Control Of Goal-directed Behavior Research Articles

Differential regulation of G protein-coupled receptor-associated proteins in the caudate and the putamen of cynomolgus macaques following chronic ethanol drinking.

The dorsal striatum is composed of the caudate nucleus and the putamen in human and non-human primates. These two regions receive different cortical projections and are functionally distinct. The caudate is involved in the control of goal-directed behaviors, while the putamen is implicated in habit learning and formation. Previous reports indicate that ethanol differentially influences neurotransmission in these two regions. Because neurotransmitters primarily signal through G protein-coupled receptors (GPCRs) to modulate neuronal activity, the present study aimed to determine whether ethanol had a region-dependent impact on the expression of proteins that are involved in the trafficking and function of GPCRs, including G protein subunits and their effectors, protein kinases, and elements of the cytoskeleton. Western blotting was performed to examine protein levels in the caudate and the putamen of male cynomolgus macaques that self-administered ethanol for 1 year under free access conditions, along with control animals that self-administered an isocaloric sweetened solution under identical operant conditions. Among the 18 proteins studied, we found that the levels of one protein (PKCβ) were increased, and 13 proteins (Gαi1/3, Gαi2, Gαo, Gβ1γ, PKCα, PKCε, CaMKII, GSK3β, β-actin, cofilin, α-tubulin, and tubulin polymerization promoting protein) were reduced in the caudate of alcohol-drinking macaques. However, ethanol did not alter the expression of any proteins examined in the putamen. These observations underscore the unique vulnerability of the caudate nucleus to changes in protein expression induced by chronic ethanol exposure. Whether these alterations are associated with ethanol-induced dysregulation of GPCR function and neurotransmission warrants future investigation.

Inflammatory processes in the prefrontal cortex induced by systemic immune challenge: Focusing on neurons

Peripheral immune challenge induces neurobiological alterations in the brain and related neuropsychiatric symptoms both in humans and other mammals. One of the best known physiological effects of systemic inflammation is sickness behavior. However, in addition to this depression-like state, there are other cognitive outcomes of peripherally induced neuroinflammation that can be linked to the dysfunction of higher-order cortical areas, such as the prefrontal cortex (PFC). As the physiological activity of the PFC is largely based on the balanced interplay of excitatory pyramidal cells and inhibitory interneurons, it may be hypothesized that neuroinflammatory processes result in a shift of excitatory/inhibitory balance, which is a common hallmark of several neuropsychiatric conditions. Indeed, many data suggest that peripherally induced neuroinflammation is strongly associated with molecular and functional changes in PFC neurons leading to disturbances in their synaptic networks. Different experimental approaches may cause some incongruence in the reviewed data. However, it is commonly agreed that acute systemic inflammation leads to changes in the excitatory/inhibitory balance in the PFC by proinflammatory signaling at the brain borders and in the brain parenchyma. These cellular changes result in altered local and brain-wide network activity inducing disturbances in the top-down control of goal-directed behavior and cognition regulated by the PFC. Lipopolysaccharide (LPS)-treated rodents are the most widely used experimental models of peripherally induced neuroinflammation, so the majority of the reviewed data come from studies utilizing the LPS model. This may limit their general interpretation regarding the neuronal effects of peripheral immune activation. In addition, several biological variables (e.g., sex, age) can influence the PFC effects of systemic immune challenge, not only the nature and severity of immune activation. Therefore, it would be desirable to investigate inflammation-related neuronal changes in the PFC using other models of systemic inflammation as well, and to focus on the targeted fine-tuning of the affected cell types via common molecular mechanisms of the immune and nervous systems.

Chemogenetic Perturbation of the Posterior But Not Anterior Cerebellum Reduces Voluntary Ethanol Consumption.

