Dominant motor control theories propose that the brain uses an internal forward model in combination with a copy of the motor command (efference copy) to predict the sensory consequences of our movements (McNamee and Wolpert, 2019; Shadmehr et al., 2008; Wolpert and Flanagan, 2001). For example, the brain predicts the upcoming touch as one reaches toward an object. These predictions allow for the correction of motor errors without relying on sensory feedback that suffers from intrinsic delays (Shadmehr et al., 2010), thereby improving the estimation of the current state of our body by combining the predicted touch with the actual sensory input (Scott, 2004; Shadmehr et al., 2008; Todorov and Jordan, 2002). These predictions attenuate the perception of the self-generated input (sensory reafference) compared to that of externally generated input (Davidson and Wolpert, 2005; Franklin and Wolpert, 2011; Wolpert and Kawato, 1998) and allow to infer whether the cause of the sensory input is the self or the environment (Brown et al., 2013; Corlett, 2020; Idei et al., 2022). A classic example of this attenuation is that we are unable to tickle ourselves with our own touch, yet we are easily tickled by the touch of others (Blakemore et al., 2000a). The attenuation of sensory reafference – also referred to as sensory cancelation – is thought to be necessary to compensate for the limited capacity of the sensory systems by optimally prioritizing the perception of more informative externally generated stimuli (Bays and Wolpert, 2008; McNamee and Wolpert, 2019). Thus, the attenuation model proposes that we dampen perceptual representations of expected self-generated stimuli to reduce redundancy and to highlight behaviorally relevant unexpected externally generated stimuli.

In contrast, an alternative theoretical framework proposes that predictions, including those arising from our motor commands, should not attenuate but instead enhance sensory signals, thereby allowing for sharper (i.e. more accurate) representations of predicted compared to unpredicted sensory events (Press et al., 2020; Press and Yon, 2019; Yon et al., 2021). This enhancement account – also referred to as the sharpening account – posits that predictions based on our motor commands are equivalent to expectations formed by statistical regularities in sensory input or from prior knowledge (e.g. at the North Pole, one expects to see a polar bear rather than an elephant) and that these predictions should bias our perception towards our expectations. The proposal mainly stems from experimental research outside the domain of action and argues that weak, noisy, or ambiguous sensory input that is in line with prior expectations should be enhanced to achieve, on average, more accurate representations. For example, we are more biased to report the presence of visual events that are statistically likely to occur rather than unlikely events (Chalk et al., 2010; Wyart et al., 2012), more sensitive to low-level visual features that are in line with prior expectations (Stein and Peelen, 2015; Teufel et al., 2018), and show greater biases when perceiving visual events that are congruent with our expectations (Hudson et al., 2016a; Hudson et al., 2016b). Such effects are thought to result from mechanisms that increase the ‘gain’ of expected information by altering the weights of different sensory signals (Summerfield and de Lange, 2014). Thus, the enhancement model proposes that we amplify the perceptual representations of expected compared to unexpected sensory input.

In the somatosensory domain, evidence supporting the attenuation model has repeatedly shown that touch delivered to one (passive) hand by the other (active) hand (i.e. self-generated touch) is perceived as weaker or less ticklish (Asimakidou et al., 2022; Bays et al., 2006; Bays et al., 2005; Blakemore et al., 1999; Kilteni et al., 2021; Kilteni et al., 2020; Kilteni et al., 2019; Kilteni et al., 2018; Kilteni and Ehrsson, 2022; Kilteni and Ehrsson, 2017a; Kilteni and Ehrsson, 2017b; Knoetsch and Zimmermann, 2021; McNaughton et al., 2023; Weiskrantz et al., 1971; Wolpe et al., 2016) and evokes attenuated somatosensory cortical activation (Blakemore et al., 1998a; Hesse et al., 2010; Kilteni and Ehrsson, 2020; Shergill et al., 2014; Shergill et al., 2013) compared to a touch of identical intensity applied on the passive hand that is externally generated. In contrast, evidence supporting the enhancement model has shown that the action of the active hand results in an increase in the perceived intensity of touch on the passive hand, provided that the active hand never receives touch simultaneously with the passive hand (i.e. hands do not make contact) (Thomas et al., 2022). This enhancement finding has been recently used to support the sharpening model and to argue that attenuation effects are due to unspecific gating processes caused by the simultaneous double tactile stimulation of the two hands rather than action prediction (Press et al., 2023).

