Paradoxical evidence weighting in confidence judgments for detection and discrimination.

When making discrimination decisions between two stimulus categories, subjective confidence judgments are more positively affected by evidence in support of a decision than negatively affected by evidence against it. Recent theoretical proposals suggest that this "positive evidence bias" may be due to observers adopting a detection-like strategy when rating their confidence-one that has functional benefits for metacognition in real-world settings where detectability and discriminability often go hand in hand. However, it is unknown whether, or how, this evidence-weighting asymmetry affects detection decisions about the presence or absence of a stimulus. In four experiments, we first successfully replicate a positive evidence bias in discrimination confidence. We then show that detection decisions and confidence ratings paradoxically suffer from an opposite "negative evidence bias" to negatively weigh evidence even when it is optimal to assign it a positive weight. We show that the two effects are uncorrelated and discuss our findings in relation to models that account for a positive evidence bias as emerging from a confidence-specific heuristic, and alternative models where decision and confidence are generated by the same, Bayes-rational process.

Stable Cortical Body Maps Before and After Arm Amputation.

Neuroscientists have long debated the adult brain's capacity to reorganize itself in response to injury. A driving model for studying plasticity has been limb amputation. For decades, it was believed that amputation triggers large-scale reorganization of cortical body resources. However, these studies have relied on cross-sectional observations post-amputation, without directly tracking neural changes. Here, we longitudinally followed adult patients with planned arm amputations and measured hand and face representations, before and after amputation. By interrogating the representational structure elicited from movements of the hand (pre-amputation) and phantom hand (post-amputation), we demonstrate that hand representation is unaltered. Further, we observed no evidence for lower face (lip) reorganization into the deprived hand region. Collectively, our findings provide direct and decisive evidence that amputation does not trigger large-scale cortical reorganization.

A demonstration of cone function plasticity after gene therapy in achromatopsia.

Recent advances in regenerative therapy have placed the treatment of previously incurable eye diseases within arms' reach. Achromatopsia is a severe monogenic heritable retinal disease that disrupts cone function from birth, leaving patients with complete colour blindness, low acuity, photosensitivity and nystagmus. While successful gene-replacement therapy in non-primate models of achromatopsia has raised widespread hopes for clinical treatment, it was yet to be determined if and how these therapies can induce new cone function in the human brain. Using a novel multimodal approach, we demonstrate for the first time that gene therapy can successfully activate dormant cone-mediated pathways in children with achromatopsia (CNGA3- and CNGB3-associated, 10-15 years). To test this, we combined functional MRI population receptive field mapping and psychophysics with stimuli that selectively measure cone photoreceptor signalling. We measured cortical and visual cone function before and after gene therapy in four paediatric patients, evaluating treatment-related change against benchmark data from untreated patients (n = 9) and normal-sighted participants (n = 28). After treatment, two of the four children displayed strong evidence for novel cone-mediated signals in visual cortex, with a retinotopic pattern that was not present in untreated achromatopsia and which is highly unlikely to emerge by chance. Importantly, this change was paired with a significant improvement in psychophysical measures of cone-mediated visual function. These improvements were specific to the treated eye, and provide strong evidence for successful read-out and use of new cone-mediated information. These data show for the first time that gene replacement therapy in achromatopsia within the plastic period of development can awaken dormant cone-signalling pathways after years of deprivation. This reveals unprecedented neural plasticity in the developing human nervous system and offers great promise for emerging regenerative therapies.

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

The study of 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 deficiencies: a congenital group-individuals who were born with a partial arm, and an acquired group-who lost their arm following amputation 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 the congenital group started using an artificial arm as toddlers, they produced increased error noise and directional errors when reaching to visual targets, relative to the acquired group who performed similarly to controls. However, the earlier an individual with a congenital limb difference was fitted with an artificial arm, the better their motor control was. Since we found no group differences when reaching without visual feedback, we suggest that the ability to perform efficient visual-based corrective movements is highly dependent on either biological or artificial arm experience at a very young age. Subsequently, opportunities for sensorimotor plasticity become more limited.

Robotic hand augmentation drives changes in neural body representation.

Humans have long been fascinated by the opportunities afforded through augmentation. This vision not only depends on technological innovations but also critically relies on our brain's ability to learn, adapt, and interface with augmentation devices. Here, we investigated whether successful motor augmentation with an extra robotic thumb can be achieved and what its implications are on the neural representation and function of the biological hand. Able-bodied participants were trained to use an extra robotic thumb (called the Third Thumb) over 5 days, including both lab-based and unstructured daily use. We challenged participants to complete normally bimanual tasks using only the augmented hand and examined their ability to develop hand-robot interactions. Participants were tested on a variety of behavioral and brain imaging tests, designed to interrogate the augmented hand's representation before and after the training. Training improved Third Thumb motor control, dexterity, and hand-robot coordination, even when cognitive load was increased or when vision was occluded. It also resulted in increased sense of embodiment over the Third Thumb. Consequently, augmentation influenced key aspects of hand representation and motor control. Third Thumb usage weakened natural kinematic synergies of the biological hand. Furthermore, brain decoding revealed a mild collapse of the augmented hand's motor representation after training, even while the Third Thumb was not worn. Together, our findings demonstrate that motor augmentation can be readily achieved, with potential for flexible use, reduced cognitive reliance, and increased sense of embodiment. Yet, augmentation may incur changes to the biological hand representation. Such neurocognitive consequences are crucial for successful implementation of future augmentation technologies.

