett 2003b; Perrett et al., 2009; Wring properties of mirror neurons in monkeys’ ventral pre-
Puce & Perrett, 2003). Importantly, STS neurons respond- motor cortex, some of which discharge maximally during
ing to implied actions failed to respond to static postures observation of the last phases of grasping (Umiltà et al.,
that corresponded to the starting-point of the actions but 2001), others stop Wring when the target object has been
showed a strong response to the articulated static postures achieved, while others continue to discharge also during the
that corresponded to the end-point of the actions (Jellema & active holding phase (Gallese, Fadiga, Fogassi, & Rizzol-
Perrett, 2003b). In a similar vein, the response of some STS atti, 1996). Furthermore, in keeping with the response of
neurons to static body postures was inXuenced by which STS neurons (Baker et al., 2001), mirror neurons are acti-
action was previously observed, with a strong, selective vated also when the Wnal part of an object grasp action can-
response to a particular body action, independently from not be seen but only predicted from contextual information
the compatibility between the action end-points and the (Umiltà et al., 2001). Thus, mirror neurons may code the
static body posture (Jellema & Perrett, 2003a). This sug- outcome of goal-directed actions even when it is not appar-
gests that the Wring of STS neurons is inXuenced by the per- ent in the visual scene.
ceptual history of the action sequence in which a body However, information about hidden actions may be
posture is presented. Furthermore, STS neurons increased obtained from previous perceptual experience with the
their response after occlusion of the action (Baker, Keysers, action sequence or from the motor representation used dur-
Jellema, Wicker, & Perrett, 2001) suggesting that they ing action execution. The response of the STS neurons to
derive hidden information from the representations body posture is inXuenced by previous action perception
acquired during perception of the previous phases of the experiences (Jellema & Perrett, 2003a), suggesting these
action. neurons may use visual information to form a representa-
tion of ongoing actions (Perrett et al., 2009). This represen-
Motor representation of implied actions tation is purely visual, as STS neurons do not respond
during action execution (Rizzolatti & Craighero, 2004).
Recent studies have shown that perception of implied Conversely, mirror neurons matching action observation
motion of human bodies and body parts engenders activa- and execution may allow observers to use previous motor
tion not only in occipito-temporal areas, but also within the experience with similar actions in order to predict the future
motor system (Proverbio et al., 2009; Urgesi, Moro, Cand- course of impending actions ahead of their realization
idi, & Aglioti, 2006a). Using single-pulse TMS, we showed (Wilson & Knoblich, 2005).
that passive viewing of static images implying body actions
triggers mirror motor facilitation (Urgesi et al., 2006), sug- InXuence of visual and motor expertise on the action
gesting that the frontal node of the human mirror neuron observation network
system that matches action observation and execution (di
Pellegrino et al., 1992; Rizzolatti & Craighero, 2004) may Using motor representations to predict others’ actions is
play a major role in the extrapolation of dynamic information dependent upon the observers’ motor repertoire. While the

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studies on whether the action observation network responds immobilization or fatigue conditions). Behavioral studies in
also to robotic movements have given controversial Wnd- the Weld of motor control have shown that observational
ings, several neuroimaging studies have shown that the practice of movements leads to subsequent improved motor
activation of the action observation network is modulated performance (Ashford, Bennett, & Davids, 2006), although
by the observer’s motor experience (Calvo-Merino, Glaser, the extent of the beneWt may be lower as compared to physi-
Grèzes, Passingham, & Haggard, 2005; Calvo-Merino, cal practice. Thus, action observation may induce the forma-
Grèzes, Glaser, Passingham, & Haggard, 2006; Cross, tion of a motor memory trace also in the absence of actual
Hamilton, & Grafton, 2006; Cross, Kraemer, Hamilton, muscle contraction and perception of sensory feedback (Ste-
Kelley, & Grafton, 2009a; Orgs et al., 2008; Reithler et al., fan et al., 2005). Interference with primary motor cortex
2007). In particular, observing dance moves engenders activity disrupts the consolidation of motor memories
greater activation of temporal and fronto-parietal areas in acquired after physical (Muellbacher et al., 2002) and obser-
expert dancers than in participants naïve to the observed vational (Brown, Wilson, & Gribble, 2009) practice, thus
dance style (Calvo-Merino et al., 2005). Since, however, suggesting that both types of training induce changes of
not only motor but also visual experience is acquired during movement representation in primary motor cortex (Censor &
action performance, in a subsequent study the authors Cohen, 2011). Interestingly, several movement properties
(Calvo-Merino et al., 2006) compared the activations of the may be acquired during observational practice, including
action observation network of male and female dancers coordination patterns (Boutin, Fries, Panzer, Shea, & Blan-
during observation of gender-speciWc dance moves. This din, 2010; Gruetzmacher, Panzer, Blandin, & Shea, 2011;
allowed them to control for the relative role of visual expe- Heyes & Foster, 2002; Vinter & Perruchet, 2002), timing
rience, which was common to both types of moves, and (Hayes, Timmis, & Bennett, 2009; Hayes, Elliott, & Bennett,
motor experience, which was speciWc for one’s own gender 2010; Ong & Hodges, 2010) and force scaling (Mattar &
moves. The authors found that premotor, parietal and cere- Gribble, 2005; Porro, Facchin, Fusi, Dri, & Fadiga, 2007).
bellum activations were higher when participants viewed Furthermore, like for physical training, the observation-
dance moves belonging to their’ own gender repertoire, related improvements in a speciWc motor task can be
while temporal areas were not modulated. transferred to a diVerent one (Buchanan & Wright, 2011;
Similar results were obtained in another series of neuro- Gruetzmacher et al., 2011; Hayes et al., 2010; Mattar &
imaging investigations that compared the changes in the Gribble, 2005; Ong & Hodges, 2010; Shea, Wright, Wulf, &
activation of the action observation network after observa- Whitacre, 2000). Importantly, however, motor skill acquisition
tional and physical learning of dance moves (Cross et al., is not identical when mediated by physical or observational
2006; Cross et al., 2009a; Cross, Hamilton, Kraemer, Kel- practice, as suggested by the Wnding that their combination
ley, & Grafton, 2009b). Largely overlapping increases of resulted in a greater advantage on transfer tasks (Shea et al.,
activation, mainly in right premotor and left inferior parie- 2000). Advantages of physical practice, for example, transfer
tal areas, were found during observation of trained and to motor tasks with analogous motor coordinates (i.e., using
watched moves (as compared to untrained moves) (Cross the opposite limb to control the same pattern of muscle con-
et al., 2009a). Moreover, speciWc modulation for the trained tractions and joint angles). In contrast, advantages of obser-
as compared to the watched moves was found in two areas vational practice transfer to tasks with diVerent motor
of the right dorsal premotor cortex. The opposite contrast coordinates but analogous visuo-spatial coordinates (i.e.,
did not reveal any speciWc activation for the watched using the opposite limb to control movements with the same
moves. These results suggest that action execution may direction in the external space) (Gruetzmacher et al., 2011).
shape the fronto-parietal sectors of the action observation Moreover, unlike execution, observation of visuo-motor
network more intensively and extensively than action adaptation to reaching movements in a perturbed environ-
observation (Cross et al., 2009a). ment did not induce any after-eVects (i.e. the unwanted per-
sistence of compensatory movements in an environment that
InXuence of visual and motor expertise on action execution do not require such compensations) (Ong & Hodges, 2010).
This suggests that observational practice does not allow
The notion of common representations of executed and updating internal sensorimotor models of actions.
observed actions has a notable interest in applicative Welds
like neurorehabilitation (Garrison, Winstein, & Aziz-Zadeh, InXuence of visual and motor expertise on action
2010; Small et al., 2010) and sport learning (Holmes & perception
Calmels, 2008). Indeed, the possibility to acquire new
motor skills through action observation may be helpful when It is well known that visual experience improves discrimi-
physical practice cannot take place (e.g. in injury-related nation of visual stimuli (Sasaki, Nanez, & Watanabe, 2010),