The attenuation (or cancellation) and enhancement (or sharpening) models present strikingly different hypotheses regarding how action influences the perception of sensory input and are supported by contradictory experimental evidence, leading to debates between researchers (Corlett, 2020; Fuehrer et al., 2022; Kilteni and Ehrsson, 2022; Press et al., 2023; Press et al., 2020; Thomas et al., 2022). Clarifying how predictions about the sensory consequences of our movements affect our perception is fundamental to understanding the interaction between perception and action (McNamee and Wolpert, 2019; Shadmehr et al., 2008; Wolpert and Flanagan, 2001) but also for clinical and neurobiological theories of psychosis spectrum disorders, such as schizophrenia (Blakemore et al., 2000b; Corlett et al., 2019; Frith, 2005a; Frith, 2005b; Frith, 2019; Frith et al., 2000; Shergill et al., 2014; Shergill et al., 2005) and schizotypy (Asimakidou et al., 2022), as well as functional movement disorders (Pareés et al., 2014), Parkinson’s disease (Wolpe et al., 2018), and aging (Wolpe et al., 2016).

The present study aimed to contribute to this debate by directly contrasting the enhancement and attenuation models within the same paradigm. To this end, the same force discrimination task employed in earlier studies reporting attenuation (Asimakidou et al., 2022; Bays et al., 2006; Bays et al., 2005; Kilteni et al., 2023; Kilteni et al., 2021; Kilteni et al., 2020; Kilteni et al., 2019; Kilteni and Ehrsson, 2022; Timar et al., 2023) and enhancement (Thomas et al., 2022) was used to determine (a) whether movement of the right hand enhances the perceived magnitude of touch applied on the left hand when the two hands do not make contact (Experiments 1 and 2) and (b) whether attenuation effects are due to double tactile stimulation caused by the contact of the two hands or action prediction (Experiment 3). Capitalizing on the fact that any conclusion about whether action prediction ‘attenuates’ or ‘enhances’ the perception of the somatosensory input needs to be made with a comparison to one’s somatosensory perception in the absence of action, we also included a condition in which participants passively received externally generated touch, with which we compared the participants’ perception in all experimental conditions. Consequently, if participants perceive a touch as less or more intense during action than in the absence of action, we can infer that the received touch was attenuated or enhanced, respectively. This is a critical methodological novelty compared to previous studies (Bays et al., 2006; Thomas et al., 2022) because if such baseline conditions are missing, the same patterns of results can be incorrectly attributed to ‘attenuation’ or ‘enhancement’: for example, if one condition produces less attenuation than another, it may be interpreted as enhancement, and vice versa. Indeed, it was this baseline condition that turned out to be critical for distinguishing between attenuation and enhancement effects. All studies and analysis plans were pre-registered on the Open Science Framework prior to data collection (Methods).

In the present study, two opposing hypotheses regarding how action influences somatosensory perception were contrasted: the attenuation model and the enhancement model. Our findings demonstrate that action does not enhance (Experiments 1 and 2) but attenuates the predicted touch (Experiment 3) compared to identical touch in the absence of action. Before discussing the findings, it is important to emphasize that to draw conclusions about whether perception is attenuated or enhanced in an experimental condition including action, it is necessary to include a baseline condition without action. Only comparing conditions that include action prevents differentiating a genuine effect of enhancement (or attenuation) from an effect of reduced attenuation (or enhancement) in one of the two conditions. This also applies to experimental manipulations that contrast predicted with unpredicted somatosensory stimuli during action (see Thomas et al., 2022 for such comparisons). To this end, a baseline condition of pure somatosensory exafference (i.e. externally generated touch) was included in all the present experiments that allowed us to distinguish the direction of the effects.

