Our brains capacity to adapt, known as cortical plasticity, is integral to our successful functioning in daily life, as well as rehabilitation from injury. A key model for exploring the extent, and consequences of, cortical plasticity is upper-limb loss (via amputation or congenital absence). Here, the cortical hand territory in the primary somatosensory cortex (hereafter S1), suffers an extreme loss of sensory input in tandem with dramatic alterations of motor behaviour (Makin et al., 2013a; Muret and Makin, 2021). The functional and perceptual correlates of amputation-related plasticity are currently debated (Makin and Bensmaia, 2017; Ortiz-Catalan, 2018). In particular, it is not clear whether functional cortical reorganisation is restricted to early life development or can also occur in adults.

Traditionally, research assessing cortical plasticity after upper-limb loss has followed the tenet that neighbouring body parts of the missing hand, and the lower face in particular, shift and encroach into the deprived hand area. This emphasis on the lip representation stems from early electrophysiological work in non-human primates, where numerous studies demonstrated an ‘upside-down’ facial somatotopy, with the lower face immediately neighbouring the hand (Dreyer et al., 1975; Merzenich et al., 1978; Sur et al., 1982; Cusick et al., 1986; Lin and Sessle, 1994; Manger et al., 1995; Manger et al., 1996; Jain et al., 2001; Cerkevich et al., 2014). Here, the lips and/or chin inputs have been shown to remap into the deprived hand area after sensory loss (Pons et al., 1991; Jain et al., 1997), leading to the well-accepted assumption that remapping is determined by cortical proximity (Buonomano and Merzenich, 1998; Nardone et al., 2013). Thereafter, human measurement of topographic shifts has tended to focus on that of the lips, where researchers have reported that shifted lip representation towards and into the deprived hand area is significantly associated with phantom limb pain (PLP) intensity (Flor et al., 1995; Birbaumer et al., 1997; Lotze et al., 2001; Grüsser et al., 2001; Foell et al., 2014). PLP is a neuropathic pain syndrome experienced in the missing, amputated limb by the majority of amputees (Limakatso et al., 2019). This condition is commonly thought to arise from maladaptive cortical plasticity in S1 (although see Makin, 2021), specifically from a signal mismatch between the missing hand representation and the remapped inputs of the lips in the deprived hand area (Ramachandran and Hirstein, 1998).

The research focus on lip cortical remapping in amputees is based on the assumption that the lips neighbour the hand representation. However, this assumption goes against the classical upright orientation of the face in S1 (Penfield, 1950; Schwartz et al., 2004; Roux et al., 2018; Sato et al., 2005; Willoughby et al., 2020), as first depicted in Penfield’s Homunculus and in later intracortical recordings and stimulation studies (Penfield, 1950; Schwartz et al., 2004; Roux et al., 2018; Sato et al., 2005), with the upper-face (i.e. forehead) bordering the hand area. Furthermore, neuroimaging studies in humans studying face topography provided contradictory evidence for the past 30 years. While a few neuroimaging studies provided partial evidence in support of the traditional upright face organisation (Willoughby et al., 2020), other studies supported the inverted (or ‘upside-down’) somatotopic organisation of the face, similar to that of non-human primates (Yang et al., 1993; Servos et al., 1999). Other studies suggested a segmental organisation (Moulton et al., 2009), or even a lack of somatotopic organisation (Iannetti et al., 2003; Nguyen et al., 2004; Kopietz et al., 2009), whereas many studies provided inconclusive or incomplete results (Mogilner et al., 1994; Hoshiyama et al., 1996; Disbrow et al., 2003; Nevalainen et al., 2006). Together, the available evidence does not successfully converge on face topography in humans. In line with the upright organisation originally suggested by Penfield, recent work reported that the shift in the lip representation towards the missing hand in amputees was minimal (Makin et al., 2015; Raffin et al., 2016), and likely to reside within the face area itself. Surprisingly, there is currently no research that considers the representation of other facial parts, in particular the upper-face (e.g. the forehead), in relation to plasticity or PLP. Detailed mapping of the upper and lower face is therefore needed to assess typical topography of facial sensorimotor organisation, as well as remapping after limb loss.

