Morality and the Brain: The Right Hemisphere and Doing Right.

Morality, the set of shared attitudes and practices that regulate individual behavior to facilitate cohesion and well-being, is a function of the brain, yet its localization is uncertain. Neuroscientific study of morality has been conducted by examining departures from moral conduct after neurologic insult and by functional neuroimaging of moral decision-making in cognitively intact individuals. These investigations have yielded conflicting results: Acquired sociopathy, a syndromic surrogate for acquired immorality, has been reported predominantly after right frontotemporal lesions, whereas functional neuroimaging during moral decision-making has demonstrated bilateral activation. Although morality is bilaterally represented, the right hemisphere is clinically more critical in light of focal lesion data suggesting that moral behavior is subserved by a network of right frontotemporal structures and their subcortical connections. Evolution may have endowed the brain with bilaterally represented but unilaterally right-dominant morality. The unilateral dominance of morality permits concentration of an essential social cognitive function to support the perceptual and executive operations of moral behavior within a single hemisphere; the bilateral representation of morality allows activation of reserve tissue in the contralateral hemisphere in the event of an acquired hemispheric injury. The observed preponderance of right hemisphere lesions in individuals with acquired immorality offers a plausible hypothesis that can be tested in clinical settings. Advances in the neuroscience of morality promise to yield potentially transformative clinical and societal benefits. A deeper understanding of morality would help clinicians address disordered conduct after acquired neurologic insults and guide society in bolstering public health efforts to prevent brain disease.

The First Neuroethics Meeting: Then and Now.

It wasn't until 2002 that more than 150 neuroscientists, bioethicists, doctors of psychiatry and psychology, philosophers, and professors of law and public policy came together to chart the boundaries, define the issues, and raise some of the ethical implications tied to advances in brain research. On the 15 th anniversary of the Neuroethics: Mapping the Field conference in San Francisco, we asked three of the original speakers to reflect on how far the neuroethics field has come in 15 years-and where the field may be going in the next 15.

Blending computational and experimental neuroscience.

The launch of the United States' BRAIN Initiative brings with it a new era in systems neuroscience that is being driven by innovative neurotechnologies, increases in computational power and network-style artificial intelligence. A new conceptual framework for understanding cognitive behaviours based on the dynamical patterns of activity in large populations of neurons is emerging.

Brains, genes, and primates.

One of the great strengths of the mouse model is the wide array of genetic tools that have been developed. Striking examples include methods for directed modification of the genome, and for regulated expression or inactivation of genes. Within neuroscience, it is now routine to express reporter genes, neuronal activity indicators, and opsins in specific neuronal types in the mouse. However, there are considerable anatomical, physiological, cognitive, and behavioral differences between the mouse and the human that, in some areas of inquiry, limit the degree to which insights derived from the mouse can be applied to understanding human neurobiology. Several recent advances have now brought into reach the goal of applying these tools to understanding the primate brain. Here we describe these advances, consider their potential to advance our understanding of the human brain and brain disorders, discuss bioethical considerations, and describe what will be needed to move forward.

Putting big data to good use in neuroscience.

Big data has transformed fields such as physics and genomics. Neuroscience is set to collect its own big data sets, but to exploit its full potential, there need to be ways to standardize, integrate and synthesize diverse types of data from different levels of analysis and across species. This will require a cultural shift in sharing data across labs, as well as to a central role for theorists in neuroscience research.

Exploring the causal underpinnings of determination, resolve, and will.

In this issue of Neuron, Parvizi et al. (2013) show that mild electrical stimulation using depth electrodes in the brains in human patients reliably elicits a highly specific configuration of cognitive-emotional-motivational responses to persevere in the face of danger. The underlying mechanisms involve distributed networks, both cortical and subcortical.

Problems with measuring peripheral oxytocin: can the data on oxytocin and human behavior be trusted?

Research on the neurobiological and behavioral effects of oxytocin (OT), as well as on its possible therapeutic applications, has intensified in the past decade. Accurate determination of peripheral OT levels is essential to reach meaningful conclusions and to motivate, support and inform clinical interventions. Different, but concordant, methods for measuring plasma OT have been developed over the past four decades, but since 2004 several commercially available methods have been favored in research with humans. Evaluation of these methods reveals that they lack reliability when used on unextracted samples of human fluids, and that they tag molecules in addition to OT, yielding estimates that are wildly discrepant with an extensive body of earlier findings that were obtained using methods that are well validated, but more laborious. An accurate, specific, and readily available method for measuring OT that can be adopted as the standard in the field is urgently needed for advances in our understanding of OT's roles in cognition and behavior.

Modulating social behavior with oxytocin: how does it work? What does it mean?