The results of Experiment 1 showed a robust attenuation of the touch applied to the passive left hand when the two hands made contact (contact condition), but neither attenuation nor enhancement was caused by the mere movement of the right hand (no-contact condition). In contrast, the perceived intensity of touch in the no-contact condition was similar to that of the baseline (i.e. externally generated touch). These findings are in line with the results of Bays et al., 2006, who found no change in the participants’ somatosensory perception when the two hands did not make contact, but do not replicate those of Thomas et al., 2022, who observed enhancement of touch in a no-contact condition. Experiment 2 further investigated the source of the previously reported enhancement effects and showed that they are in fact driven by the reference condition used: enhancement was observed only relative to a condition in which the touch was applied rapidly following a cue to inhibit the movement (i.e. a ‘do not move’ cue), as in previous research (Thomas et al., 2022), but not relative to an externally generated touch condition (our baseline). In support of this claim, Experiment 2 showed that a cue to inhibit the movement results in a significant reduction in the perceived intensity of the imperative stimulus (i.e. touch on the passive hand) compared to baseline perception. This is in line with previous evidence showing reduced amplitudes of somatosensory evoked potentials for tactile stimuli presented shortly following a cue to inhibit a movement (Hoshiyama and Sheean, 1998), reduced perceived amplitude for tactile stimuli under the mere expectation to move (Voss et al., 2008; Voss et al., 2006), and reduced encoding of sensory stimuli following motor inhibition (Chiu and Egner, 2015b; Yebra et al., 2019). Therefore, rather than participants’ perception being enhanced in the no-contact condition, it is a reduction of the perceived intensity following the cue to inhibit the movement that leads to an apparent enhancement. In contrast, by including a novel externally generated touch condition as a baseline that involved neither motor planning nor response inhibition, it became clear that there were no enhancement effects.

Experiment 3 demonstrated that action attenuates the predicted touch even if the active hand does not receive simultaneous tactile stimulation with the passive hand. When participants simply moved their right hand to trigger the touch on their left hand (no-contact trials in Experiments 1 and 2), no change in their somatosensory perception was found. This suggests that an arbitrary mapping between the movement of the right hand and the delivery of touch on the left hand is insufficient to elicit attenuation. In contrast, when participants expected that they would touch their own hand (Experiment 3), significant attenuation was observed, even when the active hand unexpectedly missed the contact (no-contact trials, 20%). This pattern of results can be explained by the attenuation model, which suggests that the self-generation of a stimulus is in itself not sufficient for predictive attenuation, but that the action and its predicted sensory feedback are consistent with touching one’s own body (Bays and Wolpert, 2008). Thus, a sensorimotor context that closely resembles tapping directly on the left index finger with the right index finger (self-touch) was critical for predictive attenuation (Bays et al., 2006). The findings of Experiment 3 also contradict the suggestion that attenuation on a passive hand reflects a nonpredictive gating of tactile input during movement of the active hand (Press et al., 2023; Thomas et al., 2022). Indeed, several earlier studies showed gating effects only on the moving limb (movement effector) and not on the passive limb (Chapman et al., 1987; Cohen and Starr, 1987; Colino et al., 2014; Papakostopoulos et al., 1975; Pertovaara et al., 1992; Rushton et al., 1981), and we have also recently shown that experimental paradigms identical to the one used in the present study produce attenuation effects but not gating effects on the passive hand (Kilteni and Ehrsson, 2022). Overall, the results suggest that attenuation effects are driven by action prediction and not the simultaneous tactile stimulation of the two hands. From an ecological point of view, in every self-touch behavior, we necessarily receive somatosensory input on the active hand and the body part that passively receives touch (‘touchant-touché’ [Schütz-Bosbach et al., 2009]), and it is within these sensorimotor contexts that the brain forms predictions about the somatosensory consequences on multiple body parts (Bays and Wolpert, 2008).

