The Unconscious Formation of Motor and Abstract Intentions

Three separate fMRI studies were conducted to study the neural dynamics of free decision formation. In Study 1, we first searched across the brain for spatiotemporal patterns that could predict the specific outcome and timing of free motor decisions to make a left or right button press (Soon et al., 2008). In Study 2, we replicated Study 1 using ultra-high field fMRI for improved temporal and spatial resolution to more accurately characterize the evolution of decision-predictive information in prefrontal cortex (Bode et al., 2011). In Study 3, to unequivocally dissociate high-level intentions from motor preparation and execution, we investigated the neural precursors of abstract intentions as participants spontaneously decided to perform either of two mental arithmetic tasks: addition or subtraction (Soon et al., 2013). Across the three studies, we consistently found that upcoming decisions could be predicted with ~60% accuracy from fine-grained spatial activation patterns occurring a few seconds before the decisions reached awareness, with very similar profiles for both motor and abstract intentions. The content and timing of the decisions appeared to be encoded in two functionally dissociable sets of regions: frontopolar and posterior cingulate/ precuneus cortex encoded the content but not the timing of the decisions, while the pre-supplementary motor area encoded the timing but not the content of the decisions. The choice-predictive regions in both motor and abstract decision tasks overlapped partially with the default mode network. High-resolution imaging in Study 2 further revealed that as the time-point of conscious decision approached, activity patterns in frontopolar cortex became increasingly stable with respect to the final choice.:Abstract 1 1. General Introduction 5 2. Study 1: Decoding the Unconscious Formation of Motor Intentions 21 3. Study 2: Temporal Stability of Neural Patterns Involved in Intention Formation 56 4. Study 3: Decoding the Unconscious Formation of Abstract Intentions 89 5. General Discussion 119 References 14

Remedial Effects of Motivational Incentive on Declining Cognitive Control In Healthy Aging and Parkinson's Disease

The prospect of reward may provide a motivational incentive for optimizing goal-directed behavior. Animal work demonstrates that reward-processing networks and oculomotor-control networks in the brain are connected through the dorsal striatum, and that reward anticipation can improve oculomotor control via this nexus. Due perhaps to deterioration in dopaminergic striatal circuitry, goal-directed oculomotor control is subject to decline in healthy seniors, and even more in individuals with Parkinson's disease (PD). Here we examine whether healthy seniors and PD patients are able to utilize reward prospects to improve their impaired antisaccade performance. Results confirmed that oculomotor control declined in PD patients compared to healthy seniors, and in healthy seniors compared to young adults. However, the motivational incentive of reward expectation resulted in benefits in antisaccade performance in all groups alike. These findings speak against structural and non-modifiable decline in cognitive control functions, and emphasize the remedial potential of motivational incentive mechanisms in healthy as well as pathological aging