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including body actions (Calvo-Merino, Urgesi, Orgs, Zawi, 2007; Aglioti et al., 2008). Previous studies on this
Aglioti, & Haggard, 2010; Maslovat, Hodges, Krigolson, & issue (Abernethy & Zawi, 2007; Abernethy et al., 2008;
Handy, 2010). This may not be surprising because action Aglioti et al., 2008; Farrow & Abernethy, 2003; Weissen-
execution is typically coupled with the visual perception of steiner et al., 2008) used temporal occlusion paradigms
one’s own movements (Maslovat et al., 2010). Importantly, where video presentation was interrupted at diVerent
action execution could improve the visual discrimination of instants after the beginning of the action, showing videos of
full (Hecht, Vogt, & Prinz, 2001) and point-light (Casile & increasing length. In these conditions, the presentation of
Giese, 2006) displays of the same action even if no visual body- versus ball-related cues was confounded with
feedback was provided during the execution phase. Further- increasing viewing time and cumulative presentation of
more, visual perception improvements were obtained after more information available on the action. Thus, it could not
active as well passive movement execution (Hecht et al., be established whether the superior perceptual abilities of
2001), thus suggesting that the aVerent kinesthetic signals motor experts reXected faster processing speed and need of
are suYcient to establish a visual representation of the less information to make a decision or were speciWcally
action. related to their capacity to read body kinematics. Here we
Studies of motor and visual experts suggest that motor used a modiWed temporal occlusion paradigm in which vol-
experience is extremely relevant for action perception. In leyball Xoating services were presented. Importantly, only
particular, behavioral investigations of elite athletes have the body movements or only the ball trajectory were
shown that motor experts present superior abilities not only shown. This procedure allowed us to highlight the comple-
in the execution of complex actions, but also in the predic- mentary roles of motor and visual expertise on the repre-
tion and anticipation of the behavior of other players (Aber- sentation of body action and object motion, respectively.
nethy & Zawi, 2007; Abernethy, Zawi, & Jackson, 2008; We expected that only motor experts should be able to pre-
Aglioti et al., 2008; Farrow & Abernethy, 2003; Weissen- dict the outcome of the Xoating service by observing the
steiner, Abernethy, Farrow, & Müller, 2008). These studies initial body movements, while watchers should not show
have typically used a temporal occlusion paradigm, in any advantage as compared to novices. On the other hand,
which the presentation of sport actions is interrupted at both athlete and watcher groups are expected to outperform
diVerent delays from onset. Domain-speciWc motor experts novices in the perception of the ball trajectory. Indeed, their
and naive participants are required to predict the direction previous visual experience with volleyball should allow
or the correctness of the action after viewing only the initial athletes and watchers to build more accurate models of the
body kinematics of the model player or also the visual object motion as compared to that of novices, who can use
consequences of the movements (e.g., ball trajectory). only general internal models of the object motion under
Research in a variety of diVerent sports shows that motor 1-G gravity conditions to understand the ball trajectory
experts are more accurate than expert observers (e.g., (Zago & Lacquaniti, 2005).
coaches) and naïve participants in predicting the fate of the The second aspect explores whether the congruence
observed actions after viewing the initial body movements. between the spatial perspective of the observer and of the
This suggests elite athletes ‘read’ the body kinematics of model (Alaerts, Heremans, Swinnen, & Wenderoth, 2009;
the observed actions (Abernethy & Zawi, 2007; Smeeton & Maeda, Kleiner-Fisman, & Pascual-Leone, 2002; Urgesi,
Huys, 2010). Crucially, in a previous study we showed that Candidi, Fabbro, Romani, & Aglioti, 2006b; Christensen,
the superior predictive abilities of elite basketball players Ilg, &, Giese 2011, this issue) inXuences action prediction.
with respect to naïve and expert observers were associated To this aim, we tested whether the action knowledge
to diVerential motor activation during observation of the acquired by doing or watching inXuenced diVerently the
early phases of erroneous versus correct shots (Aglioti perception of the actions performed by back-facing models
et al., 2008). Thus, achieving excellence in sports might be (viewed according to a Wrst person perspective), or by
related, in part, to the Wne-tuning of speciWc anticipatory front-facing models (viewed according to a third-person
‘resonant’ mechanisms that allow for an earlier and more perspective). Previous studies have suggested that motor
accurate prediction of the future of others’ actions. activation during action observation is higher for actions
viewed from a spatial perspective which is compatible
Aims of the present study rather than incompatible with the actor’s perspective (Ala-
erts et al., 2009; Maeda et al., 2002; Urgesi et al., 2006b).
In two psychophysics studies, we explored action observa- Furthermore, observers are more accurate in perceptual pre-
tion and prediction in volleyball. Three novel, important dictions for videos showing self-produced actions than for
aspects of the issue were investigated. The Wrst concerns videos showing similar actions produced by other individu-
the role of body or context cues in shaping diVerent predic- als (Knoblich & Flach, 2001; Knoblich, Seigerschmidt,
tion strategies in motor and visual experts (Abernethy & Flach, & Prinz, 2002). In keeping with the notion that