More recent electrophysiological studies in monkeys demonstrated that much of the face remapping observed in the primary sensorimotor cortex following upper-limb deafferentation does not result from cortico-cortical plasticity, but instead arises from plasticity in the brainstem (Florence and Kaas, 1995; Kambi et al., 2014). This important finding also highlights the limited explanatory power of local activity increase that has been historically used to infer changes to the representational features of a given brain. Does it reflect a gain modulation of input coming from the brainstem, or increase of local processing of the face input in the hand area? While it remains challenging to dissociate these two contributions, alternative analysis tools are becoming increasingly popular for mining richer information of the processing underlying activity in a given cortical region. Multivariate analyses are sensitive to more subtle changes in representational content, that are not accessible with the traditional univariate approach. In the context of facial remapping, if the deprived hand area updates its local processing to include facial information (and does not just display more facial activity), we would expect it to show greater information about representational features relevant to different facial parts.

Remapping after upper-limb loss has also been documented in individuals born without a hand (hereafter one-handers), who do not experience PLP (Makin et al., 2013b). Here, it has been shown that the representation of multiple body parts, including the residual arm, legs and lips, remapped into the missing hand territory (Hahamy et al., 2017; Hahamy and Makin, 2019). Importantly, cortical remapping in this group does not depend on cortical proximity. With regard to the lips, a recent transcranial magnetic stimulation (TMS) study has reported functionally relevant lip activity in the deprived hand area of one-handers (Amoruso et al., 2021), showcasing that reported remapping may also be functional. It was proposed that the observed remapping of various body parts could have been shaped by compensatory behaviour (Hahamy et al., 2017), as these body parts are all used by one-handers to compensate for their missing hand function, but this hypothesis awaits validation.

Here, we conducted a mapping of face cortical organisation to determine facial orientation (upright versus inverted) in the primary sensorimotor cortex in 22 two-handed controls, 17 amputees, and 21 one-handers using an active functional MRI paradigm, where participants were visually instructed to move their forehead, nose, lips or tongue. This paradigm was chosen because it enabled simultaneous bilateral activation of S1 within individual participants, providing a within-participants control design. We also explored the representation of multiple facial movements, which have not been previously studied in the context of deprivation-triggered brain plasticity. To measure the extent of cortical remapping of the upper (forehead) and lower (lips) face in relation to the deprived (or non-dominant) hand area across all groups, we used surface-based topographic comparisons. For this purpose, we employed up-to-date methodology to harvest traditional measures, that is winner-takes-all assessment of surface coverage in the hand and face areas, followed by cortical (geodesic) distances, and similarity analysis of selectivity maps. Furthermore, to go beyond the gross topographic properties of face representation, we used multivariate representational similarity analysis (RSA). This approach allows us to characterise more subtle alterations in the relationship between facial activity patterns (forehead, nose, lips, tongue) in the deprived hand and the face areas.

We found that, in line with Penfield’s original description, facial topography was arranged in an upright manner, with the forehead (i.e. upper-face) bordering the hand area across all groups. Contrary to traditional theories (Flor et al., 2006), we did not find evidence for facial remapping of either the lips or the (cortically neighbouring) forehead, into the deprived hand area of amputees. We did, however, observe significant remapping of multiple face parts (upper and lower face) in the one-handers group, validating our methodology as suitable for identifying remapping effects. Interestingly, remapping of the cortical neighbour (upper-face) within the one-handers group was away from the missing hand area, while the lips and tongue representations shifted towards the deprived cortex, hinting that the underlying mechanism of remapping is more complex than simple cortical proximity.

It is a well-accepted notion, rooted in non-human primate electrophysiological data, that upper-limb amputation triggers cortical remapping of the assumed neighbour – the lower face – into the missing hand area. This previous work predominantly characterised remapping by investigating shifts of the lip representation (Flor et al., 1995; Birbaumer et al., 1997; Lotze et al., 2001; Grüsser et al., 2001; Foell et al., 2014; Karl et al., 2001; MacIver et al., 2008). However, by focusing on only one face part, activity elicited by other facial parts (such as the forehead) are not taken into account (see Muret and Makin, 2021). Here, we explored the relationship of face-to-hand remapping in controls and one-handed groups, and used both univariate (topographic) and multivariate (representational structure) methods to investigate in detail the information content of the face in both the deprived and intact hand and face areas. We found evidence for an upright somatotopy of the face across all groups, confirming that the cortical neighbour to the hand in humans is the upper, not lower, face. We further found little evidence for remapping of all tested facial parts in amputees, with no significant relationship to the presence of PLP. As a positive control, we also recruited individuals that were born without a hand (one-handers), who have previously shown cortical remapping across multiple body parts (Hahamy et al., 2017; Hahamy and Makin, 2019; Amoruso et al., 2021). Across multiple facial parts (forehead, lips and tongue), one-handers showed evidence for a complex pattern of face remapping in the deprived hand area, with consistent and converging evidence across analysis approaches. Finally, we demonstrate that facial representation in amputees’ deprived hemisphere is more similar to two-handed controls than to one-handers. Together, our findings demonstrate that the face representation in humans is highly plastic, but that this plasticity is restricted by the developmental stage of input deprivation, rather than cortical proximity.