Interestingly, the magnitude of attenuation was greater in the contact compared to the no-contact condition by approximately 35% in Experiment 3. This could suggest that the attenuation effect observed in the contact condition arises from multiple sources, one being the motor-based prediction and another source being the simultaneous touch on the active hand. However, this interpretation is inconsistent with the findings that somatosensory attenuation has been previously observed in the absence of simultaneous tactile stimulation, for example, when imagining but not executing the right hand movement (Kilteni et al., 2018) or just before the hands make contact (Bays et al., 2005). Similarly, no somatosensory attenuation is observed in the absence of action prediction, even if the two hands receive simultaneous tactile stimulation; for example, the passive displacement of the right hand towards the left hand that is accompanied by a double touch (Kilteni et al., 2020) or the mere delivery of simultaneous tactile stimulation on both hands (Bays et al., 2005) does not produce attenuation. Instead, our finding that the magnitude of attenuation was greater in the contact than the no-contact condition in Experiment 3 may instead reflect a decrease of attenuation in the no-contact trials, due to the unexpected omission of contact influencing the perceived magnitude of touch in a postdictive manner. Specifically, the unexpected omission of contact in the minority of trials (20%) could be seen as a form of stimulus omission akin to so-called ‘silent oddballs’ that are known to generate prediction errors (Busse and Woldorff, 2003; Karamürsel and Bullock, 2000; SanMiguel et al., 2013a; SanMiguel et al., 2013b). That is, although participants clearly attenuated the touch predicted by their movement in the no-contact trials, their expectation of contact was necessarily violated. In one of our previous studies (Kilteni et al., 2019) using the same force discrimination task as in the present study we showed in three experiments that participants experienced the touch to be stronger (i.e. less attenuation) and more ticklish when triggering a prediction error (<17% frequency). Similarly, research outside the domain of action (Kusnir et al., 2020) has found that perception is improved for tactile events delivered at unpredictable compared to predictable temporal onsets, also suggesting that tactile perception is facilitated in the presence of prediction error. In line with these previous results, the decreased attenuation we observed in the no-contact condition could be attributed to the prediction error. Additionally, this violation could be considered a novel event given its infrequency. Novel events can have several consequences for cognition, including transient enhancements of perception (Schomaker and Meeter, 2012), facilitated encoding of information into working memory (Mayer et al., 2011) as well as changes in the allocation of attentional resources in a postdictive manner (for a review of the effects of novelty on cognition see Schomaker and Meeter, 2015).

Some authors have criticized the comparison of perceived intensity of sensory stimuli in self-generated and externally generated conditions (Press et al., 2020; Yon et al., 2018; Yon et al., 2021). A frequent criticism is that tactile input may be ‘predicted’ in self-generated conditions through action and ‘unpredicted’ in externally generated (passive) conditions that do not involve action. This concern can be ruled out since in all three experiments, the stimulus in the baseline (i.e. externally generated touch) condition was delivered at a fixed timepoint (800 ms after the cue); therefore, participants could predict it in the absence of motor-based predictions. Second, it has been argued that somatosensory attenuation findings may be driven by dual-task requirements (Press et al., 2020) present only in self-generated conditions that could increase the working memory load or result in a shift of attention towards the active hand. A similar criticism has been suggested in which the mere movement of the active hand in self-generated conditions can result in a nonpredictive gating of tactile input. However, these explanations can also be ruled out, given that dual-task requirements and movement of the active hand were present in the contact and no-contact conditions of all three experiments without concomitant attenuation effects. Indeed, if dual-task requirements, or the mere presence of movement during the perceptual task, were sufficient to cause attenuation effects, then attenuation would have been observed in the no-contact conditions of Experiments 1 and 2. Furthermore, if the mere simultaneous tactile stimulation on the active hand was responsible for the attenuation effect, we would not have observed attenuation in the no-contact condition of Experiment 3. Finally, alternative explanations based on differences in other psychophysical parameters, movement kinematics, or timings between experiments can also be ruled out.

How can the findings that action prediction attenuates touch be reconciled with those showing that expectations outside the domain of action improve sensory perception (Press et al., 2020)? While it is difficult to directly compare these lines of research because of differences in the sensory modality investigated, the task designs used, and the perceptual measures employed, we speculate that there are numerous possible reasons why action prediction may not have the same effect on perception as prediction mechanisms outside the domain of action. First, research on action-based predictions concerns ubiquitous associations between actions and their sensory consequences that we are continually exposed to throughout the lifespan. For example, we are constantly exposed to associations between our motor behaviors and their tactile consequences during self-touch, even as early as 13 weeks in utero (Kurjak et al., 2003). In contrast, research on sensory expectations outside the domain of action primarily concerns arbitrary associations between stimuli that are typically learned only during the time course of a given task. It is, therefore, conceivable that separable mechanisms may operate to predict action effects versus stimulus-stimulus associations (see Dogge et al., 2019a; Dogge et al., 2019b for discussion). Second, higher-level expectations, such as explicit prior knowledge that a specific sensory event is likely, might not operate in the same way as lower-level action predictions; for example, it has been proposed that action-based predictions inhibit expected stimuli, while sensory expectations potentiate the expected sensory inputs (de Lange et al., 2018). Most importantly, from a theoretical perspective, attenuating the predicted sensory consequences of actions does not necessarily mean that the brain forms inaccurate representations of the world but instead indicates a flexible strategy that prioritizes more informative externally generated events.