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observed actions are perceived using internal models of the mean of 14.58 § 4.3 years (athletes; mean age = 24.33
sensory consequences of executed actions (Hommel et al., years, SD = 5.3); 12 supporters who had never practiced
2001; Prinz, 1997), these results suggest that action predic- volleyball but attended continuously to volleyball matches
tion is maximally facilitated for close match between pre- for at least 10 years (watchers; mean age = 27.5 years,
dicted and actual action outcomes like in the case of actions SD = 8.97); 12 age-matched participants with no experi-
that are self-produced or belonging to the observers’ motor ence with volleyball (novices; mean age = 29.42 years,
repertoire. Since we assume that the predicted sensory con- SD = 3.5). All participants were right-handed according to
sequences of executed actions match to a greater extent the a standard handedness inventory (Briggs & Nebes, 1975),
back- than the front-facing actions, we expected that the reported normal or corrected-to-normal visual acuity in
perceptual advantage of athletes, who may use internal both eyes and were naive about the purposes of the experi-
models of actions established during action execution, ment. Participants gave their written informed consent and
should be maximal for the back-facing action videos. This received information about the experimental hypothesis
eVect should be speciWc for motor experience with body only after the experimental tests were completed. The
movements. In contrast, the inXuence of visual experience procedures were approved by the ethics committee of the
on the athletes’ and watchers’ ability to perceive the ball ScientiWc Institute (IRCCS) “E. Medea” and were in accor-
trajectory should be comparable for back- and front-facing dance with the ethical standards of the 1964 Declaration of
conditions, because both groups may have learned visual Helsinki.
models of the ball trajectory from Wrst- and third perspec-
tives. Failing to Wnd any speciWc modulation of spatial per- Stimuli and task
spective on the prediction abilities of motor experts would
suggest that internal action models are not constrained by Two female expert model players (aged 18 and 20 years)
spatial perspective. were videotaped while they performed series of volleyball
Finally, we compared longitudinally the ability of ado- Xoating services directed to the right or to the left of the
lescents practicing volleyball to predict the outcome of opposite court. The videos were taken from the back (back
domain-speciWc actions before and after a physical, obser- view) and from the front (front view) of the shooting
vational or control training. In experiment 2, comparable player. The model players were instructed to direct the
practice periods for the physical and observational training shots to the farthest border of the court, thus making more
were given. This allowed us to establish whether the supe- likely the execution of services in which the ball landed in
rior perceptual abilities of athletes are speciWcally related to (In-shots) or out (Out-shots) the court. For each model and
motor practice or simply due to their greater visual experi- viewing perspective, we selected 8 In-shots (4 directed to
ence with domain speciWc actions. Furthermore, to match the left and 4 to the right of the court) and 8 Out-shots
the instructions given to the subjects, the same verbal (4 directed to the left and 4 to the right of the court). Thus,
instructions were given to both execution and observation a total of 64 videos were used. Videos were presented in a
groups. Finally, the videos used in the observational train- modiWed temporal occlusion paradigm (Abernethy & Zawi,
ing were focused on both the moving bodies and the ball 2007; Aglioti et al., 2008). Using the Adobe Premiere soft-
motion. This procedure allowed us to avoid that any diVer- ware (Adobe Systems Incorporated, San Jose, CA), each
ential perceptual eVects of the physical and observational video was split into two 1,250-ms clips with respect to the
training might be ascribed purely to relative attention to the frame showing the hand-ball contact. Half of the clips
body or to the ball in the two groups. Overall, our approach lasted from the beginning of the action to the instant at
allowed us to explore the speciWc inXuence of motor and which the hand touches the ball, thus showing only the
visual experience on the plastic modeling of action predic- body kinematics; the other half from the frame showing the
tion abilities. hand-ball contact to the initial falling trajectory of the ball
(when it was approximately at the height of the net), thus
showing only the ball trajectory (Fig. 1). This resulted into
Experiment 1 presentation of a total of 128 clips: 2 models £ 2
directions £ 2 In- or Out-shots £ 2 cues £ 8 shot exem-
Methods plars.

Participants Procedure

In experiment 1, we tested three groups of female right- Each participant was tested in a single experimental session
handed adult individuals (age 27.1, range 18–45 years): 12 lasting approximately 45 min. Back-view and front-view
volleyball athletes (C or B series) practicing volleyball for a clips were presented in two separate 64-trial blocks and the

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Fig. 1 Examples of the starting snapshots of videos showing back- models are shown in the two perspectives. The depicted exemplars are
and front-facing body kinematics and ball trajectory. Two diVerent taken from In shots

order of the blocks was counterbalanced between subjects. anced between participants. Participants were instructed to
A short rest was allowed before proceeding to a diVerent be as accurate and fast as possible. Response accuracy and
block. The order of trials with In- and Out-shots, left and the time taken to respond from the onset of the prompt
right services, and body and ball cues was randomized. frame (latency) were recorded and analyzed.
Before proceeding to each experimental block, participants
completed a 5-trial practice block, with random selection of Data analysis
trials from the diVerent experimental conditions.
Participants sat 57 cm away from a 15-inch monitor (res- We calculated the percentage of correct responses (accu-
olution, 1,024 £ 768 pixels; refresh frequency, 60 Hz), racy) and the mean latency of each participant in each
where the video stimuli appeared on a black background experimental condition (32 trials per cell). Individual accu-
and subtended a 22.5° £ 15° region. Stimulus-presentation racy and mean latency values were entered into two sepa-
timing, randomization, and data recording were controlled rate mixed model ANOVAs with group (athletes, watchers,
by using E-prime V1.2 software (Psychology Software novices) as between-subjects variable and viewing perspec-
Tools Inc., Pittsburgh, PA) running on a laptop computer. tive (back, front) and cue (body, ball) as within-subjects
Movie presentation was obtained by presenting 25 snap- variables. All pair-wise comparisons were performed using
shots at a 20 Hz rate for a total of 1,250 ms for each stimu- the Duncan post-hoc test. A signiWcance threshold of
lus clip. P < 0.05 was set for all statistical analyses. Data are
A trial started with the presentation of a central 3° £ 3° reported as mean § standard error of the mean (SEM).
Wxation cross lasting 500 ms, which was followed by the
presentation of the stimulus clip at the centre of the moni-
tor. At the end of clip presentation a prompt frame was pre- Results
sented requesting the participants to press, respectively
with the left or the right index Wnger, the left or the right Accuracy
button of the computer track pad to decide whether the shot
was in or out of the Weld. Association between in- or out- The ANOVA on accuracy (Fig. 2a) revealed signiWcant
responses and left or right button presses was counterbal- main eVect of group (F2,33 = 18.95, P < 0.001, p2 = 0.534),

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Psychological Research (2012) 76:542–560 549

