The creativity of architects.

TA builds on the state of mind (SoM) framework to offer the novelty-seeking model (NSM). The model relates curiosity to creativity but this commentary focuses on creativity: (i) It assesses the SoM + NSM model of creativity-in-the-lab, showing that the focus on semantic networks is inadequate. (ii) It discusses architectural design to sketch ideas for a theory of "big C" creativity.

From spatial navigation via visual construction to episodic memory and imagination.

This hybrid of review and personal essay argues that models of visual construction are essential to extend spatial navigation models to models that link episodic memory and imagination. The starting point is the TAM-WG model, combining the Taxon Affordance Model and the World Graph model of spatial navigation. The key here is to reject approaches in which memory is restricted to unanalyzed views from familiar places, and their later recall. Instead, we will seek mechanisms for imagining truly novel scenes and episodes. We thus introduce a specific variant of schema theory and VISIONS, a cooperative computation model of visual scene understanding in which a scene is represented by an assemblage of schema instances with links to lower-level "patches" of relevant visual data. We sketch a new conceptual framework for future modeling, Visual Integration of Diverse Multi-Modal Aspects, by extending VISIONS from static scenes to episodes combining agents, actions and objects and assess its relevance to both navigation and episodic memory. We can then analyze imagination as a constructive process that combines aspects of memories of prior episodes along with other schemas and adjusts them into a coherent whole which, through expectations associated with diverse episodes and schemas, may yield the linkage of episodes that constitutes a dream or a narrative. The result is IBSEN, a conceptual model of Imagination in Brain Systems for Episodes and Navigation. The essay closes by analyzing other papers in this Special Issue to assess to what extent their results relate to the research proposed here.

The super-learning hypothesis: Integrating learning processes across cortex, cerebellum and basal ganglia.

Despite wide evidence suggesting anatomical and functional interactions between cortex, cerebellum and basal ganglia, the learning processes operating within them --often viewed as respectively unsupervised, supervised and reinforcement learning-- are studied in isolation, neglecting their strong interdependence. We discuss how those brain areas form a highly integrated system combining different learning mechanisms into an effective super-learning process supporting the acquisition of flexible motor behaviour. The term "super-learning" does not indicate a new learning paradigm. Rather, it refers to the fact that different learning mechanisms act in synergy as they: (a) affect neural structures often relying on the widespread action of neuromodulators; (b) act within various stages of cortical/subcortical pathways that are organised in pipeline to support multiple sensation-to-action mappings operating at different levels of abstraction; (c) interact through the reciprocal influence of the output compartments of different brain structures, most notably in the cerebello-cortical and basal ganglia-cortical loops. Here we articulate this new hypothesis and discuss empirical evidence supporting it by specifically referring to motor adaptation and sequence learning.

Dysfunctions of the basal ganglia-cerebellar-thalamo-cortical system produce motor tics in Tourette syndrome.

Motor tics are a cardinal feature of Tourette syndrome and are traditionally associated with an excess of striatal dopamine in the basal ganglia. Recent evidence increasingly supports a more articulated view where cerebellum and cortex, working closely in concert with basal ganglia, are also involved in tic production. Building on such evidence, this article proposes a computational model of the basal ganglia-cerebellar-thalamo-cortical system to study how motor tics are generated in Tourette syndrome. In particular, the model: (i) reproduces the main results of recent experiments about the involvement of the basal ganglia-cerebellar-thalamo-cortical system in tic generation; (ii) suggests an explanation of the system-level mechanisms underlying motor tic production: in this respect, the model predicts that the interplay between dopaminergic signal and cortical activity contributes to triggering the tic event and that the recently discovered basal ganglia-cerebellar anatomical pathway may support the involvement of the cerebellum in tic production; (iii) furnishes predictions on the amount of tics generated when striatal dopamine increases and when the cortex is externally stimulated. These predictions could be important in identifying new brain target areas for future therapies. Finally, the model represents the first computational attempt to study the role of the recently discovered basal ganglia-cerebellar anatomical links. Studying this non-cortex-mediated basal ganglia-cerebellar interaction could radically change our perspective about how these areas interact with each other and with the cortex. Overall, the model also shows the utility of casting Tourette syndrome within a system-level perspective rather than viewing it as related to the dysfunction of a single brain area.

Toward the Language-Ready Brain: Biological Evolution and Primate Comparisons.

The approach to language evolution suggested here focuses on three questions: How did the human brain evolve so that humans can develop, use, and acquire languages? How can the evolutionary quest be informed by studying brain, behavior, and social interaction in monkeys, apes, and humans? How can computational modeling advance these studies? I hypothesize that the brain is language ready in that the earliest humans had protolanguages but not languages (i.e., communication systems endowed with rich and open-ended lexicons and grammars supporting a compositional semantics), and that it took cultural evolution to yield societies (a cultural constructed niche) in which language-ready brains could become language-using brains. The mirror system hypothesis is a well-developed example of this approach, but I offer it here not as a closed theory but as an evolving framework for the development and analysis of conflicting subhypotheses in the hope of their eventual integration. I also stress that computational modeling helps us understand the evolving role of mirror neurons, not in and of themselves, but only in their interaction with systems "beyond the mirror." Because a theory of evolution needs a clear characterization of what it is that evolved, I also outline ideas for research in neurolinguistics to complement studies of the evolution of the language-ready brain. A clear challenge is to go beyond models of speech comprehension to include sign language and models of production, and to link language to visuomotor interaction with the physical and social world.