Firstly, our univariate analyses at both group and individual levels provides converging and clear evidence for an upright orientation of the face in controls, amputees and one-handers. Contrary to previous neuroimaging studies reporting an inverted facial somatotopy (Servos et al., 1999; Yang et al., 1993; similar to primates Manger et al., 1996; Jain et al., 2001), or a lack of somatotopic organisation (Iannetti et al., 2003; Nguyen et al., 2004; Kopietz et al., 2009), here we found that the forehead representation borders the hand representation, followed by the nose, the lips, and the tongue located most laterally. These discrepancies may arise from the challenge to find a robust and reliable method to stimulate face parts, and thus elicit detectable cortical activation. In this context, it may be argued that it is difficult to achieve isolated execution of specific facial muscles when performing gross movements without impacting sensory processing of neighbouring facial parts. For instance, tongue movements in our paradigm (e.g. touching the roof of the mouth with the tongue), may be best considered as a holistic inner mouth movement, and forehead movements may be best considered as engaging the upper-face. While this critique is valid, it may also be relevant (although to a smaller degree) for passive paradigms, as stimulation can induce waves that propagate through the skin (Manfredi et al., 2012; Sofia and Jones, 2013; Shao et al., 2016) and Pacinian receptors were found to activate during stimulation of remote sites (Edin and Abbs, 1991; Prsa et al., 2019). Despite this caveat, both our univariate and multivariate analyses showed that we were successful in isolating sensorimotor representations of the various movements (forehead, nose, lips, and tongue) within our regions of interest (see Appendix 1—figure 2 for validation of our approach using vibrotactile stimulation). In other words, even if somatosensory information is overlapping across movements, there is still enough distinct information to separate representational patterns. This finding indicates the suitability of our motor paradigm for teasing apart facial somatotopy, allowing us to characterise the face in greater detail than previously attempted. According to the confirmed upright organisation, if cortical remapping of neighbours exists, we would expect to see the forehead shifting towards and into the hand area – not the lips.

When looking at face-to-hand remapping in amputees, where the remapping of cortical neighbours has been the prevalent explanation for PLP, we find little evidence of shifts of locality and remapping towards the deprived hand area for facial parts, including the neighbour (forehead) and hypothesised neighbour (lips). This was further confirmed by the Jaccard analysis showing that amputees’ maps were more similar to the ones of controls than of one-handers in the deprived hemisphere, as well as similar spatial representation of amputees’ phantom thumb movements relative to controls (Appendix 1—figure 3). Our univariate results are further partially supported by our multivariate analysis, where we find no significant changes in dissimilarity of activity patterns across facial parts in amputees’ deprived hand area relative to controls. These results support previous work reporting minimal cortical remapping after amputation (Makin et al., 2013b; Kikkert et al., 2018; Raffin et al., 2012), suggesting that in amputees this area might be functionally unipotent – pertaining to hand-related activity alone and lacking the ability to reorganise after hand loss. However, due to some inconclusive Bayes Factor in our key analyses, we cannot strongly conclude that remapping does not occur in this group. This could be attributed to our relatively small sample (further recruitment was prevented due to Covid-19 restrictions), and in particular, the small proportion of amputees experiencing PLP (11 out of 17). However, pain is not a necessary condition for deprivation-triggered remapping (Recanzone et al., 1992; Wang et al., 1995; Andoh et al., 2020) and vice versa, PLP has been shown to be experienced in absence of remapping (De Nunzio et al., 2018). Moreover, previous studies reporting significant difference between amputees who experienced PLP and those who do not, often employed similar (or smaller) sample sizes (Lotze et al., 2001), indicating that the expected effect of remapping should be substantial.