The cerebellum communicates with brain areas critically involved in control of goal-directed behaviors including the prefrontal and orbitofrontal cortices and midbrain and basal ganglia structures. In particular, the posterior cerebellum is important for cognitive flexibility and has been implicated in alcohol and drug-related memory. We hypothesized that the cerebellum, through its multiple connections to reward-related brain circuitry, regulates alcohol consumption. To test this, we expressed inhibitory designer receptors exclusively activated by designer drugs (DREADDs) in molecular layer interneurons (MLIs) in anterior (IV-V) or posterior (VI-VIII) cerebellar lobules of male and female mice and activated them during alcohol drinking sessions. In a home-cage drinking paradigm, alcohol consumption was significantly decreased by clozapine-N-oxide (CNO) or deschloroclozapine (DCZ) administration in male mice expressing DREADDs in posterior but not anterior lobules. CNO/DCZ injections did not affect drinking in DREADD expressing female mice or in male mice expressing the control vector. Activation of DREADDs expressed in anterior or posterior lobules had no effect on sucrose or quinine consumption in male or female mice. During operant self-administration sessions, DCZ decreased the number of licks and bouts in male but not female mice expressing DREADDs in posterior lobules with no effect in control vector mice. Performance on an accelerated rotarod was unaffected by chemogenetic manipulation while distance traveled in the open field was decreased by DREADD activation in anterior but not posterior lobules. These results indicate that neuronal activity within the posterior cerebellar cortex plays an important role in the control of alcohol consumption in male mice.

Perception-Action Integration Is Altered in Functional Movement Disorders.

Although functional neurological movement disorders (FMD) are characterized by motor symptoms, sensory processing has also been shown to be disturbed. However, how the integration of perception and motor processes, essential for the control of goal-directed behavior, is altered in patients with FMD is less clear. A detailed investigation of these processes is crucial to foster a better understanding of the pathophysiology of FMD and can systematically be achieved in the framework of the theory of event coding (TEC). The aim was to investigate perception-action integration processes on a behavioral and neurophysiological level in patients with FMD. A total of 21 patients and 21 controls were investigated with a TEC-related task, including concomitant electroencephalogram (EEG) recording. We focused on EEG correlates established to reflect perception-action integration processes. Temporal decomposition allowed to distinguish between EEG codes reflecting sensory (S-cluster), motor (R-cluster), and integrated sensory-motor processing (C-cluster). We also applied source localization analyses. Behaviorally, patients revealed stronger binding between perception and action, as evidenced by difficulties in reconfiguring previously established stimulus-response associations. Such hyperbinding was paralleled by a modulation of neuronal activity clusters, including reduced C-cluster modulations of the inferior parietal cortex and altered R-cluster modulations in the inferior frontal gyrus. Correlations of these modulations with symptom severity were also evident. Our study shows that FMD is characterized by altered integration of sensory information with motor processes. Relations between clinical severity and both behavioral performance and neurophysiological abnormalities indicate that perception-action integration processes are central and a promising concept for the understanding of FMD. © 2023 The Authors. Movement Disorders published by Wiley Periodicals LLC on behalf of International Parkinson and Movement Disorder Society.

Longitudinal effects of early psychosocial deprivation on macaque executive function: Evidence from computational modelling.

Executive function (EF) describes a group of cognitive processes underlying the organization and control of goal-directed behaviour. Environmental experience appears to play a crucial role in EF development, with early psychosocial deprivation often linked to EF impairment. However, many questions remain concerning the developmental trajectories of EF after exposure to deprivation, especially concerning specific mechanisms. Accordingly, using an 'A-not-B' paradigm and a macaque model of early psychosocial deprivation, we investigated how early deprivation influences EF development longitudinally from adolescence into early adulthood. The contribution of working memory and inhibitory control mechanisms were examined specifically via the fitting of a computational model of decision making to the choice behaviour of each individual. As predicted, peer-reared animals (i.e. those exposed to early psychosocial deprivation) performed worse than mother-reared animals across time, with the fitted model parameters yielding novel insights into the functional decomposition of group-level EF differences underlying task performance. Results indicated differential trajectories of inhibitory control and working memory development in the two groups. Such findings not only extend our knowledge of how early deprivation influences EF longitudinally, but also provide support for the utility of computational modelling to elucidate specific mechanisms linking early psychosocial deprivation to long-term poor outcomes.