It is important to emphasize that this study investigated the perception of self-generated touch during an action directed toward one’s own body. Thus, perception was assessed on a passive body part that was the target of a movement of an active body part (i.e. moving the right hand to touch the static left hand). This focus was chosen because extensive previous research has shown deteriorated performance when detecting or discriminating touch (both self-generated and externally generated) delivered on a moving limb, a phenomenon referred to as movement-related gating (Chapman et al., 1996; Chapman et al., 1987; Voudouris and Fiehler, 2021; Voudouris and Fiehler, 2017; Williams et al., 1998). By assessing somatosensory perception on a static limb, we were able to isolate the effect of action prediction on somatosensory perception from other perceptual phenomena related to the movement per se including movement-related gating (Chapman et al., 1987; Cohen and Starr, 1987; Colino et al., 2014; Kilteni and Ehrsson, 2022; Papakostopoulos et al., 1975; Pertovaara et al., 1992; Rushton et al., 1981). However, it is interesting to note that during the active exploration of objects through touch, referred to as active touch, self-generated touch also occurs, albeit only on the moving limb. Studies on active touch have observed improved perception of object features, such as texture or edge orientation when actively exploring an object (Morley et al., 1983; Pruszynski et al., 2018) compared to passive exposure (Bensmaia et al., 2008; Peters et al., 2015). At first, the improved perception observed during active touch might appear paradoxical to observations of movement-related gating and predictive attenuation effects: if anything, active touch should result in worse perception than passive touch. However, it has been shown that humans optimize their exploratory movements to minimize these effects and maximize the quality of their sensory feedback. For example, it is known that the degree of gating decreases as movement speed slows (Angel and Malenka, 1982; Chapman et al., 1996; Schmidt et al., 1990) and it has been proposed that participants optimize the speed of movement during active touch to avoid high speeds associated with tactile gating (Cybulska-Klosowicz et al., 2011). Similarly, when exploring objects through active touch, weaker forces are typically applied (e.g. <1 N [Olczak et al., 2018; Tanaka et al., 2014]) that elicit less attenuation than stronger forces, as shown in the classic force-matching task (Shergill et al., 2003). Thus, differences in the task context might primarily influence how one moves (e.g. the act of swatting a fly from one’s arm versus feeling for a bump on one’s skin), which can have disparate ensuing effects on perception depending on the movement kinematics.

Debates between researchers supporting attenuation or enhancement are useful and fruitful for scientific dialogue and advancement. The present study revisited recent findings on somatosensory enhancement during action and showed that when the requirement to inhibit the action is removed from the baseline, action predictions do not enhance but attenuate the received somatosensory input. Our results are in strong alignment with animal studies showing that action attenuates the predicted sensory consequences (for reviews, see [Brooks and Cullen, 2019; Crapse and Sommer, 2008; Cullen, 2004; Schneider and Mooney, 2018; Straka et al., 2018]). For example, crickets and mice suppress auditory reafferent signals but maintain their sensitivity to external sounds (Audette et al., 2022; Poulet and Hedwig, 2003; Poulet and Hedwig, 2006; Schneider et al., 2018), the weakly electric fish attenuates its electrosensory reafference to respond to externally generated electrical signals (Cullen, 2004; Sawtell, 2017), and primates attenuate their vestibular reafference and activate vestibular-related reflexes only when the vestibular input is exafferent (Brooks et al., 2015; Cullen, 2012; Roy and Cullen, 2004). To this end, the results of the present study prompt a reappraisal of recent experimental findings upon which theoretical frameworks proposing a perceptual enhancement by action prediction are based.

Data (PSEs and JNDs) are publicly available along with pre-registered analyses plans on the OSF (Experiment 1: https://osf.io/9jkqt, Experiment 2: https://osf.io/hs8au, Experiment 3: https://osf.io/pxy87).

1. 1. Job X
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   ID 9jkqt. Attenuation versus enhancement of self-generated touch.

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Author details

1. Xavier Job

   Department of Neuroscience, Karolinska Institute, Stockholm, Sweden

   Contribution

   Conceptualization, Formal analysis, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6017-5983

2. Konstantina Kilteni

   1. Department of Neuroscience, Karolinska Institute, Stockholm, Sweden
   2. Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen, Netherlands

   Contribution

   Conceptualization, Writing – original draft, Writing – review and editing

   For correspondence

   konstantina.kilteni@ki.se

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-6887-6434

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