Our findings seem contradictory to the many previous studies reporting lip remapping in amputees (Flor et al., 1995; Birbaumer et al., 1997; Lotze et al., 2001; Grüsser et al., 2001; Foell et al., 2014; Karl et al., 2001; MacIver et al., 2008). A major difference with regard to these previous studies, which predominantly focused on a single part of the face, lies in the fact that our study was the first to assess the mapping and potential remapping of multiple facial parts at once. By focusing on the lips only, previous designs excluded other facial parts which may have elicited greater activity in certain areas of S1, resulting in a less accurate delineation of the lip-selective representation. Such down-sampling of body maps, therefore, can lead to biased results and interpretation (Muret and Makin, 2021). While our design is not exempt from this limitation, the fact that we assessed other parts of the face may explain why our results diverged from previous findings.

We did find anecdotal evidence for remapping for the tongue within the deprived hand area in amputees. This was a surprising result, as the tongue is not a cortical neighbour to the hand and was not specifically hypothesised to remap in amputees. We also found that amputees demonstrated a different amount of facial information across the two hand areas. Although this multivariate result was not significantly different to that of controls, it demonstrates the plausibility that an inter-hemisphere imbalance may exist to a certain degree, albeit any relationship to PLP is tenuous. While these latter results require further validation, they support our premise that cortical proximity of representations may not be a necessity for remapping to occur. In this context, as our tongue condition could also be classed as an ‘inner mouth’ movement, it is important to note that previous work addressing sensorimotor representations of the mouth and the larynx have demonstrated both lateral and more medial ‘hotspots’ (i.e. a ‘double’ representation Eichert et al., 2020). The potential tongue remapping in amputees, therefore, may reflect changes in the medial mouth representation, but this would need to be investigated further. Moreover, even if the remapping we observed here goes against the theory of cortical proximity, it can still arise from representational proximity at the subcortical level, in particular at the brainstem level (Florence and Kaas, 1995 ; Kambi et al., 2014). While challenging in humans, mapping both the cuneate and trigeminal nuclei would be critical to provide a more complete picture regarding the role of proximity in remapping.

We did find converging and conclusive evidence for cortical remapping of multiple facial parts, both neighbours (forehead) and non-neighbours (lips and tongue) in our congenital one-handed group. Here, the pattern of remapping is strikingly different to that of cortical neighbourhood theories. Specifically, the location of the cortical neighbour – the forehead – is shown to shift away from the deprived hand area, which is subsequently more activated by the lips and the tongue than is the intact hand area. The increase of facial activity in the deprived hand area is in turn supported by our multivariate results, whereby significantly greater information content for the face was found in the deprived hand area for one-handers when compared to controls. One-handers’ deprived hand area, therefore, seems to have increased discriminability between different facial movements. It is difficult to ascertain from our study the drivers of this remapping. It has been suggested previously that remapping within this group may be driven by functionally-relevant behaviour substituting for the loss of the limb (Hahamy et al., 2017; Amoruso et al., 2021). Together with the recent evidence that lip information content is already significant in the hand area of two-handed participants (Muret et al., 2022), compensatory behaviour since developmental stages might further uncover (and even potentiate) this underlying latent activity. Alternative explanations relate to an overall and unspecific release of inhibition (i.e. decreased GABA) in the missing hand area, allowing for latent activity of other body parts to be detected (Hahamy et al., 2017). While speculative, our results tend to support the former, as we report remapping for facial parts which have the ability to compensate for hand function, for example using the lips and/or mouth to manipulate an object, and a lack of remapping into the hand area for those that cannot (the forehead and nose). This increased activity from body parts compensating for hand function may represent a stabilising mechanism, aimed at preserving the integrity of the sensorimotor network and its function (Muret and Makin, 2021). The deprived hand area in one-handers, therefore, may reflect domain specificity – suitable for adapting to multiple body parts (Amoruso et al., 2021), which may preserve the role of the hand area by sustaining its hand-function related information content.

