Biology subjects

Makin, T. R.

Publications and source records attributed to Makin, T. R..

16 recordsLinked to original sources

bioRxiv2026

Spatial localisation of touch on a robotic limb in the absence of direct haptic feedback

Artificial limbs typically lack somatosensory receptors, limiting users access to haptic feedback about movement outcomes. However, indirect tactile signals generated during object interaction are transmitted to the body-limb interface and may serve as a haptic feedback source. This study investigates whether such indirect tactile signals convey distinct information that can support localisation of touch on an artificial body part. Using an additional robotic thumb (The Third Thumb, Dani Clode Design), we measured tactile signals at the interface between the Third Thumb and the hand following stimulation from four vibration motors positioned along the Thumb. We then assessed whether participants could discriminate these signals and integrate them with different Thumb postures to spatially localise touch. Measurements showed that each stimulation site produced distinct tactile signatures at the interface. Behaviourally, participants localised touch at above-chance levels. Representational similarity analyses further revealed that performance was best explained by the spatial positions of stimuli, rather than by limb-based coordinates or indirect signal patterns alone, indicating flexible mapping of indirect touch signals onto the Third Thumb posture. Together, these findings demonstrate that indirect tactile signals transmitted through an artificial limb can be discriminated and flexibly remapped to support spatial localisation of touch.

neuroscience↗

bioRxiv2025

Motor Learning Outside the Body: Broad Skill Generalisation with an Extra Robotic Limb

Our ability to transfer motor skills across tools and contexts is what makes modern technology usable. The success of motor augmentation devices, such as supernumerary robotic limbs, hinges on users capacity for generalised motor performance. We trained participants over seven days to use an extra robotic thumb (Third Thumb, Dani Clode Design), worn on the right hand and controlled via the toes. We tested whether motor learning was confined to the specific tasks and body parts involved in controlling and interacting with the Third Thumb, or whether it could generalise beyond them. Participants showed broad skill generalisation across tasks, body postures, and even when either the Third Thumb or the controller was reassigned to a different body part, suggesting the development of abstract, body-independent motor representations. Training also reduced cognitive demands and increased the sense of agency over the device. However, participants still preferred using their biological hand over the Third Thumb when given the option, suggesting that factors beyond motor skill generalisation, cognitive effort, and embodiment must be addressed to support the real-world adoption of such technologies.

neuroscience↗

bioRxiv2025

Muscle control of an extra robotic digit

Controlling an extra robotic finger requires the brain to adapt existing motor signals. While most current strategies exploit physical movement, there is growing interest in harnessing muscle activity directly via surface electromyography (EMG) as a more seamless interface. We systematically compared muscle- (EMG) and movement-based (force sensor) control of a Third Thumb. Using identical instructions and a counterbalanced within-participants design, we assessed initial skill, learning, and cognitive load across a variety of tasks, enabling a blinded comparison across control modalities. Both control modalities afforded successful Third Thumb control and learning, although force control consistently delivered better performance. Despite execution differences, learning rates and cognitive loads were comparable, with a similar evoked sense of agency. Signal analyses showed performance was predicted by real-time force sensor parameters but not by EMG, reflecting distinct control dynamics. Nonetheless, EMG training led to greater skill transfer to force control, suggesting it may better support generalisable learning. These findings challenge the assumption that proximity to neural signals ensures better control. Although EMG underperformed in execution, it showed unique advantages, including enhanced generalisation and access to richer signals, highlighting the need for improved real-time decoding to fully exploit its potential.

neuroscience↗

bioRxiv2025

Developing a Sensory Representation of an Artificial Body Part

Somatosensory feedback is essential for motor control, yet artificial limbs are thought to lack such feedback. We investigated how the body and brain gather informative sensory signals from a wearable augmentation interface (a robotic digit for motor augmentation), and whether naturally-mediated feedback can support technological embodiment. Participants intuitively interpreted natural feedback across perceptual tasks, performing comparably to state-of-the-art artificial feedback systems. fMRI revealed an immediate and distinct, topographically-organised representation of the robotic digit. After longitudinal training, this representation was further refined, becoming more similar to the biological digits, which correlated with increased subjective somatosensory embodiment. Our findings demonstrate that wearable devices naturally provide a powerful source of feedback which is immediately integrated with our body. Long-term use can then promote device-embodiment across the sensorimotor hierarchy.

neuroscience↗

bioRxiv2024

Shaping the developing homunculus: the roles of deprivation and compensatory behaviour in sensory remapping