Does stress consistently favor habits over goal-directed behaviors? Data from two preregistered exact replication studies

Instrumental learning is controlled by two distinct parallel systems: goal-directed (action-outcome) and habitual (stimulus-response) processes. Seminal research by Schwabe and Wolf (2009, 2010) has demonstrated that stress renders behavior more habitual by decreasing goal-directed control. More recent studies yielded equivocal evidence for a stress-induced shift towards habitual responding, yet these studies used different paradigms to evaluate instrumental learning or used different stressors. Here, we performed exact replications of the original studies by exposing participants to an acute stressor either before (cf. Schwabe and Wolf, 2009) or directly after (cf. Schwabe and Wolf, 2010) an instrumental learning phase in which they had learned that distinct actions led to distinct, rewarding food outcomes (i.e., instrumental learning). Then, following an outcome devaluation phase in which one of the food outcomes was consumed until participants were satiated, action-outcome associations were tested in extinction. Despite successful instrumental learning and outcome devaluation and increased subjective and physiological stress levels following stress exposure, the stress and no-stress groups in both replication studies responded indifferently to valued and devalued outcomes. That is, non-stressed participants failed to demonstrate goal-directed behavioral control, thereby rendering the critical test of a shift from goal-directed to habitual control in the stress group inapt. Several reasons for these replication failures are discussed, including the rather indiscriminate devaluation of outcomes that may have contributed to indifferent responding during extinction, which emphasize the need to further our understanding of the boundary conditions in research aimed at demonstrating a stress-induced shift towards habitual control.

Voluntary Control of Cognitive Activity in Preschool Children: Age-dependent Changes from Ages 3-4 to 4-5.

Voluntary control of goal-directed behavior and mental activity in preschool children plays a key role in knowledge acquisition and future academic achievement. Studies of voluntary control have mainly concerned 6-8-year old children; much less is known about the ability to exercise voluntary control at early ages. Due to the high prognostic value of the level of development of voluntary control and heterogeneous development of their individual components, it seems actually useful to study age-related changes of these components in children from 3-4 to 4-5 years old. To compare age-related changes in executive functions (EF) in children age 3-4 years (mean age: 3.5±0.2 yrs; n = 49; 31 boys) and 4-5 years (mean age: 4.5±0.3 yrs; n = 70; 35 boys). To assess the different components of EF we used: 1) a qualitative group and individual testing procedure based on the principles of Luria's theory of the dynamic localization and organization of higher mental functions; and 2) a computerized testing procedure which included the Bourdon-Wiersma cancellation test, the "Hearts and Flowers" conflict test (the Dots task), and the Corsi block-tapping test. The results showed that different components of voluntary control developed at different rates (heterochronically): there were significant progressive changes from 3-4 to 4-5 years for working memory, assimilation of instructions, switching between separate actions, selective concentration on a target or task, and the distribution of attention. Some other components of EF did not show significant positive dynamics during this period. The results indicate the importance of applying the activity theory approach to the development of cognitive processes in preschool age.

A lineage explanation of human normative guidance: the coadaptive model of instrumental rationality and shared intentionality

This paper aims to contribute to the existing literature on normative cognition by providing a lineage explanation of human social norm psychology. This approach builds upon theories of goal-directed behavioral control in the reinforcement learning and control literature, arguing that this form of control defines an important class of intentional normative mental states that are instrumental in nature. I defend the view that great ape capacities for instrumental reasoning and our capacity (or family of capacities) for shared intentionality coadapted to each other and argue that the evolution of this capacity has allowed the representation of social norms and the emergence of our capacity for normative guidance.

Dysfunction of goal-directed control in patients with depression and nonsuicidal self-injury.

BackgroundNon‐suicidal self‐injury (NSSI) is a common problem associated with dangerous outcomes. Dysfunction of goal‐directed behavioral control may contribute to NSSI. To test this, we used a novel experimental paradigm (Pavlovian‐to‐Instrumental Transfer, PIT) to test whether patients with NSSI utilize Pavlovian conditioned stimuli (CSs) during goal‐directed control of ongoing behavior.MethodsThirty‐five depressed patients with NSSI (D‐NSSI) and thirty‐four healthy controls performed a PIT task. We measured the influence of positive and negative background CSs on instrumental responses for rewards.ResultsThe results showed that D‐NSSI performed significantly lower PIT than controls, and PIT measures were negatively correlated with NSSI frequency. Furthermore, in a subset of patients exhibiting high levels of compulsivity, PIT positively moderated the relationship between compulsivity and NSSI frequency.ConclusionsThese results indicate that D‐NSSI patients have difficulties in using different CSs to control ongoing behavior in a goal‐directed manner, and the dysfunction of goal‐directed control may contribute to NSSI.

A Canonical Scheme of Bottom-Up and Top-Down Information Flows in the Frontoparietal Network.