A limitation that should be acknowledged arises from the potential contribution to S1 from M1 activity. Since these cortical areas are neighbours, it is difficult to separate them with certainty. We minimised the contribution of M1 by taking multiple acquisition and pre-processing steps, including the use of anatomical delineation at the individual level, as well as a comprehensive analytical approach (e.g. both univariate and multivariate techniques). Perhaps most convincingly, our RSA evidence for remapping in congenital one-handers but not in amputees were qualitatively stronger in S1 relative to M1. Furthermore, it has been claimed that active movements may produce different cortical maps to those with passive stimulation (Flor et al., 2013; Andoh et al., 2018), and previous work demonstrating a relationship between cortical remapping and PLP tended to use passive stimulation (Flor et al., 1995; Birbaumer et al., 1997; Foell et al., 2014; Karl et al., 2001). However, we do not think this methodological difference underlies our contrasting results as movement-induced lip activity has been shown to demonstrate lip remapping before (Lotze et al., 2001; MacIver et al., 2008; Striem-Amit et al., 2018), indicating that an active paradigm is suitable for assessing cortical remapping (if it exists). Conversely, a recent study using passive lip stimulation in amputees did not find any evidence for remapping (Philip et al., 2017). Moreover, we recently ran a study which found that S1 topography and multivariate representational structure are similar across active and passive paradigms (Sanders et al., 2019) (see Appendix 1—figure 2 for a comparison between active and passive facial stimulation). In our view, the choice of an active paradigm is the most reflective of naturalistic tactile inputs in everyday life. Together with robust evidence for remapping in one-handers using all the methods tested here, our choice of active paradigm is clearly suitable to identify topographic organisation and remapping, and is practically accessible and translatable to fMRI designs.

To conclude, both our univariate and multivariate analyses found consistent evidence for a complex pattern of face remapping in congenital one-handers, in line with the theory suggesting remapping in this group reflects compensatory behaviour (Muret and Makin, 2021). This is in contrast to amputees, where we find little evidence for cortical remapping, indicating a relative stability of both the hand and face representation after limb loss. By and large, remapping measures were not linked to PLP. Our results call for a reassessment of traditional remapping theories based on cortical proximity, and future research into potential remapping of the inner mouth representation after limb loss.

The data generated and analysed during this study is available to the public on Open Science Framework (https://osf.io/xq3am/).

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Author details

1. Victoria Root

   1. WIN Centre, University of Oxford, Oxford, United Kingdom
   2. Institute of Cognitive Neuroscience, University College London, London, United Kingdom
   3. Medical Research Council Cognition and Brain Sciences Unit (CBU), University of Cambridge, Cambridge, United Kingdom

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Validation, Investigation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   Contributed equally with

   Dollyane Muret

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-0500-3206

2. Dollyane Muret

   Institute of Cognitive Neuroscience, University College London, London, United Kingdom

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Supervision, Validation, Investigation, Visualization, Methodology, Writing – original draft, Project administration, Writing – review and editing

   Contributed equally with

   Victoria Root

   For correspondence

   dollyane.muret@inserm.fr

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-2626-654X

3. Maite Arribas

   1. Institute of Cognitive Neuroscience, University College London, London, United Kingdom
   2. Department of Psychosis Studies, Institute of Psychiatry, Psychology & Neuroscience, King's College London, London, United Kingdom

   Contribution

   Data curation, Formal analysis, Writing – original draft

   Competing interests

   No competing interests declared

4. Elena Amoruso

   1. Institute of Cognitive Neuroscience, University College London, London, United Kingdom
   2. Medical Research Council Cognition and Brain Sciences Unit (CBU), University of Cambridge, Cambridge, United Kingdom

   Contribution

   Investigation, Writing – original draft, Project administration

   Competing interests

   No competing interests declared

5. John Thornton

   Wellcome Trust Centre for Neuroimaging, University College London, London, United Kingdom

   Contribution

   Resources, Writing – review and editing

   Competing interests

   No competing interests declared

6. Aurelie Tarall-Jozwiak

   Queen Mary’s Hospital, London, United Kingdom

   Contribution

   Resources, Writing – review and editing

   Competing interests

   No competing interests declared

7. Irene Tracey

   WIN Centre, University of Oxford, Oxford, United Kingdom

   Contribution

   Supervision, Writing – review and editing

   Competing interests

   No competing interests declared

8. Tamar R Makin

   1. Institute of Cognitive Neuroscience, University College London, London, United Kingdom
   2. Medical Research Council Cognition and Brain Sciences Unit (CBU), University of Cambridge, Cambridge, United Kingdom
   3. Wellcome Trust Centre for Neuroimaging, University College London, London, United Kingdom

   Contribution

   Conceptualization, Resources, Supervision, Funding acquisition, Methodology, Writing – original draft, Project administration, Writing – review and editing

   Competing interests

   Senior editor, eLife

   "This ORCID iD identifies the author of this article:" 0000-0002-5816-8979

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