Towards a Computational Comparative Neuroprimatology: Framing the language-ready brain.

We make the case for developing a Computational Comparative Neuroprimatology to inform the analysis of the function and evolution of the human brain. First, we update the mirror system hypothesis on the evolution of the language-ready brain by (i) modeling action and action recognition and opportunistic scheduling of macaque brains to hypothesize the nature of the last common ancestor of macaque and human (LCA-m); and then we (ii) introduce dynamic brain modeling to show how apes could acquire gesture through ontogenetic ritualization, hypothesizing the nature of evolution from LCA-m to the last common ancestor of chimpanzee and human (LCA-c). We then (iii) hypothesize the role of imitation, pantomime, protosign and protospeech in biological and cultural evolution from LCA-c to Homo sapiens with a language-ready brain. Second, we suggest how cultural evolution in Homo sapiens led from protolanguages to full languages with grammar and compositional semantics. Third, we assess the similarities and differences between the dorsal and ventral streams in audition and vision as the basis for presenting and comparing two models of language processing in the human brain: A model of (i) the auditory dorsal and ventral streams in sentence comprehension; and (ii) the visual dorsal and ventral streams in defining "what language is about" in both production and perception of utterances related to visual scenes provide the basis for (iii) a first step towards a synthesis and a look at challenges for further research.

Learning to grasp and extract affordances: the Integrated Learning of Grasps and Affordances (ILGA) model.

The activity of certain parietal neurons has been interpreted as encoding affordances (directly perceivable opportunities) for grasping. Separate computational models have been developed for infant grasp learning and affordance learning, but no single model has yet combined these processes in a neurobiologically plausible way. We present the Integrated Learning of Grasps and Affordances (ILGA) model that simultaneously learns grasp affordances from visual object features and motor parameters for planning grasps using trial-and-error reinforcement learning. As in the Infant Learning to Grasp Model, we model a stage of infant development prior to the onset of sophisticated visual processing of hand-object relations, but we assume that certain premotor neurons activate neural populations in primary motor cortex that synergistically control different combinations of fingers. The ILGA model is able to extract affordance representations from visual object features, learn motor parameters for generating stable grasps, and generalize its learned representations to novel objects.

Co-evolution of human consciousness and language (revisited).

This article discusses the view that human consciousness may share aspects of "animal awareness" with other species, but has its unique form because humans possess language. Two ingredients of a theory of the evolution of human consciousness are offered: the view that a précis of intended activity is necessarily formed in the brain of a human that communicates in a human way; and the notion that such a précis underwrites the uniquely human aspect of consciousness.

Ontogenetic ritualization of primate gesture as a case study in dyadic brain modeling.

This paper introduces dyadic brain modeling - the simultaneous, computational modeling of the brains of two interacting agents - to explore ways in which our understanding of macaque brain circuitry can ground new models of brain mechanisms involved in ape interaction. Specifically, we assess a range of data on gestural communication of great apes as the basis for developing an account of the interactions of two primates engaged in ontogenetic ritualization, a proposed learning mechanism through which a functional action may become a communicative gesture over repeated interactions between two individuals (the 'dyad'). The integration of behavioral, neural, and computational data in dyadic (or, more generally, social) brain modeling has broad application to comparative and evolutionary questions, particularly for the evolutionary origins of cognition and language in the human lineage. We relate this work to the neuroinformatics challenges of integrating and sharing data to support collaboration between primatologists, neuroscientists and modelers that will help speed the emergence of what may be called comparative neuro-primatology.

A neuroinformatics of brain modeling and its implementation in the Brain Operation Database BODB.

We present principles for an integrated neuroinformatics framework which makes explicit how models are grounded on empirical evidence, explain (or not) existing empirical results and make testable predictions. The new ontological framework makes explicit how models bring together structural, functional, and related empirical observations. We emphasize schematics of the model's operation linked to summaries of empirical data (SEDs) used in both the design and testing of the model, with tests comparing SEDs to summaries of simulation results (SSRs) from the model. We stress the importance of protocols for models as well as experiments. We complement the structural ontology of nested brain structures with a functional ontology of Brain Operating Principles (BOPs) for observed neural function and an ontological framework for grounding models in empirical data. We present an implementation of this ontological framework in the Brain Operation Database (BODB), an environment in which modelers and experimentalists can work together by making use of their shared empirical data, models and expertise.

Action and language mechanisms in the brain: data, models and neuroinformatics.