Fig. 2 Results of experiment 1. The graphs show the mean accuracy back- (left panels) or front-facing (right-panels) perspectives. Error
(a) and latency (b) of adult athletes, expert watchers and novices in bars indicate standard errors. Asterisks indicate signiWcant between-
predicting the fate of volleyball Xoating services viewed from the group diVerences in the diVerent experimental conditions

with athletes (72.23%) performing better than watchers outperformed novices in all conditions (all Ps < 0.04). In
(60.58%; P < 0.001) and novices (56.33%; P < 0.001), who contrast, watchers outperformed novices only in under-
in turn had comparable performance (P = 0.122). The standing the ball trajectory in the back- (P = 0.007), but not
eVects of viewing perspective (F1,33 = 4.76, P = 0.036, in the front-view clips (P = 0.383). The performance of
p2 = 0.126), cue (F1,33 = 19.8, P < 0.001, p2 = 0.375) and watchers and novices was comparable for the predictions
their two-way interaction (F1,33 = 7.07, P = 0.012, based on body kinematics, independently from the viewing
p2 = 0.176) were also signiWcant. Performance was better perspective (both Ps > 0.26). Better performance of athletes
for ball than for body cue clips when they were taken from as compared to watchers was observed for predictions
the back- (70.06 vs. 59.67%; P < 0.001) but not when they based on reading the kinematics of the model’s body when
were taken from the front-view perspective (62.58 vs. viewed from the back (P = 0.01) but not when viewed from
59.89%; P = 0.197). Furthermore, participants were more the front (P = 0.112). On the other hand, athletes were bet-
accurate for the back- than for the front-view perspective ter than watchers in predicting the ball trajectory in both
when clips showed the ball trajectory (P = 0.001), but not viewing perspectives (both Ps < 0.029).
when clips showed the body movements (P = 0.914). The
cue £ group (F2,33 < 1, p2 = 0.025) and the viewing Latencies
perspective £ group interactions (F2,33 = 2.259, P = 0.132,
p2 = 0.116) were non-signiWcant. Crucially, however, a The ANOVA on latencies (Fig. 2b) revealed a signiWcant
signiWcant three-way interaction was found (F2,33 = 5.35, main eVect of group (F2,33 = 8, P = 0.001, p2 = 0.326),
P = 0.01, p2 = 0.245). Pair-wise post-hoc tests revealed with novices (2,437.59 ms) being overall slower than ath-
better performance in perceiving the ball trajectory in the letes (1,764.03 ms; P < 0.001) and watchers (2,072.91 ms,
back-view perspective as compared to the other conditions P = 0.038), while the diVerence between athletes and
for athletes (all Ps < 0.003) and watchers (all Ps < 0.004), watchers did not reach signiWcance (P = 0.076). The eVects of
but not for novices (all Ps > 0.35). Furthermore, athletes viewing perspective (F1,33 = 13.54, P = 0.001, p2 = 0.291),

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550 Psychological Research (2012) 76:542–560

cue (F1,33 = 395.85, P < 0.001, p2 = 0.923) and their provided by initial and last part of the ball trajectory reX-
interaction (F1,33 = 17.75, P < 0.001, p2 = 0.35) were also ected speciWc visual or motor expertise. Both athletes and
signiWcant. Post-hoc tests showed that ball cue clips were watchers were most accurate when they predicted the fate
responded to faster than body cue clips in both back- of the shots on the basis of the ball trajectory viewed from
(1,834.07 ms vs. 2,486.66 ms; P < 0.001) and front-view the back, while novices were similarly inaccurate in under-
perspectives (1,593.69 ms vs. 2,451.61 ms; P < 0.001). standing back- and front-viewed ball trajectories. Further-
However, participants were faster for the front- than back- more, although, athletes were better than novices and
view perspective when clips showed the ball trajectory watchers in understanding the ball trajectory in both
(P < 0.001), but not when videos showed the body move- perspectives, watchers outperformed novices only in the
ments (P = 0.317). The eVects of cue and viewing perspec- back-view perspective. This would suggest that the visual
tive on latencies were comparable for the three groups, experience gained by watchers improved only their ability
since non-signiWcant two- or three-way interaction with to use the last part of the ball trajectory, when the ball was
group was obtained (in all cases, F2,33 < 2.1, P > 0.14, falling down to the Xoor. In contrast, athletes could use also
p2 < 0.12). the initial part of the ball trajectory which was visible in
both the back- and front-view videos.
In contrast to the strong eVect of the viewing perspective
Discussion on the ability to understand the ball trajectory, responses
based on reading the body kinematics were not aVected by
In experiment 1 we compared the action prediction abilities the viewing perspective in any group. Importantly, how-
of adult athletes, who had both motor and visual expertise, ever, while athletes outperformed watchers in understand-
watchers, who had only visual expertise, and novices, who ing the ball trajectory of videos taken from both back- and
had no experience with the observed actions. Stimuli front-viewing perspectives, they were better than watchers
showed the body movement or the ball trajectory phases of in reading the body kinematics only for videos taken from
volleyball Xoating services viewed from the back or from the back-view perspective, which was compatible with the
the front. This allowed the manipulation of the recognition Wrst-person experience of the observed actions. This would
cue and of the correspondence between the model player’s suggest that motor experience with actions played a major
and the observer’s viewing perspectives. The results role in determining the superior perceptual abilities of ath-
showed that athletes outperformed novices in predicting the letes. Thus, the results of experiment 1 are in keeping and
outcome of actions on the basis of body kinematics and ball further corroborate previous studies (Abernethy & Zawi,
trajectory, independently from the viewing perspective. It is 2007; Abernethy et al., 2008; Aglioti et al., 2008; Farrow &
worth noting that watchers outperformed novices only in Abernethy, 2003; Weissensteiner et al., 2008) showing that
predictions based on the ball trajectory when videos were expert athletes, but not expert watchers, are equipped with
taken from the back-view perspective. fast and generally accurate perceptual mechanisms. These
Back- and front-view videos diVered in the possibility to mechanisms allow experts to read the kinematics of others’
see the last part of the ball trajectory, which was partially body movements and the initial phases of the resulting
hidden in the front-view perspective. This inXuenced the object motion in order to anticipate the future course of
performance of athletes and watchers in predicting the fate observed actions ahead of realization. Importantly, rather
of the shots on the basis of the ball trajectory, with less than presenting videos of increasing length as in the typical
accurate and faster responses to front- than to back-view temporal occlusion paradigm used in previous studies
ball trajectory clips. The trade-oV between accuracy and (Abernethy & Zawi, 2007; Abernethy et al., 2008; Aglioti
latency for front- as compared to back-view videos might et al., 2008; Farrow & Abernethy, 2003; Weissensteiner
reXect a change of the response criterion due to the absence et al., 2008), we presented clips with a Wxed duration and
of relevant information from the last part of the videos. showing only the body kinematics or only the ball trajec-
Since the two perspectives were presented in a blocked tory. This allowed us to relate the superior perceptual abili-
design, in the absence of important information from the ties of athletes to their use of body movement information.
last part of front-ball videos participants might have ana- However, athletes outperformed watchers also in under-
lyzed only the initial part and planned earlier a response to standing the ball trajectory and not only in reading the body
be provided after the appearance of the prompt frame. Cru- movements. This last result is in keeping with our previous
cially, this strategic use of the visual information to adjust study (Aglioti et al., 2008) with basketball players, who
the velocity of response was non-speciWc for the level of provided better predictions than watchers when videos
motor and/or visual expertise, since all the three groups of showed only the initial body movements as well as when
participants showed similar eVects on response latencies. In they showed the initial body movements and the ball trajec-
contrast, their ability to use appropriately the information tory. While the pattern of results of the above study could