The power of prediction

Unsere Fähigkeit Erwartungen auszubilden über zukünftige Ereignisse oder die Ergebnisse unserer eigenen Handlungen ist von entscheidender Bedeutung für zielgerichtetes Verhalten. Obgleich diese Ansicht einer Reihe von einflussreichen theoretischen Strömungen in der kognitiven Psychologie zugrunde liegt, bleiben bislang wichtige Fragen dazu offen. Jüngere Ergebnisse aus neurophysiologischen und Bildgebungsstudien legen nahe, dass bewusste Ziele – ähnlich expliziter Erwartung – nicht mehr sind als ein Nebenprodukt von automatischen und unbewussten Aktivierungsmustern und keinen echten Einfluss auf Verhalten haben. Explizite subjektive Erwartung wird vielmals abgetan, weil sie kein zuverlässiges Maß „wahrer“ mentaler Prozesse darstelle, oder weil sie nicht notwendig sei, um Verhalten zu erklären. In der vorliegenden Arbeit beschäftige ich mich mit der Rolle expliziter subjektiver Erwartung und untersuche die folgenden Fragen: (1) Sind verbalisierte Vorhersagen ein valider Indikator für intern generierte Erwartungen? (2) Haben solche Erwartungen tatsächlich einen Einfluss auf Handlungsvorbereitung? (3) Worin besteht der Unterschied zwischen Erwartungen, die auf äußeren oder inneren Quellen beruhen? Die Ergebnisse aus drei Studien, die ich im Rahmen dieser Dissertation durchgeführt habe, belegen, dass verbalisierte Vorhersagen ein valider Indikator von subjektiver Erwartung sind. Subjektive Erwartung kann nicht als ein Nebenprodukt von Handlungsvorbereitung verstanden werden, sondern trägt zu dieser bei und spielt daher eine wichtige Rolle in der Handlungskontrolle. Selbstgenerierte Erwartung beeinflusst gegenüber Cue-induzierter Erwartung verstärkt frühe Aufmerksamkeits- und zentrale Entscheidungsprozesse. Selbstgenerierte Vorhersagen führen zu verstärkten Verhaltenseffekten, und zwar sowohl bei Stimuluserwartungen als auch bei Erwartungen bezüglich einer Aufgabe. Subjektive Erwartung ist demnach auch entscheidend für kognitive Kontrolle.Our ability to form expectations about future events or the results of our own actions is crucial for efficient behavior. While this notion underlies a range of influential theoretical approaches in cognitive psychology, from reinforcement learning to ideo-motor theory, a number of open questions remain. Recent results from neurophysiological and brain imaging studies suggest that conscious intention – or explicit expectation – is nothing more than a by-product of automatic and unconscious activation patterns, without any real impact on behavior. Explicit subjective expectation has been dismissed by many researchers who regard it either an unreliable measure of “true” mental processes, or question its necessity in explaining behavior in general. In the current work, I focus on the role of explicit subjective expectation and attempt to answer the following questions: (1) Are verbalized predictions a valid indicator of internally generated expectations? (2) Do such expectations really affect action preparation? (3) What are the differences between expectations arising from external or internal sources? Results from the three studies conducted within the scope of this dissertation demonstrate that verbalized predictions are in fact a valid indicator of subjective expectation and are suitable for use in experimental paradigms. Also, subjective expectation cannot be described as a mere by-product of preparation, but instead feeds into preparation and therefore plays a role in action control. Self-generated expectation was shown to involve early attentional and central decision processes to a greater degree than cue-induced expectation. Consequently, self-generated predictions entailed greater behavioral effects not only for stimulus expectations, but also for expectations regarding a task set. Subjective expectation is therefore also elemental in cognitive control

Do we dance because we walk? The impact of regular vestibular experience on the early development of beat production and perception

Movement to music is a universal human behaviour (Savage, Brown, Sakai & Currie, 2015). Whilst the strong link between music and movement is clearly bidirectional, the origins are not clear. Studying the emergence of rhythmic skills through infancy provides a window into the perceptual and physical attributes, experience, and contexts necessary, to attain the basics of human musicality. This thesis asks whether the human experience of bipedal locomotion, as a primary source of regular vestibular information, is crucial for sensorimotor synchronisation (SMS), spontaneous motor tempo (SMT), and impacts rhythm perception. The first experiment evidences the emergence of tempo-flexibility when moving to music between 10- and 18-months-of-age. The following study is the first to show that experience of locomotion, including from infant carrying, predicts the temporal matching of infant movement to music. Curious if carrying practices influence the very rhythms that we naturally produce, a large-scale correlational study finds infant SMT is predicted by parent height, but not infant’s own body size, such that infants with taller caregivers show a slower SMT than those with shorter caregivers. We contend that this reflects infant experience of being carried by their caregiver. The fourth experiment confirms that experience of being carried at a novel tempo can alter the rhythms infant spontaneously produce. Finally, we asked how information from being carried during locomotion might be changing rhythm perception; specifically, if infants show greater activation of their sensorimotor system when hearing rhythms that match the tempo at which they were carried. Combined, these studies present a highly original piece of research into the ways in which early experiences of locomotion may impact fundamental musical skill

Semantic and motor processes in infant perception of object-directed and tool-mediated action