Some of the most dramatic examples of neuroplasticity in the human brain follow congenital sensory deprivation, though we have limited understanding of the plasticity mechanisms driving such large-scale remapping. Hand loss due to congenital limb differences (CLD) offers a unique temporal dissociation of developmental neuroplasticity mechanisms: While sensory deprivation is congenital, compensatory motor behaviours develop progressively across childhood. Using paediatric neuroimaging and semi-ecological behavioural analysis in children (5-7 years old) and adults (>25 years old) with unilateral upper-limb CLD, we studied deprivation- and use-dependent plasticity in the deprived primary somatosensory cortex and beyond. We reveal that global remapping, encompassing the entire sensory homunculus, is established early and maintained in adulthood. We demonstrate that deprivation-driven homeostatic plasticity can drive this global remapping, with Hebbian-based compensatory learning further contributing to inter-individual differences both in childhood and adulthood. Our findings emphasise the early establishment and stability of cortical maps, despite extensive daily-life behavioural adaptation.

neuroscience↗

bioRxiv2023

Stable Cortical Body Maps Before and After Arm Amputation

The adult brains capacity for cortical reorganization remains debated. Using longitudinal neuroimaging in three adults, followed up to five years before and after arm amputation, we compared cortical activity elicited by movement of the hand (pre-amputation) versus phantom hand (post-amputation) and lips (pre/post-amputation). We observed stable representations of both hand and lips. By directly quantifying activity changes across amputation, we overturn decades of animal and human research, demonstrating amputation does not trigger large- scale cortical reorganization.

neuroscience↗

bioRxiv2023

Should bionic limb control mimic the human body? Impact of control strategy on bionic hand skill learning

A longstanding engineering ambition has been to design anthropomorphic bionic limbs: devices that look like and are controlled in the same way as the biological body (biomimetic). The untested assumption is that biomimetic motor control enhances device embodiment, learning, generalization, and automaticity. To test this, we compared biomimetic and non-biomimetic control strategies for able-bodied participants when learning to operate a wearable myoelectric bionic hand. We compared motor learning across days and behavioural tasks for two training groups: Biomimetic (mimicking the desired bionic hand gesture with biological hand) and Arbitrary control (mapping an unrelated biological hand gesture with the desired bionic gesture). For both trained groups, training improved bionic limb control, reduced cognitive reliance, and increased embodiment over the bionic hand. Biomimetic users had more intuitive and faster control early in training. Arbitrary users matched biomimetic performance later in training. Further, arbitrary users showed increased generalization to a novel control strategy. Collectively, our findings suggest that biomimetic and arbitrary control strategies provide different benefits. The optimal strategy is likely not strictly biomimetic, but rather a flexible strategy within the biomimetic to arbitrary spectrum, depending on the user, available training opportunities and user requirements.

neuroscience↗

bioRxiv2021

Gender Imbalance in the Editorial Activities of a Researcher-led Journal

Editorial decision-making is a fundamental element of the scientific enterprise. We examined whether contributions to editorial decisions at various stages of the publication process is subject to gender disparity, based on analytics collected by the biomedical researcher-led journal eLife. Despite efforts to increase women representation, the board of reviewing editors (BRE) was men-dominant (69%). Moreover, authors suggested more men from the BRE pool, even after correcting for mens numerical over-representation. Although women editors were proportionally involved in the initial editorial process, they were under-engaged in editorial activities involving reviewers and authors. Additionally, converging evidence showed gender homophily in manuscripts assignment, such that men Senior Editors over-engaged men Reviewing Editors. This tendency was stronger in more gender-balanced scientific disciplines. Together, our findings confirm that gender disparities exist along the editorial process and suggest that merely increasing the proportion of women might not be sufficient to eliminate this bias.

scientific communication and education↗

bioRxiv2021

Beyond body maps: information content of specific body parts is distributed across the somatosensory Homunculus

The somatosensory homunculus in primary somatosensory cortex (S1) is topographically organised, with relatively high selectivity to each body part in its primary area. This dominant feature may eclipse other organising principles in S1. Recent multivariate methodologies allow us to identify representational features beyond selectivity, e.g., information content, providing new opportunities to characterise the homunculus. Using Representational Similarity Analysis, we asked whether body part information content can be identified in S1 beyond the primary area of a given body part. Representational dissimilarities in fMRI activity patterns were compared between different body parts (face, hand and feet) and subparts (e.g., fingers), and between different actions performed with the same body part. Throughout the S1 homunculus, we identified significant dissimilarities between non-primary body parts (e.g., between the hand and the lips in the foot area). We also observed significant dissimilarities between body subparts in distant non-primary areas (e.g., different face parts in the foot area). Finally, we could significantly dissociate between two movements performed by one body part (e.g., the hand) well beyond its primary area (e.g., in the foot and face areas), even when focusing the analysis along the most topographically organised sub-region of S1, Brodmann area 3b. Altogether, our results demonstrate that body part and action-related information content is more distributed across S1 homunculus than previously considered. While this finding does not revoke the general topographic organising principle of S1, it reveals yet unexplored underlying information contents that could be harnessed for rehabilitation, as well as novel brain-machine interfaces.