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Psychological Research (2012) 76:542–560 551

be ascribed to the combination of body and object cues in Group allocation procedure
longer videos, the present results show that experts are bet-
ter than watchers also in perceiving the ball trajectory pre- Participants were pseudorandomly allocated to three train-
sented in isolation. This suggests an overall better ing groups using a matching procedure based on their per-
performance of motor experts also in the visual representa- formance in a preliminary Xoating service execution test.
tion of the ball trajectory. This ensured comparable baseline levels in action execu-
The diVerence between expert athletes and watchers may tion abilities. In a preliminary session, participants were
reXect either the greater extent of athletes’ visual experi- administered a Xoating service execution test. Placed in a
ence with volleyball actions, which was higher than that of central position behind the bottom line of the volleyball
supporters, or the additive combination of visual and motor court (9 m from the net), participants performed 32 Xoating
experience allowing a better description of movements and services to the opposite court. According to a pseudoran-
their visual consequences. Comparing diVerent groups of dom sequence determined by the examiner, 16 services
individuals with acquired domain speciWc expertise, should be directed to the left (zone 1) and 16 services to the
indeed, allows testing the eVect of extensive practice peri- right (zone 5) of the opposite court. The net height was set
ods, which can be hardly used in longitudinal studies, but at 224 cm. Each service received a 3-point score: 0, wrong
prevents to clearly disentangle the speciWc roles of motor service; 1, service in court but in the wrong zone; 2, service
and visual experience in action perception. Furthermore, in court in the correct zone. Participants were divided into
studies based on comparison of diVerent groups do not take three groups on the basis of their total precision score in the
into account the possible preexisting interindividual diVer- test (mean = 11.31, SD = 14.41): low (precision scores: 0;
ences that may have determined the superior abilities of N = 19), intermediate (precision scores: 1–17; N = 13), and
athletes in sport as compared to novices or watchers. To high performers (precision scores: 18–45; N = 13). Partici-
clarify the speciWc roles of motor and visual expertise on pants from each of three performance groups were ran-
the Wne tuning of anticipatory perceptual abilities, in exper- domly selected and allocated sequentially to the three
iment 2 we adopted a longitudinal approach comparing the training groups: execution (N = 15, 13 females; mean
perceptual abilities of adolescents practicing volleyball age = 12.13 years, SD = 1.36), observation (N = 15, 14
before and after a physical or a observational training about females; mean age = 12 years, SD = 1.07), control (N = 15,
the Xoating services in volleyball. 14 females; mean age = 11.73 years, SD = 1.1). This
ensured a comparable presence of high, intermediate and
lower performers in the three training groups, with a resulting
Experiment 2 comparable baseline execution performance (F2,42 = 1.42,
P = 0.252).
Methods
Training procedure
Participants
After the initial testing session, twelve 2-h training sessions
We recruited from a local volleyball sport club 45 (41 were administered to each group, with a frequency of
females) right-handed children aged 10–14 years (mean 4 days per week for 3 weeks. Each training session con-
age = 11.96 years, range 10–14 years) with no previous for- sisted of a common training (90 min) and a diVerentiated
mal training on volleyball Xoating service. All participants training (30 min). During the common training, the individ-
were right-handed according to a standard handedness uals of all groups participated together to a standard volley-
inventory (Briggs & Nebes, 1975), reported normal or cor- ball training with no practice or reference to Xoating
rected-to-normal visual acuity in both eyes and were naive services. The common training was aimed at allowing par-
about the purposes of the experiment. Participants gave ticipants to continue regularly their volleyball practice in a
their informed consent and their parents gave written controlled setting that matched the general experience of
informed consent for participation in the research. Detailed the three groups with volleyball actions. During the diVer-
information on the phases and procedures of the study were entiated training, each group received separately a speciWc
provided at the time of recruitment, but participants and intervention. The execution group was involved in a tradi-
their parents received information about the experimental tional training requiring the repeated performance of Xoat-
hypothesis only after the study phases were completed. The ing services after presentation of a model and following a
procedures were approved by the ethics committee of the series of instructions on how to perform the Xoating ser-
ScientiWc Institute (IRCCS) “E. Medea” and were in accor- vice. The observation group was involved in an observa-
dance with the ethical standards of the 1964 Declaration of tional training in which participants watched videos of
Helsinki. Xoating services and heard the same instructions provided

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552 Psychological Research (2012) 76:542–560