Goal-directed behavior often involves temporal separation and flexible context-dependent association between sensory input and motor output. The control of goal-directed behavior is proposed to lie in the frontoparietal network, but the computational architecture of this network remains elusive. Based on recent rodent studies that measured and manipulated projection neurons in the frontoparietal network together with findings from earlier primate studies, we propose a canonical scheme of information flows in this network. The parietofrontal pathway transmits the spatial information of a sensory stimulus or internal motor bias to drive motor programs in the frontal areas. This pathway might consist of multiple parallel connections, each controlling distinct motor effectors. The frontoparietal pathway sends the spatial information of cognitively processed motor plans through multiple parallel connections. Each of these connections could support distinct spatial functions that use the motor target information, including attention allocation, multi-body part coordination, and forward estimation of movement state (i.e., forward models). The parallel pathways in the frontoparietal network enable dynamic interactions between regions that are tuned for specific goal-directed behaviors. This scheme offers a promising framework within which the computational architecture of the frontoparietal network and the underlying circuit mechanisms can be delineated in a systematic way, providing a holistic understanding of information processing in this network. Clarifying this network may also improve the diagnosis and treatment of behavioral deficits associated with dysfunctional frontoparietal connectivity in various neurological disorders including Alzheimer’s disease.

If I do A, B will happen: Dissecting circuits detecting causal relations between actions and outcomes in marmoset prefrontal cortex

In this issue of Neuron, Duan etal. (2021) use pharmacological manipulation to reveal opposing influences of anterior cingulate and orbitofrontal cortex of marmosets on decisions that are based on action-outcome associations.

GOALIATH: a theory of goal-directed behavior

Commonsense and theorizing about action control agree in assuming that human behavior is (mainly) driven by goals, but no mechanistic theory of what goals are, where they come from, and how they impact action selection is available. Here I develop such a theory that is based on the assumption that GOALs guide Intentional Actions THrough criteria (GOALIATH). The theory is intended to be minimalist and parsimonious with respect to its assumptions, as transparent and mechanistic as possible, and it is based on representational assumptions provided by the Theory of Event Coding (TEC). It holds that goal-directed behavior is guided by selection criteria that activate and create competition between event files that contain action-effect codes matching one or more of the criteria—a competition that eventually settles into a solution favoring the best-matching event file. The criteria are associated with various sources, including biological drives, acquired needs (e.g., of achievement, power, or affiliation), and short-term, sometimes arbitrary, instructed aims. Action selection is, thus, a compromise that tries to satisfy various criteria related to different driving forces, which are also likely to vary in strength over time. Hence, what looks like goal-directed action emerges from, and represents an attempt to satisfy multiple constraints with different origins, purposes, operational characteristics, and timescales—which among other things does not guarantee a high degree of coherence or rationality of the eventual outcome. GOALIATH calls for a radical break with conventional theorizing about the control of goal-directed behavior, as it among other things questions existing cognitive-control theories and dual-route models of action control.

How children with neurodevelopmental disorders can benefit from the neurocognitive effects of exercise

Exercise is an integral part of children’s lives, and research in educational settings has found that regular engagement promises improvements in executive function (i.e. top-down control of goal-directed behavior). Based on recent advances in understanding the moderators and the neurocognitive mechanisms of these effects, we highlight a potential application of exercise in the treatment of executive dysfunction. Even though different neurodevelopmental disorders are characterized by a heterogeneity in core symptoms, children affected by these disorders often face common executive function deficits. So far, exercise has not been recognized as an alternative or additional treatment for this specific cognitive impairment. The limited experimental evidence in children with neurodevelopmental disorders provides a first indication that regular exercise engagement benefits executive function. However, we identified key research questions that need to be answered before a prescription of exercise to children with executive dysfunction can be encouraged in clinical practice.

Association between alexithymia and impaired reward valuation in patients with fronto-insular damage.