Actions are the translation of internal states such as intentions into overt gestures and goals. Actions are communicative, because by observing another’s overt behaviour we can infer that person’s internal states. Infants’ abilities to execute actions are limited by developing motor processes. Their capacity to make inferences from others’ behaviour is hindered by their inability to engage in perspective-taking and other advanced social cognitive processes. Nonetheless, extensive evidence shows that infants perceive actions as goal-directed sequences that are meaningful, and that they respond to observed actions with motor resonance. The aims of this thesis were to determine how semantic and motor processing of observed action develop in infancy, whether these processes develop separately or in conjunction with one another, and how infants’ abilities to execute and plan actions affects ability to detect semantic and motor differences between actions. These aims were achieved by studying how infants processed grasping actions that varied on different dimensions. In Chapter 1, the literature on infant action perception from social, motor and semantic perspectives is reviewed and the objectives of the thesis are described. In Chapter 2, the ability of 16-month-olds to discriminate between the uses of a novel tool when motor simulation processes are uninformative was investigated. In Chapter 3, the attentional and semantic neural correlates of processing of observed grasps were measured in 9-month-olds, 11.5-month-olds, and adults. In Chapter 4, motor activation in 10-month-old infants in response to motorically similar but semantically distinct grasping actions was related to infants’ action planning skills. The results of these experiments show that there is a complex interplay between motor and semantic constituents of the action processing system, and that this interplay is developmentally dynamic. The implications of the results for understanding action processing in development are considered in Chapter 5

Effects of pro-social motives and reward on children's prospective memory and inhibitory control

Prospective memory (PM) refers to the ability to remember to perform an intended action in the future whereas inhibitory control (IC) is defined as the capacity to stop a prepotent or automatic response in favour of a more correct or pertinent response in a certain situation. The main aim of the present thesis was to investigate the role of motivational factors – such as the social values of goals and presence of rewards – in children’s PM and IC abilities by using an event-based PM task (EBPM) and a RI task (Go/No-Go) modelled after Brandimonte et al. (2011). RI plays a pivotal role in executive control (e.g., Andres, 2003; Aron, 2007; Logan, 1985a; Miyake et al., 2000; Stuphorn & Schall, 2006). RI concerns the ability to withhold responses that are unsuitable or not useful, so facilitating behaviour that is both flexible and goal-orientated in constantly changing environments. RI ability is called upon frequently in everyday life to prevent us from committing potentially harmful actions, for example, from stepping into the road when a car comes around a corner without seeing you. Go/No-go task is typically used in order to measure RI ability. Only two types of stimuli are used in the conventional Go/No-go task: a Go stimulus, and a No-go stimulus. Instructions are given to participants to respond rapidly, usually by pressing a button, only when Go stimuli appear, whereas they have to refrain from pressing the button on the presentation of No-go stimuli; response inhibition refers to the ability to stop oneself responding to No-go stimuli. Usually, Go stimuli appear more frequently in this kind of task, such as to predispose a participant to responding and increase the amount of inhibitory effort needed to not respond when presented with No-go stimuli (Simmonds et al., 2008). Performance in RI paradigms can be thought of as an independent “horse race” in which there is a go process prompted by go stimuli, and a stop process prompted by No-go stimuli (Logan & Cowan, 1984; Logan, Van Zandt, Verbruggen, & Wagenmakers, 2014; Verbruggen & Logan, 2009b). RI is successful when the stop process is underway before the go process and no response is made (signal-inhibit); when the go process starts before the stop process, response inhibition is unsuccessful as a response is made inappropriately (signal-respond).This choice wase made because these two tasks mentioned above seem to involve two types of intention that differ with respect to the direction of the intended action. While both the EBPM and RI tasks require forming, maintaining, and realising delayed intentions, the EBPM task involves the overt execution of the intended action, while a response inhibition task necessitates the suppression of the predominant response (Brandimonte et al., 2011). In summary, these two tasks could be considered as being in parallel in all respects apart from the response direction, as the former consists of remembering to perform an action, whereas the latter involves refraining from acting. Of particular importance are the EF abilities needed for each task type; task-shifting abilities are predominantly required during EBPM tasks when a participant must switch from the ongoing activity to the execution of a certain action when a target cue is seen. The RI task, by contrast, principally involves inhibitory control abilities, as the participant must remember not to perform the ongoing activity when a target cue appears. Despite this contrast, switching an inhibition are not mutually exclusive; switching is still involved in an RI task, inasmuch as the participant switches from the task of performing (ongoing), to the task of not performing (RI – no response). Nevertheless, the pivotal role in the RI task is played by the active suppression of actions that would be counterproductive to the achievement of the predefined goal of the task. This comparison of RI and EBPM tasks applies particularly to these two kinds of task, as both of them involve performing two types of task simultaneously, that is the ongoing task and additionally either the PM or the RI task. However, the same comparison cannot be made between TBPM and RI tasks, since the TBPM is not based on the appearance of a target cue, which however does occur during the RI and EBPM tasks. Indeed, in TBPM tasks, intentions must be executed only after a certain period of time has passed or at a predetermined point in time in the future (Wang et al., 2008); for example, recalling the need to take medicine at a given interval, or remembering to call a friend on their birthday. By contrast, in EBPM tasks an external cue should in theory remind the participant to perform the intended action (Talbot & Kerns, 2014); examples include remembering to pass on a message to a friend when they are next seen.Overall, the present research includes three large studies testing children aged 4-5 years, 6-7 years, 7-8 years, 10-11 years, and an adult group. Specifically, the following research issues have been explored: a) whether motivational factors, such as pro-sociality and reward, can have effects on children’s memory for intentions and response inhibition (RI) and, if any, b) whether these effects differ as a function of task (PM and RI); c) whether children’s PM and RI performance differ from that of adults when pro-sociality is involved. Results highlighted a significant interaction between Pro-sociality and Task, indicating that children had worse PM performance under the condition with pro-sociality (Study 1). In contrast, pro-sociality improved adults’ but not children’s performance (Study 3), as qualified by a significant interaction between Pro-sociality and Age. Significant effects of Reward emerged when Task factor (Go/No-go task / PM task) was partialled out (Study 2), showing that children performed better in conditions with a reward but only in the Go/No-Go task. A significant effect of Task was found in Study 2 and Study 3, such that participants had higher scores in Go/No-Go than in PM tasks. In conclusion, this Ph.D. project adds to the evidence that while PM and IC may have some commonalities because they are both linked to intentions (to do something or not to do something), they seem to rely on different mechanisms as indicated by the differential effects of Task, Pro-sociality, and Reward