to the execution group. The control group watched videos Results


of volleyball defense actions in which the Xoating services
were cut out and heard instructions on the volleyball Accuracy
actions.
The videos for the observational and control training The four-way ANOVA on accuracy revealed a signiWcant
were created for this experimental research and focused on four-way interaction between group, viewing perspective,
the technical movements and game of volleyball. They had cue, and session (F2,42 = 3.29, P = 0.047, p2 = 0.136). The
a total duration of 30 min and showed a series of action follow-up three-way ANOVA for the back-view condition
scenes that were presented at natural speed or slowed down (Fig. 3a) revealed a signiWcant main eVect of cue
by 25% to improve understanding of the action dynamics. (F1,42 = 30.04, P < 0.001, p2 = 0.417), with better perfor-
The videos showed two expert female athletes as technical mance for ball (75%) than body videos (64.33%). No other
models taken from the front or from the back-viewing per- main eVects or two-way interactions were signiWcant (all
spective. The video on the Xoating service in volleyball that Fs < 2.73, Ps > 0.07, p2s < 0.12). Crucially a signiWcant
was presented to the observation group reproduced the three-way interaction was obtained for the back-view con-
whole or a partial dynamic of diVerent Xoating services, dition (F2,42 = 5.5, P = 0.008, p2 = 0.207). Duncan post-
with 360-grade shots. The video allowed the observer to hoc test showed that, after the training, the performance of
understand the key elements of the service technique. The the execution group improved in predicting the fate of the
visual presentation of the actions was accompanied by ver- shots on the basis of body kinematics (62.5 § 3.24% vs.
bal descriptions explaining the appropriate movements. The 68.54 § 2.9%; P = 0.041), but not on the basis of the ball
same verbal descriptions were given to the observation and trajectory (74.79 § 3.74% vs. 75.63 § 3.62%; P = 0.754).
execution groups. The video on the defense in volleyball In contrast, the observation group improved in predic-
that was presented to the control group showed defense tions based on the ball trajectory (71.83 § 3.15% vs.
game actions, with the service phase being hidden from 80.67 § 2.74%; P = 0.004), but not on the body kinemat-
view. Also in this video, the visual presentation of the ics (65.83 § 2.72% vs. 62.46 § 3.25%; P = 0.253). No
defense actions was accompanied by verbal descriptions change was, instead, observed for the control group in
explaining the appropriate movements. reading body kinematics (65 § 3.81% vs. 61.67 § 3.91%;
P = 0.259) or in understanding the ball trajectory
Stimuli and task (74.38 § 4.85% vs. 72.71 § 5.22%; P = 0.532). The fol-
low-up three-way ANOVA for the front-view condition
The eVects of the three types of interventions were evalu- (Fig. 4a) revealed non-signiWcant main eVects (all
ated by administering before (Pre) and after (Post) training, Fs < 3.8, Ps > 0.05, p2s < 0.09) or interactions (all
the same task and stimuli used in experiment 1. Pre- and Fs2,42 < 1).
post-training evaluations were conducted at an interval of at
least 1 month. Latencies

Data analysis The four-way ANOVA on response latencies (Figs. 3b, 4b)
showed signiWcant eVects of viewing perspective
We calculated the percentage of correct responses (accu- (F1,42 = 19.25, P < 0.001, p2 = 0.314), cue (F1,42 = 542.94,
racy) and the mean latency of each participant in each P < 0.001, p2 = 0.928) and of their interaction
experimental condition. Individual accuracy and mean (F1,42 = 30.48, P < 0.001, p2 = 0.421). In contrast, the
latency values were entered into two separate mixed model other main eVects and two-, three-, and four-way interac-
ANOVA with group (execution, observation, control) as tions did not reach signiWcance (all Fs < 3.38, Ps > 0.07,
between-subjects variable and viewing perspective (back, p2s < 0.08). Participants were faster in predicting the fate
front), cue (body, ball), and session (pre, post) as within- of the shots by understanding the ball trajectory than by
subjects variable. A signiWcant four-way interaction was reading the body kinematics when videos where taken from
further explored by means of two separate group £ the back- (1,882.25 ms vs. 2,360.88 ms; P < 0.001) and
cue £ session mixed-model ANOVAs, conducted sepa- front-view perspectives (1,663.95 ms vs. 2,286.12 ms;
rately for the back- and front-view conditions. All pair-wise P < 0.001). Furthermore, participants were faster for the
comparisons were performed using the Duncan post-hoc front- than for the back-view perspective, independently
test. A signiWcant threshold of P < 0.05 was set for all sta- from cue (both Ps < 0.001), but the eVect of perspective
tistical analyses. Data are reported as mean § standard was stronger for understanding the ball trajectory than for
error of the mean (SEM). reading body kinematics.

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Psychological Research (2012) 76:542–560 553

Fig. 3 Results of experiment 2 relative to the back-view stimuli. The Error bars indicate standard errors. Asterisks indicate signiWcant
graphs show the mean accuracy (a) and latency (b) in predicting the between-sessions and between-cues diVerences in the diVerent experi-
fate of volleyball Xoating services viewed from the back-view perspec- mental groups
tive before and after the execution, observation and control trainings.

Discussion corresponds to the pattern of performance of adult expert


watchers tested in experiment 1, who were better than nov-
In experiment 2 we performed a psychophysics investiga- ices in understanding the ball trajectory shown from back-,
tion of sport-speciWc action prediction abilities in adoles- but not from front-view perspective. Furthermore, the
cents before and after the acquisition of perceptual and motor experience of the execution group may have not been
motor skills related to volleyball. We found a double disso- eVective in allowing a better understanding of the model’s
ciation between physical versus observational practice and body movements when viewed from the front-view, third-
improvements in predicting the fate of observed actions on person perspective. Indeed, even after several years of
the basis of body kinematics versus ball trajectory. This practice, adult athletes tested in experiment 1 did not out-
double dissociation rules out that the spurious eVects of perform watchers in reading the body kinematics when the
task diYculty and training complexity might explain the model’s body was viewed from the front. Importantly, the
selectivity of the eVects of the two training procedures on improvement of the execution group was selective for pre-
the two tasks. Improvements in accuracy were not accom- dicting the fate of others’ actions by reading their body
panied by changes in latency of responses, thus ruling out movements, and did not extend to the performance in
speed-accuracy trade-oV eVects. Furthermore, the eVects of understanding the ball trajectory, which was improved only
the training were speciWc for the back view condition. That after observational practice. Thus, the results of experiment
no speciWc change was obtained in the front-view condition 2 allowed us to better clarify the results of experiment 1 on
may be due to the fact that the ball was not fully visible adult athletes, who were better than novices in perceiving
when the videos were recorded from a front-view perspec- the body kinematics and ball trajectory. Indeed, since ath-
tive and thus the observation group could not use their letes had a greater motor and visual expertise with volley-
improved abilities in understanding the ball trajectory. This ball actions as compared to novices and also to expert

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554 Psychological Research (2012) 76:542–560