Among its many roles in body and brain, oxytocin influences social behavior. Understanding the precise nature of this influence is crucial, both within the broader theoretical context of neurobiology, social neuroscience and brain evolution, but also within a clinical context of disorders such as anxiety, schizophrenia, and autism. Research exploring oxytocin's role in human social behavior is difficult owing to its release in both body and brain and its interactive effects with other hormones and neuromodulators. Additional difficulties are due to the intricacies of the blood-brain barrier and oxytocin's instability, which creates measurement issues. Questions concerning how to interpret behavioral results of human experiments manipulating oxytocin are thus made all the more pressing. The current paper discusses several such questions. We highlight unresolved fundamental issues about what exactly happens when oxytocin is administered intranasally, whether such oxytocin does in fact reach appropriate receptors in brain, and whether central or peripheral influences account for the observed behavioral effects. We also highlight the deeper conceptual issue of whether the human data should be narrowly interpreted as implicating a specific role for oxytocin in complex social cognition, such a generosity, trust, or mentalizing, or more broadly interpreted as implicating a lower-level general effect on general states and dispositions, such as anxiety and social motivation. Using several influential studies, we show how seemingly specific, higher-level social-cognitive effects can emerge via a process by which oxytocin's broad influence is channeled into a specific social behavior in a context of an appropriate social and research setting. This article is part of a Special Issue entitled Oxytocin, Vasopressin, and Social Behavior.

Can Innate, modular "foundations" explain morality? Challenges for Haidt's Moral Foundations Theory.

Jonathan Haidt's Moral Foundations Theory is an influential scientific account of morality incorporating psychological, developmental, and evolutionary perspectives. The theory proposes that morality is built upon five innate "foundations," each of which is believed to have been selected for during human evolution and, subsequently, tuned-up by learning during development. We argue here that although some general elements of Haidt's theory are plausible, many other important aspects of his account are seriously flawed. First, innateness and modularity figure centrally in Haidt's account, but terminological and conceptual problems foster confusion and ambiguities. Second, both the theory's proposed number of moral foundations and its taxonomy of the moral domain appear contrived, ignoring equally good candidate foundations and the possibility of substantial intergroup differences in the foundations' contents. Third, the mechanisms (viz., modules) and categorical distinctions (viz., between foundations) proposed by the theory are not consilient with discoveries in contemporary neuroscience concerning the organization, functioning, and development of the brain. In light of these difficulties, we suggest that Haidt's theory is inadequate as a scientific account of morality. Nevertheless, the theory's weaknesses are instructive, and hence, criticism may be useful to psychologists, neuroscientists, and philosophers attempting to advance theories of morality, as well as to researchers wishing to invoke concepts such as innateness and modularity more generally.

Control: conscious and otherwise.

Social psychologists have shown human decisions to be sensitive to numerous ordinary, possibly nonconscious, situational contingencies, motivating the view that control is largely illusory, and that our choices are largely governed by such external contingencies. Against this view is evidence that self-control and goal-maintenance are regularly displayed by humans and other animals, and evidence concerning neurobiological processes that support such control. Evolutionarily speaking, animals with a robust capacity to exercise control - both conscious and nonconscious - probably enjoyed a selective advantage. Counterbalancing data thus point to an account of control that sees an important role for nonconscious control in action and goal maintenance. We propose a conceptual model of control that encompasses such nonconscious control and links in-control behavior to neurobiological parameters.

The impact of neuroscience on philosophy.

In the last two decades, neuroscience has profoundly transformed how we understand learning, decision making, self, and social attachment. Consequently, traditional philosophical questions about mind and morality have been steered in new directions.

The significance of neuroscience for philosophy.

The ground is shifting under the traditional approaches to problems in the philosophy of mind. Earlier doctrines concerning the independence of cognition from the brain now appear untenable. As neuroscience uncovers more about the organization and dynamics of the brain, it becomes increasingly evident that theories about our nature must be informed by neuroscientific data. Consistent with this progress, we may expect that philosophical problems about the mind will be productively addressed and perhaps radically transformed by a convergence of neuroscientific, psychological and computational research.

Neurophilosophy: the early years and new directions.

Neurophilosophy embraces the hypothesis that what we call "the mind" is in fact a level of brain activity. A corollary of this hypothesis states that we can learn much about the reality of mental function by studying the brain at all levels of organization. Until fairly recently, many philosophers preferred to believe that important domains of mental function could never be addressed using the tools of empirical science. Nevertheless, co-evolutionary progress by psychology and the neurosciences on many topics, including consciousness, free will and the nature of knowledge, have meant that such convictions need to be updated. Some large-scale mind-brain problems have not yet been solved, and do require significant theoretical innovation. In particular the problem of how to understand the true nature of representations remains unsolved.