Do we dance because we walk? The impact of regular vestibular experience on the early development of beat production and perception

Movement to music is a universal human behaviour (Savage, Brown, Sakai & Currie, 2015). Whilst the strong link between music and movement is clearly bidirectional, the origins are not clear. Studying the emergence of rhythmic skills through infancy provides a window into the perceptual and physical attributes, experience, and contexts necessary, to attain the basics of human musicality. This thesis asks whether the human experience of bipedal locomotion, as a primary source of regular vestibular information, is crucial for sensorimotor synchronisation (SMS), spontaneous motor tempo (SMT), and impacts rhythm perception. The first experiment evidences the emergence of tempo-flexibility when moving to music between 10- and 18-months-of-age. The following study is the first to show that experience of locomotion, including from infant carrying, predicts the temporal matching of infant movement to music. Curious if carrying practices influence the very rhythms that we naturally produce, a large-scale correlational study finds infant SMT is predicted by parent height, but not infant’s own body size, such that infants with taller caregivers show a slower SMT than those with shorter caregivers. We contend that this reflects infant experience of being carried by their caregiver. The fourth experiment confirms that experience of being carried at a novel tempo can alter the rhythms infant spontaneously produce. Finally, we asked how information from being carried during locomotion might be changing rhythm perception; specifically, if infants show greater activation of their sensorimotor system when hearing rhythms that match the tempo at which they were carried. Combined, these studies present a highly original piece of research into the ways in which early experiences of locomotion may impact fundamental musical skill

Action-Related Representations

Theories of grounded cognition state that there is a meaningful connection between action and cognition. Although these claims are widely accepted, the nature and structure of this connection is far from clear and is still a matter of controversy. This book argues for a type of cognitive representation that essentially combines cognition and action, and which is foundational for higher-order cognitive capacities