Fig. 4 Results of experiment 2 relative to the front-view stimuli. The tive before and after the execution, observation and control trainings.
graphs show the mean accuracy (a) and latency (b) in predicting the Error bars indicate standard errors. Asterisks indicate signiWcant
fate of volleyball Xoating services viewed from the front-view perspec- between-cues diVerences in the diVerent experimental groups

watchers, we can attribute their higher body movement per- manipulation of the type of training in experiment 2 makes
ception abilities to their motor experience, and their higher unlikely that diVerential attentional deployment in execu-
ball trajectory perception abilities to their visual experi- tion and observation training may inXuence action predic-
ence. In contrast, expert watchers could use only their tion abilities. Future studies, using for example eye
visual experience to understand the ball trajectory, but not movement recording, are needed to explore systematically
the body kinematics. Importantly, it is unlikely that the the type of information used by execution and observation
eVects of experiment 2 might be simply ascribed to the trainees during action demonstrations.
diVerential attention to body or to ball cues paid during the
training by the participants of the execution and observa-
tion groups. Indeed, while in natural learning contexts one General discussion
cannot control for which aspects of the performance are
attended to by players and observers, in experiment 2 we Several studies indicate that action perception and execu-
controlled for the type of information presented to the tion rely upon largely overlapping cognitive systems
observation group participants. It is worth noting that the (Hommel et al., 2001; Prinz, 1997) and neural substrates
observation training videos directed the participants’ atten- (Grafton, 2009; Rizzolatti & Craighero, 2004). It is less
tion to body movements as well as to their consequences on clear, however, how shared representations of action execu-
the ball trajectory. Furthermore, all the participants were tion and perception develop and whether visual and motor
motivated to learn to perform the volleyball Xoating service experiences are equivalent in their ability to establish novel
and likely attended to the demonstrations of its accurate perceptuo-motor representations of actions.
body kinematics. We cannot rule out that attentional factors In the present study we aimed to investigate the diVeren-
may play a role in building the diVerent perceptual abilities tial contribution exerted by motor and visual experience to
of athletes and watchers. However, our experimental the development of the experts’ abilities to predict the

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Psychological Research (2012) 76:542–560 555

outcome of the actions of other individuals. Results showed the associative sequence learning hypothesis on the devel-
that learning ‘by observation’ may foster visual representa- opment of mirror neuron system (Catmur, Walsh, & Heyes,
tions of actions that are used to describe and understand the 2009; Heyes, 2010), which claims that perceptuo-motor
visual dynamics of the actions and of the related contexts representations of actions emerge from a sensorimotor
(e.g., the ball trajectory). In contrast, learning ‘by doing’ learning process. Indeed, planning and execution of actions
may allow for motor, simulative representations of actions is contingently (Cook et al., 2010) and contiguously (Key-
that are used to predict and anticipate the future behaviors sers & Perrett, 2004) associated with the perception of the
of other individuals by reading their body kinematics sensory (kinesthetic and visual) consequences of move-
(Schütz-Bosbach & Prinz, 2007; Urgesi et al., 2010; Wilson ments in the actor’s body and/or of the sensory (visual) sig-
& Knoblich, 2005). nals coming from the body of other individuals performing
The use of these simulative representations in the predic- similar actions in imitative settings. Evidence favoring an
tion of others’ action, however, might be constrained by the associative sequence learning account of mirror neuron sys-
spatial correspondence between executed and observed tem development derives from behavioural (Heyes et al.,
actions. We found that both the improvements of the execu- 2005), TMS (Catmur, Walsh, & Heyes, 2007) and neuroim-
tion training group on body action prediction and the aging (Catmur, Gillmeister, Bird, Liepelt, Brass, & Heyes,
improvements of the observation training group on ball 2008) studies showing a reversal of the typical somatotopic
trajectory understanding were limited to the back-view mirror motor activation after repeated exposure to incon-
perspective. No Wrm conclusion on the role of spatial per- gruent associations of movement execution and action
spective can be drawn from these results, because the observation (e.g., doing a index Wnger movement while see-
absence of any eVects of visual expertise on object motion ing a little Wnger movement). Learning sensorimotor con-
perception in the front-facing perspective might be due tingencies seems not to be constrained by the biological
to fact the last part of the ball trajectory was missing. plausibility of the visual stimulus, since mirror motor facil-
However, a similar account cannot fully explain why long- itation may occur also during observation of a symbolic cue
(experiment 1) and short-term (experiment 2) motor exper- which has been contingently associated with action execu-
tise aVected reading the kinematics of body movements tion (Petroni, Baguear, & Della-Maggiore, 2010). Impor-
only when viewed from the back-view perspective (likely tantly, repeated exposure to contingent associations
linked to Wrst-person experience of the observed actions). between an observed action and an informative symbolic
This might suggest that the compatibility between the cue did not inXuence mirror motor facilitation (Petroni
action representation established during physical practice et al., 2010), thus suggesting that visuo-motor rather than
and the visual percept may inXuence the action prediction visuo-visual associations are required to shape mirror neu-
abilities of athletes. These results are in keeping with stud- ron system activation. The results of the present study are
ies showing higher mirror motor facilitation for actions in keeping with the necessity of experiencing contingent
viewed from a spatial perspective which is compatible sensorimotor associations to forge novel perceptuo-motor
rather than incompatible with the actor’s perspective (Ala- representations of actions, but suggest that these action rep-
erts et al., 2009; Maeda et al., 2002; Urgesi et al., 2006b) resentations allow to predict the future states of others’
and a better perceptual prediction performance for one’s behaviors (Press, Heyes, & Kilner, 2011).
own than others’ actions (Knoblich & Flach, 2001; Knob- While the present data show that direct physical practice
lich et al., 2002). These studies suggest a more accurate and is required to establish novel simulative action representa-
rapid activation of internal, simulative models when tions that can be used to predict others’ actions, they do not
observing actions that match closer the predicted sensorial elucidate which speciWc aspect of physical as compared to
consequences of executed actions. Accordingly, the visual observational practice was critical. Action execution and
appearance of a back-facing, but not of a front-facing, mov- observation diVer for many aspects, including motor plan-
ing body may match directly the internal motor representa- ning and prediction of the sensory consequences of move-
tions of actions established during physical training, thus ments, muscle contractions, kinesthetic perception of the
explaining the superior performance of motor experts as moving body parts and sensorimotor contingencies. Action
compared to visual experts and novices in predicting the recognition is improved by repeated action execution in the
outcome of back-facing, but not front-facing, actions on the absence of any visual feedback (Casile & Giese, 2006;
basis of the initial body movements. Hecht et al., 2001) and even in the absence of any voluntary
The exclusive eVect of physical as compared to observa- action planning as during passive movements (Hecht et al.,
tional practice on the development of predictive action per- 2001). These results would suggest that the contextual per-
ception abilities would suggest that establishing simulative ception of the visual feedback of the movement or active
action representations requires the association between motor planning are not necessary to reinforce or establish
action execution and observation. This is in keeping with new perceptuo-motor representation of actions while