neuroscience↗

bioRxiv2021

Somatosensory signals from the controllers of an extra robotic finger support motor learning

Considerable resources are being invested to provide bidirectional control of substitutive and augmentative motor interfaces through artificial somatosensory feedback. Here, we investigated whether intrinsic somatosensory information, from body part(s) proportionally controlling an augmentation device, can be utilised to infer the devices state and position, to better support motor control and learning. In a placebo-controlled design, we used local anaesthetic to attenuate somatosensory inputs to the big toes while participants learned to operate a toe-controlled robotic extra finger (Third Thumb) using pressure sensors. Motor learning outcomes were compared against a control group who received sham anaesthetic. The availability of somatosensory cues about the amount of exerted pressure generally facilitated acquisition, retention and transfer of motor skills, and performance under cognitive load. Motor performance was not impaired by anaesthesia when tasks involved close collaboration with the biological fingers, indicating that the brain could close the gap of the missing pressure signals by alternative means, including feedback from other body parts involved in the motor task. Together, our findings demonstrate that there are intrinsic natural avenues to provide surrogate position information to support motor control of an artificial body part, beyond artificial extrinsic signalling.

neuroscience↗

bioRxiv2021

Finger representation in the cortex of the congenitally blind

Hand representation in the primary somatosensory cortex (S1) is thought to be shaped by experience. Individuals with congenital blindness rely on their sense of touch for completing daily tasks that in sighted people would be informed by vision, and possess superior tactile acuity. It has therefore been proposed that their S1 hand representation should differ from that of sighted individuals. Alternatively, it has been proposed that the improved tactile acuity in blind individuals is due to cross-modal plasticity, when regions in the occipital and temporal cortex are typically used for processing vision become activated by touch. We probed finger representation using psychophysics and 7T fMRI (1 mm3 resolution) in three individuals with bilateral anophthalmia, a rare condition in which both eyes fail to develop, as well as sighted controls. Despite anophthalmic individuals increased reliance on touch and superior tactile acuity, we found no evidence that they had more pronounced hand representation in S1. This is in line with recent research highlighting the stability of early sensory cortex, despite altered sensorimotor experience in adulthood. Unlike sighted controls, anophthalmic individuals activated the left human middle temporal complex (hMT+) during finger movement. This area did not express any hallmark of typical sensorimotor organisation, suggesting this and previously reported activity does not indicate low-level sensorimotor hand representation. However, left hMT+ contained some single finger information, beyond that found in sighted controls. This latter finding suggests that when the developmentally flexible area hMT+ is unaffected by retinal input, it can acquire novel cross-modal processes, which are potentially unrelated to the areas function in sighted people. As such, our findings highlight the opportunity for other organising principles, beyond domain specific plasticity, in shaping cross-modal reorganisation.

neuroscience↗

bioRxiv2021

Early life experience sets hard limits on motor learning as evidenced from artificial arm use

The study or artificial arms provides a unique opportunity to address long-standing questions on sensorimotor plasticity and development. Learning to use an artificial arm arguably depends on fundamental building blocks of body representation and would therefore be impacted by early-life experience. We tested artificial arm motor-control in two adult populations with upper-limb deficiency: congenital one-handers - who were born with a partial arm, and amputees - who lost their biological arm in adulthood. Brain plasticity research teaches us that the earlier we train to acquire new skills (or use a new technology) the better we benefit from this practice as adults. Instead, we found that although one-hander started using an artificial arm as toddlers, they produced increased error noise and directional errors when reaching to visual targets, relative to amputees who performed similarly to controls. However, the earlier a one-hander was fitted with an artificial arm the better their motor control was. We suggest that visuomotor integration, underlying the observed deficits, is highly dependent on either biological or artificial arm experience at a very young age. Subsequently, opportunities for sensorimotor plasticity become more limited.

neuroscience↗

bioRxiv2020

Malleability of the cortical hand map following a finger nerve block

Individual fingers in the primary somatosensory cortex (S1) are known to be represented separately and adjacently, forming a cortical hand map. Electrophysiological studies in monkeys show that finger amputation triggers increased selectivity to the neighbouring fingers within the deprived S1, causing local reorganisation. Neuroimaging research in humans, however, shows persistent S1 finger representation of the missing hand, even decades after amputation. We aimed to resolve these apparently contrasting evidence by examining finger representation in humans following pharmacological amputation using single-finger nerve block and 7T neuroimaging. We hypothesised that beneath the apparent selectivity of individual fingers in the hand map, peripheral and central processing is distributed across fingers. If each finger contributes to the cortical representation of the others, then localised input loss will weaken finger representation across the hand map. For the same reason, the non-blocked fingers will stabilise the blocked fingers representation, resulting in persistent representation of the blocked finger. Using univariate selectivity profiling, we replicated the electrophysiological findings of local S1 reorganisation. However, more comprehensive analyses confirmed that local blocking reduced representation of all fingers across the entire hand area. Importantly, multivariate analysis demonstrated that despite input loss, representation of the blocked finger remained persistent and distinct from the unblocked fingers. Computational modelling suggested that the observed findings are driven by distributed processing underlying the topographic map, combined with homeostatic mechanisms. Our findings suggest that the long-standing depiction of the somatosensory hand map is misleading. As such, accounts for map reorganisation, e.g. following amputation, need to be reconsidered.