We assess the challenges of studying action and language mechanisms in the brain, both singly and in relation to each other to provide a novel perspective on neuroinformatics, integrating the development of databases for encoding ­ separately or together ­ neurocomputational models and empirical data that serve systems and cognitive neuroscience.

Language is handy but is it embodied?

Part 1 provides Arbib's reflections on the influence of Marc Jeannerod on his career. Part 2 recalls the Mirror System Hypothesis (MSH) for the evolution of the language-ready brain, a theory which emphasizes the role of manual action in grounding language evolution, thus giving one meaning for "language is handy". Part 3 then joins in current debates over the notion of whether or not language is embodied. Our overall argument is that embodiment is a graded rather than binary concept, and that embodiment provides the evolutionary and developmental core of concepts and language, but that the modern human brain supports abstraction processes that make embodiment little relevant in a wide range of language use. We urge that, rather than debate the extent of embodiment, attention should turn to the integration of empirical studies with computational modeling to delineate in detail processes of abstraction, generalization, metaphor and more, bridging between modeling of neural mechanisms in macaque that may be posited for the brain of the last monkey-human common ancestor (LCA-m) and computational modeling of human language processing. Part 4 suggests that variants of construction grammar are well-suited to the latter task.

Modeling the BOLD correlates of competitive neural dynamics.

Winner-take-all models are commonly used to model decision-making tasks where one outcome must be selected from several competing options. Related random walk and diffusion models have been used to explain such processes and apply them to psychometric and neurophysiological data. Recent model-based fMRI studies have sought to find the neural correlates of decision-making processes. However, due to the fact that hemodynamic responses likely reflect synaptic rather than spiking activity, the expected BOLD signature of winner-take-all circuits is not clear. A powerful way to integrate data from neurophysiology and brain imaging is by developing biologically plausible neural network models constrained and testable by neural and behavioral data, and then using Synthetic Brain Imaging - transforming the output of simulations with the model to make predictions testable against neuroimaging data. We developed a biologically realistic spiking winner-take-all model comprised of coupled excitatory and inhibitory neural populations. We varied the difficulty of a decision-making task by adjusting the contrast, or relative strength of inputs representing two response options. Synthetic brain imaging was used to estimate the BOLD response of the model and analyze its peak as a function of input contrast. We performed a parameter space analysis to determine values for which the model performs the task accurately, and given accurate performance, the distribution of the input contrast-BOLD response relationship. This underscores the need for models grounded in neurophysiological data for brain imaging analyses which attempt to localize the neural correlates of cognitive processes based on predicted BOLD responses.

Recognizing speech in a novel accent: the motor theory of speech perception reframed.

The motor theory of speech perception holds that we perceive the speech of another in terms of a motor representation of that speech. However, when we have learned to recognize a foreign accent, it seems plausible that recognition of a word rarely involves reconstruction of the speech gestures of the speaker rather than the listener. To better assess the motor theory and this observation, we proceed in three stages. Part 1 places the motor theory of speech perception in a larger framework based on our earlier models of the adaptive formation of mirror neurons for grasping, and for viewing extensions of that mirror system as part of a larger system for neuro-linguistic processing, augmented by the present consideration of recognizing speech in a novel accent. Part 2 then offers a novel computational model of how a listener comes to understand the speech of someone speaking the listener's native language with a foreign accent. The core tenet of the model is that the listener uses hypotheses about the word the speaker is currently uttering to update probabilities linking the sound produced by the speaker to phonemes in the native language repertoire of the listener. This, on average, improves the recognition of later words. This model is neutral regarding the nature of the representations it uses (motor vs. auditory). It serve as a reference point for the discussion in Part 3, which proposes a dual-stream neuro-linguistic architecture to revisits claims for and against the motor theory of speech perception and the relevance of mirror neurons, and extracts some implications for the reframing of the motor theory.

Mirror neurons: functions, mechanisms and models.

Mirror neurons for manipulation fire both when the animal manipulates an object in a specific way and when it sees another animal (or the experimenter) perform an action that is more or less similar. Such neurons were originally found in macaque monkeys, in the ventral premotor cortex, area F5 and later also in the inferior parietal lobule. Recent neuroimaging data indicate that the adult human brain is endowed with a "mirror neuron system," putatively containing mirror neurons and other neurons, for matching the observation and execution of actions. Mirror neurons may serve action recognition in monkeys as well as humans, whereas their putative role in imitation and language may be realized in human but not in monkey. This article shows the important role of computational models in providing sufficient and causal explanations for the observed phenomena involving mirror systems and the learning processes which form them, and underlines the need for additional circuitry to lift up the monkey mirror neuron circuit to sustain the posited cognitive functions attributed to the human mirror neuron system.

Tool use and constructions.

We examine tool use in relation to the capacity of animals for construction, contrasting tools and nests; place human tool use in a more general problem-solving context, revisiting the body schema in the process; and relate the evolution of language and of tool use.