Humans compute the anticipated reward value of stimuli in their environment in order to behave in an adaptive, goal-directed manner. This reward valuation ability is vital, and its disruption in a range of clinical populations has profound personal and social consequences. However, research has often failed to consider the reward-related functions of a central component of human emotion: conscious emotional experience. Alexithymia-a condition characterized by diminished conscious awareness of one's emotions-offers a unique opportunity to examine the link between emotional awareness and reward valuation. In the present study, we measured both acquired alexithymia and reward valuation ability in a large sample of patients with traumatic brain injuries (N = 112). Behavioral analyses provided evidence for a negative association between alexithymia and reward valuation ability. This association remained significant after controlling for several covariates in the model (anxiety, depression, posttraumatic stress disorder, and IQ). Voxel-based lesion-symptom mapping was carried out to identify brain regions-of-interest (ROIs) that, when damaged, lead to increased alexithymia and impaired reward valuation. Importantly, mediation models computed using the ROIs identified through the voxel-based lesion-symptom mapping revealed a specific indirect effect of left frontoinsular damage on impaired valuation that was mediated by increased levels of alexithymia. This indirect effect was not observed for any of the other candidate ROIs. The present study identifies a network of brain regions likely to be involved in the integration of subjective feelings and reward processes critical for the adaptive control of goal-directed behavior. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

An experimental test of the habit-goal framework: Depressive rumination is associated with heightened habitual characteristics of negative thinking but not habit-directed behavior control

Habitual thinking may underpin a heightened disposition to engage in rumination in response to negative mood, a widely held notion that has rarely been directly tested. The purpose of the current study was to investigate whether rumination is associated with habitual attributes and whether it is related to an imbalance in habit relative to goal-directed behavior control. University students (N=115) completed self-report questionnaires, a rumination induction paradigm and an outcome devaluation task that measures habitual vs goal-directed behavior control. Greater habitual characteristics of negative thinking (e.g., automaticity, lack of conscious awareness, control, and intent) were associated with ruminative brooding but not ruminative reflection and predicted more persistent dysphoric mood following rumination induction. Rumination was not, however, consistently associated with an imbalance in habit versus goal-directed behavior control. These findings indicate that depression vulnerability may be in the form of rumination being habitually triggered (without awareness or intent) with deleterious effects on mood. Although habitual, rumination may not be related to an imbalance in habit relative to goal-directed behavior control. These findings provide support for current theoretical accounts of rumination and set important boundary conditions in the search for specific factors that contribute to rumination as a habit.

NEUROSCIENCE OF MOTOR LEARNING IN ADULTS WITH NON-DOMINANT HAND

Hand dominance represents an evident example of hemispheric specialization. Though the vast majority of the population are right-handed with dominant left hemisphere, some adults have a non-dominant hand and they can use both hands equally. Evidence from neurophysiological studies established that the left dominant cerebral hemisphere integrates the motor skills the dominant hand including the precision, speed, and co-ordination of motor acts. The non-dominant hemisphere is specialized in the perception and control of goal-directed behaviour and visuospatial processing related the performed motor task. Lesions in dominant cerebral hemispheres can, thus, result in considerable compromise of the motor skills. Learning new motor skills is essential in these cases, and the capability of the cerebral hemispheres to learn new tasks has long been studied and debated. To evaluate the potential motor learning capacity of cerebral hemispheres, neuroscientists have studied the motor learning in adults with non-dominant hands. Those adults exemplify a model for testing the impact of practice and learning on the brain acquisition of new motor tasks. This article aims at reviewing the recent neuroscientific and psychological evidence about the motor learning among adults with non-motor hand.

Striatopallidal adenosine A2A receptors in the nucleus accumbens confer motivational control of goal-directed behavior

The ability to learn the reward-value and action-outcome contingencies in dynamic environment is critical for flexible adaptive behavior and development of effective pharmacological control of goal-directed behaviors represents an important challenge for improving the deficits in goal-directed behavior which may underlie seemingly disparate symptoms across psychiatric disorders. Adenosine A2A receptor (A2AR) is emerging as a novel neuromodulatory target for controlling goal-directed behavior for its unique neuromodulatory features: the ability to integrate dopamine and glutamate signaling, the “brake” constraint of various cognitive processes and the balanced control of goal-directed and habit actions. However, the contribution and circuit mechanisms of the striatopallidal A2ARs in nucleus accumbens (NAc) to control of goal-directed behavior remain to be determined. Here, we employed newly developed opto-A2AR and the focal A2AR knockdown strategies to demonstrate the causal role of NAc A2AR in control of goal-directed behavior. Furthermore, we dissected out multiple distinct behavioral mechanisms underlying which NAc A2ARs control goal-directed behavior: (i) NAc A2ARs preferentially control goal-directed behavior at the expense of habit formation. (ii) NAc A2ARs modify the animals’ sensitivity to the value of the reward without affecting the action-outcome contingency. (iii) A2AR antagonist KW6002 promotes instrumental actions by invigorating motivation. (iv) NAc A2ARs facilitate Pavlovian incentive value transferring to instrumental action. (v) NAc A2ARs control goal-directed behavior probably not through NAc-VP pathway. These insights into the behavioral and circuit mechanisms for NAc A2AR control of goal-directed behavior facilitate translational potential for A2AR antagonists in reversal of deficits in goal-directed decision-making associated with multiple neuropsychiatric disorders.