neuroscience↗

bioRxiv2020

Neurocognitive consequences of hand augmentation

From hand tools to cyborgs, humans have long been fascinated by the opportunities afforded by augmenting ourselves. Here, we studied how motor augmentation with an extra robotic thumb (the Third Thumb) impacts the biological hand representation in the brains of able-bodied people. Participants were tested on a variety of behavioural and neuroimaging tests designed to interrogate the augmented hands representation before and after 5-days of semi-intensive training. Training improved the Thumbs motor control, dexterity and hand-robot coordination, even when cognitive load was increased or when vision was occluded, and resulted in increased sense of embodiment over the robotic Thumb. Thumb usage also weakened natural kinematic hand synergies. Importantly, brain decoding of the augmented hands motor representation demonstrated mild collapsing of the canonical hand structure following training, suggesting that motor augmentation may disrupt the biological hand representation. Together, our findings unveil critical neurocognitive considerations for designing human body augmentation.

neuroscience↗

bioRxiv2019

Similar somatotopy for active and passive digit representation in primary somatosensory cortex

Scientists traditionally use passive stimulation to examine organisational properties of the primary somatosensory cortex (SI). Recent research has, however, emphasised the close and bidirectional relationship between somatosensory and motor systems. This suggests active contributions (e.g., direct inputs from the motor system to SI) should also be considered when studying SI representations. Under such a framework, discrepant results are possible when different tasks are used to study the same underlying somatosensory representation. Here we examined whether SI hand representation is consistent when compared between active and passive tasks. Moreover, we ask whether this holds when task demands and stimulus properties are not directly matched. Using 7T fMRI, we examined three key digit representation features - spatial location, activity gradient profiles and multivariate representational structure. We show that although activity was increased overall in the active task, all key features of digit somatotopy were consistent over tasks, using both traditional univariate (activity-level) and multivariate (pattern structure) measurements. Despite overall comparability, when investigating fine-grained features of somatotopy using multivariate analysis, the active task was superior for individuating the representation across digits. We suggest that the information produced by an active task is more aligned with typical, ecologically relevant patterns of hand sensory input, resulting in more optimal SI activity patterns. Our findings validate the utilisation of active tasks for studying SI somatotopy.

neuroscience↗

bioRxiv2018

Large-scale intrinsic connectivity is consistent across varying task demands

Measuring whole-brain functional connectivity patterns based on task-free ( restingstate) spontaneous fluctuations in the functional MRI (fMRI) signal is a standard approach to probing habitual brain states, independent of task-specific context. This view is supported by spatial correspondence between task- and rest-derived connectivity networks. Yet, it remains unclear whether intrinsic connectivity observed in a resting-state acquisitions is persistent during task. Here, we sought to determine how changes in ongoing brain activation, elicited by task performance, impact the integrity of whole-brain functional connectivity patterns. We employed a steadystates paradigm, in which participants continuously executed a specific task (without baseline periods). Participants underwent separate task-based (visual, motor and visuomotor) or task-free (resting) steady-state scans, each performed over a 5-minute period. This unique design allowed us to apply a set of traditional resting-state analyses to various task-states. In addition, a classical fMRI block-design was employed to identify individualized brain activation patterns for each task, allowing to characterize how differing activation patterns across the steady-states impact whole-brain intrinsic connectivity patterns. By examining correlations across segregated brain regions (nodes) and the whole brain (using independent component analysis), we show that the whole-brain network architecture characteristic of the resting-state is robustly preserved across different steady-task states, despite striking inter-task changes in brain activation (signal amplitude). Subtler changes in functional connectivity were detected locally, within the active networks. Together, we show that intrinsic connectivity underlying the canonical resting-state networks is relatively stable even when participants are engaged in different tasks and is not limited to the resting-state.\n\nNew and Noteworthy\n\nDoes intrinsic functional connectivity (FC) reflect the canonical or transient state of the brain? We tested the consistency of the intrinsic connectivity networks across different task-conditions. We show that despite local changes in connectivity, at the whole-brain level there is little modulation in FC patterns, despite profound and large-scale activation changes. We therefore conclude that intrinsic FC largely reflects the a priori habitual state of the brain, independent of the specific cognitive context.

neuroscience↗