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556 Psychological Research (2012) 76:542–560

aVerent kinaesthetic signals may be suYcient. On the other models of object motion may be particularly important both
hand, while no after-eVects have been found after watching when the body is hidden from view and when it provides
visuo-motor adaptations to perturbed environments (Ong & ambiguous or even deceptive cues. Indeed, the visual repre-
Hodges, 2010), they were still present after direct experi- sentation of object motion (e.g., the ball trajectory) may
ence of visuo-motor adaptation in a deaVerented patient add to performance when the body kinematics is not suY-
(Bernier, Chua, Bard, & Franks, 2006). This would suggest cient for a reliable prediction of the action outcome and
that the eVerent copies of the anticipated sensory conse- when performance is not heavily time constrained. On the
quences of the movements, which might be easily predicted other hand, in sport, as in any competitive social settings,
during active and passive movement execution (Hecht the opponent actors try to disguise or deceive the other
et al., 2001) but are absent during action observation, are players, thus minimizing the information available to the
required to update the internal sensorimotor model of the observer or providing misleading information that makes
action (Ong & Hodges, 2010) that is used to predict the observers error prone (Jackson, Warren, & Abernethy,
future states of others’ behavior (Hommel et al., 2001; 2006). Although motor experts seem to have acquired the
Kilner, Friston, & Frith, 2007; Wilson & Knoblich, 2005). capability to detect the incongruence and the exaggerations
The absence of a signiWcant eVect of the physical train- of the body kinematics that arise from the attempt to pro-
ing on reading the body kinematics of front-facing action vide deceptive information (Jackson et al., 2006; Sebanz &
videos might attenuate the importance of the execution ShiVrar, 2009; Cañal-Bruland, van der Kamp, & van Kest-
training for the perception of the actions performed by eren, 2010), highly competitive situations might impel fast
opponent players, who are typically viewed in volleyball and likely wrong responses to others’ moves based on
from a front-facing perspective. It should be noted, how- deceptive body movements. Indeed, a greater sensitivity of
ever, that the elite athletes, but not the watchers, tested in motor experts to opponents’ body deceptive movements
experiment 1 outperformed novices in body-based predic- has been found among expert French boxers, who had more
tions of the outcome of actions viewed from both back- and false alarms in responding to feints than intermediate and
front-facing perspectives. This is in keeping with previous novice players (Ripoll, Kerlirzin, Stein, & Reine, 1995).
studies showing improved perception abilities of motor Thus, in certain situations, such us a boxer reacting to an
experts as compared to novices for actions viewed from the opponent’s stroke (Ripoll et al., 1995) or a goalkeeper try-
front (e.g., Sebanz & ShiVrar, 2009). The suggestion is ing to intercept a penalty kick (Dessing & Craig, 2010), it is
made that coupling motor and visual experience for periods crucial to rely on the processing of the object motion rather
of time longer than those used in this study may increase than on the possibly disguising or misleading body move-
the Xexibility of internal action models and thus allow to ments of the opponent player. Our data suggest that obser-
incorporate also indirect associations between executed and vational practice is required for the ability to understand the
observed actions. In this vein, the predicted sensory conse- ball trajectory independently from body kinematics.
quences of executed actions may be coupled even with Research on motor skill acquisition showed that physical
kinematics of actions viewed from a third-person perspec- and observational practice may improve action execution at
tive. Furthermore, it is worth noting that our task was qualitatively diVerent levels (Shea et al., 2000). In particu-
purely perceptual, in that it did not require a motor inter- lar, observational practice might allow to develop a repre-
ceptive reaction with the ball shot but only a perceptual sentation of the visuo-spatial coordinates of the action but
judgment. Therefore, it cannot be excluded that diVerent not of its speciWc motor codes, in terms of joint angles and
task settings, involving interceptive reactions in competi- activation patterns (Gruetzmacher et al., 2011), which
tive games, might reveal a greater sensitivity to body cues require direct motor experience. In keeping with a hierar-
while athletes are front-faced with opponent players’ chical model of human action understanding (Grafton &
actions (Farrow & Abernethy, 2003; Streuber, Knoblich, Hamilton, 2007), actions may be represented at diVerent
Sebanz, BülthoV, & de la Rosa, 2011). levels of increasing abstraction, from action kinematics to
That action observation practice alone did not have any the action goals or intentions, which may be underpinned
eVect on the ability to read the body kinematics of the by diVerent sectors of the fronto-parietal action observation
model players does not imply that observational practice is network. Importantly, accessing the speciWc motor codes
ineVective for action perception. Indeed, we showed that based on body kinematics read-out might be important for
the participants of the observation, but not of the execution predicting the future states of others’ actions. However, we
training group in experiment 2 increased their accuracy in cannot exclude that accessing the representation of the
understanding the ball trajectory, suggesting that observing visuo-spatial pattern of the action (acquired via observa-
others interacting with objects is required to build a model tional practice) may help to understand more complex
of the dynamics of the object motion in the environment aspects of others’ actions like for example their goals or
that can be used to anticipate its future course. These visual intentions.

123
Psychological Research (2012) 76:542–560 557

In conclusion, physical and observational practice might Buchanan, J. J., & Wright, D. L. (2011). Generalization of action
provide complementary and mutually reinforcing contribu- knowledge following observational learning. Acta Psychologica,
136(1), 167–178.
tions to the superior perceptual abilities of elite athletes. Calvo-Merino, B., Glaser, D. E., Grèzes, J., Passingham, R. E., & Hag-
This would prompt for the use of diVerent learning strate- gard, P. (2005). Action observation and acquired motor skills: an
gies and of diVerent perceptual cues in motor training pro- FMRI study with expert dancers. Cerebral Cortex, 15(8), 1243–
cedures aimed at optimizing physical and perceptual sport 1249.
Calvo-Merino, B., Grèzes, J., Glaser, D. E., Passingham, R. E., & Hag-
performance. gard, P. (2006). Seeing or doing? InXuence of visual and motor
familiarity in action observation. Current Biology, 16(19), 1905–
Acknowledgments This work was supported by grants from Istituto 1910.
Italiano di Tecnologia SEED 2009 (Prot. n. 21538) and from the Min- Calvo-Merino, B., Urgesi, C., Orgs, G., Aglioti, S. M., & Haggard, P.
istero Istruzione Università e Ricerca (Progetti di Ricerca di Interesse (2010). Extrastriate body area underlies aesthetic evaluation of
Nazionale, PRIN 2009; Prot. n. 2009A8FR3Z) to C.U. and S.M.A and body stimuli. Experimental Brain Research, 204(3), 447–456.
from Istituto di Ricovero e Cura a Carattere ScientiWco ‘‘E. Medea’’ Cañal-Bruland, R., van der Kamp, J., & van Kesteren, J. (2010). An
(Ricerca Corrente 2009, Ministero della Salute) to C.U. We thank Dr. examination of motor and perceptual contributions to the recogni-
Claudia Lopes for her help in data collection. tion of deception from others’ actions. Human Movement Science,
29(1), 94–102.
Candidi, M., Urgesi, C., Ionta, S., & Aglioti, S. M. (2008). Virtual le-
sion of ventral premotor cortex impairs visual perception of bio-
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