Expert Tool Users Show Increased Differentiation between Visual Representations of Hands and Tools.

The idea that when we use a tool we incorporate it into the neural representation of our body (embodiment) has been a major inspiration for philosophy, science, and engineering. While theoretically appealing, there is little direct evidence for tool embodiment at the neural level. Using functional magnetic resonance imaging (fMRI) in male and female human subjects, we investigated whether expert tool users (London litter pickers: n = 7) represent their expert tool more like a hand (neural embodiment) or less like a hand (neural differentiation), as compared with a group of tool novices (n = 12). During fMRI scans, participants viewed first-person videos depicting grasps performed by either a hand, litter picker, or a non-expert grasping tool. Using representational similarity analysis (RSA), differences in the representational structure of hands and tools were measured within occipitotemporal cortex (OTC). Contrary to the neural embodiment theory, we find that the experts group represent their own tool less like a hand (not more) relative to novices. Using a case-study approach, we further replicated this effect, independently, in five of the seven individual expert litter pickers, as compared with the novices. An exploratory analysis in left parietal cortex, a region implicated in visuomotor representations of hands and tools, also indicated that experts do not visually represent their tool more similar to hands, compared with novices. Together, our findings suggest that extensive tool use leads to an increased neural differentiation between visual representations of hands and tools. This evidence provides an important alternative framework to the prominent tool embodiment theory.SIGNIFICANCE STATEMENT It is commonly thought that tool use leads to the assimilation of the tool into the neural representation of the body, a process referred to as embodiment. Here, we demonstrate that expert tool users (London litter pickers) neurally represent their own tool less like a hand (not more), compared with novices. Our findings advance our current understanding for how experience shapes functional organization in high-order visual cortex. Further, this evidence provides an alternative framework to the prominent tool embodiment theory, suggesting instead that experience with tools leads to more distinct, separable hand and tool representations.

Talking with Your (Artificial) Hands: Communicative Hand Gestures as an Implicit Measure of Embodiment.

When people talk, they move their hands to enhance meaning. Using accelerometry, we measured whether people spontaneously use their artificial limbs (prostheses) to gesture, and whether this behavior relates to everyday prosthesis use and perceived embodiment. Perhaps surprisingly, one- and two-handed participants did not differ in the number of gestures they produced in gesture-facilitating tasks. However, they did differ in their gesture profile. One-handers performed more, and bigger, gesture movements with their intact hand relative to their prosthesis. Importantly, one-handers who gestured more similarly to their two-handed counterparts also used their prosthesis more in everyday life. Although collectively one-handers only marginally agreed that their prosthesis feels like a body part, one-handers who reported they embody their prosthesis also showed greater prosthesis use for communication and daily function. Our findings provide the first empirical link between everyday prosthesis use habits and perceived embodiment and a novel means for implicitly indexing embodiment.

Is an artificial limb embodied as a hand? Brain decoding in prosthetic limb users.

The potential ability of the human brain to represent an artificial limb as a body part (embodiment) has been inspiring engineers, clinicians, and scientists as a means to optimise human-machine interfaces. Using functional MRI (fMRI), we studied whether neural embodiment actually occurs in prosthesis users' occipitotemporal cortex (OTC). Compared with controls, different prostheses types were visually represented more similarly to each other, relative to hands and tools, indicating the emergence of a dissociated prosthesis categorisation. Greater daily life prosthesis usage correlated positively with greater prosthesis categorisation. Moreover, when comparing prosthesis users' representation of their own prosthesis to controls' representation of a similar looking prosthesis, prosthesis users represented their own prosthesis more dissimilarly to hands, challenging current views of visual prosthesis embodiment. Our results reveal a use-dependent neural correlate for wearable technology adoption, demonstrating adaptive use-related plasticity within the OTC. Because these neural correlates were independent of the prostheses' appearance and control, our findings offer new opportunities for prosthesis design by lifting restrictions imposed by the embodiment theory for artificial limbs.

Motor control drives visual bodily judgements.

The 'embodied cognition' framework proposes that our motor repertoire shapes visual perception and cognition. But recent studies showing normal visual body representation in individuals born without hands challenges the contribution of motor control on visual body representation. Here, we studied hand laterality judgements in three groups with fundamentally different visual and motor hand experiences: two-handed controls, one-handers born without a hand (congenital one-handers) and one-handers with an acquired amputation (amputees). Congenital one-handers, lacking both motor and first-person visual information of their missing hand, diverged in their performance from the other groups, exhibiting more errors for their intact hand and slower reaction-times for challenging hand postures. Amputees, who have lingering non-visual motor control of their missing (phantom) hand, performed the task similarly to controls. Amputees' reaction-times for visual laterality judgements correlated positively with their phantom hand's motor control, such that deteriorated motor control associated with slower visual laterality judgements. Finally, we have implemented a computational simulation to describe how a mechanism that utilises a single hand representation in congenital one-handers as opposed to two in controls, could replicate our empirical results. Together, our findings demonstrate that motor control is a driver in making visual bodily judgments.