Functional network topology associated with apathy in Alzheimer's disease

BackgroundApathy, a common neuropsychiatric (NPS) in patients with mild cognitive impairment (MCI) and Alzheimer's disease (AD), is associated with structural and metabolic brain changes. However, functional connectivity changes across the brain in association with apathy remain unclear. In this study, graph theoretical measures of integration and segregation from resting state functional connectivity in MCI and AD patients with low depression scores, and healthy controls. MethodsIn MCI and AD patients with low depression scores, graph theoretical measures of integration and segregation were derived from resting state functional connectivity in patients, which were compared between those with apathy (NPS_A, n = 21) to those without NPS (NPS_None, n = 28) and those with NPS other than apathy (NPS_NA, n = 38). Additionally, the same measures were compared between AD patients and healthy controls (amyloid uptake below threshold levels). ResultsAltered whole brain global efficiency and reduced local efficiency were found in NPS_A compared to NPS_None and NPS_NA. In similar contrasts, apathy was associated with increased participation coefficient in the frontoparietal and cingulo-opercular template-based networks. A study-specific network definition also showed similar results. In comparison, AD patients showed higher modularity compared to controls at the whole brain level and higher participation coefficient in the ventral attention network. LimitationsThe severity and dimensions of apathy were not assessed. ConclusionsLoss of segregation in the frontoparietal and cingulo-opercular network, which are involved in the control of goal-directed behavior, was associated with apathy in MCI/AD. The results also suggest that network-level changes in AD patients may underlie specific NPS.

Breaking the limits of executive functions: Towards a sociocultural perspective

Executive functions refer to the neurocognitive processes that enable conscious control of goal-directed behaviour. Executive functions lay the neural and cognitive foundation for civilisation and culture with its far reaching effects in day-to-day planning, problem-solving, creativity, self-regulation, empathy and cooperative social behaviour. Though a celebrated area of research for cognitive scientists, contemporary studies find that most of the interventions that target executive functions promote academic achievement in children, but fail to transfer to real-life interpersonal situations. This suggests the need for breaking its cognitive shell and conceiving executive function development in the larger sociocultural context. Thus, the present paper inquires into the cultural space that provides collaborative learning experiences for young children to become efficient problem-solvers and empathetic social beings. After reviewing certain theoretical perspectives and research findings, it is concluded that the early experiences of scaffolding by more competent individuals, specific to the social context and culture, are the crucial determinant of positive executive function development and its expression in everyday situations. The process of such sociocultural influences seems to be mediated by learning appropriate rules that guide executive functions. The paper puts forth the need for culture-specific value-based training for children that can potentially improve the internally motivated execution of control in practical situations. Revival of activities inherent in cultures is also found to be a promising option for devising effective training methods.

Delayed rewards facilitate habit formation.

The transition from goal-directed to habitual forms of instrumental behavior is determined by variables such as the amount of training, schedules of reinforcement, the availability of choices, and exposure to drugs of abuse. Less is known about the control of goal-directed behavior when reinforcement is delayed rather than immediate. In these experiments, we investigated in rats the role of response-outcome contiguity on the control of goal-directed action, assessed through satiety-specific outcome devaluation tests. In Experiment 1 using a within-subjects design we observed goal-directed behavior after 6 days of FR1 training when the outcome was presented immediately following the lever press, but not when it was delayed for 20 s, revealing habit formation with delayed outcomes. Experiment 2 revealed that the habitual control observed with 20-s delays of reinforcement can be prevented if, immediately before each instrumental training session, the rats were exposed to the experimental context in the absence of both the lever and reinforcement. In summary, these experiments suggest that response-outcome contiguity plays an important role in the control of goal-directed actions and habits. (PsycINFO Database Record (c) 2019 APA, all rights reserved).