Understanding complexity in the
human brain
Danielle S. Bassett1 and Michael S. Gazzaniga2
Complex Systems Group, Department of Physics, University of California, Santa Barbara, CA 93106, USA
Sage Center for the Study of the Mind, University of California, Santa Barbara, CA 93106, USA

Although the ultimate aim of neuroscientific enquiry is phenomena [3] that together form the physical and biolog-
to gain an understanding of the brain and how its work- ical basis of cognition. Furthermore, the structure within
ings relate to the mind, the majority of current efforts any given scale is organized into modules – for example
are largely focused on small questions using increasing- anatomically or functionally defined cortical regions – that
ly detailed data. However, it might be possible to suc- form the basis of cognitive functions that are optimally
cessfully address the larger question of mind–brain adaptable to perturbations in the external environment.
mechanisms if the cumulative findings from these neu-
roscientific studies are coupled with complementary Spatial and temporal scaling
approaches from physics and philosophy. The brain, In the spatial domain alone, the brain displays similar
we argue, can be understood as a complex system or organization at multiple resolutions (Figure 2). In addition
network, in which mental states emerge from the inter- to the spatial distributions of molecules inside neuronal
action between multiple physical and functional levels. and non-neuronal cells, the cells themselves are heteroge-
Achieving further conceptual progress will crucially de- neously distributed throughout the brain [5]. Minicolumns
pend on broad-scale discussions regarding the proper- - vertical columns through the cortical layers of the brain -
ties of cognition and the tools that are currently contain on average 100 neurons, are roughly 30 microns
available or must be developed in order to study in diameter and form the anatomical basis of columns,
mind–brain mechanisms. subareas (e.g. V1), areas (e.g. visual cortex), lobes (e.g. the
occipital lobe), and thereby the complete cerebral cortex.
The state of the mind–brain dilemma Not only is the structural distribution of components
The human mind is a complex phenomenon built on the
physical scaffolding of the brain [1–3], which neuroscien-
tific investigation continues to examine in great detail.
However, the nature of the relationship between the mind Complex network theory: a modeling framework that defines a complex
system in terms of its subcomponents and their interactions, which together
and the brain is far from understood [4]. In this article we form a network.
argue that recent advances in complex systems theory (see Complex system: a system whose overall behavior can be characterized as
Glossary) might provide crucial new insights into this more than the sum of its parts.
Connectome: a complete connectivity map of a system. In neuroscience, the
problem. We first examine what is presently known about structural connectome is defined by the anatomical connections between
the complexity of the brain and review recent applications subunits of the brain whereas the functional connectome is defined by the
functional relations between those subunits.
of complex network theory to the study of brain connectivi-
Emergence: the manner in which complex phenomena arise from a collection
ty [2,3] (Box 1 and Figure 1). We then discuss the philo- of relatively simple interactions between system components.
sophical concept of emergence as a potential framework for Ising model: a historically important mathematical model of a phenomenon in
physics known as ferromagnetism that displays several characteristics of
the investigation of mind–brain mechanisms. We delineate complex systems including phase transitions and the emergence of collective
currently available investigative tools for the examination phenomena.
of this problem, from quantitative statistical physics to Modularity: a property of a system that can be decomposed into subcompo-
nents or ‘modules’, which can perform unique functions and can adapt or
qualitative metaphors, and discuss their relative advan- evolve to external demands.
tages and limitations. Finally, we highlight crucial areas Nonfundamental causality: the concept that parts of a system that are not its
where further work is necessary to achieve progress, in- smallest parts (i.e. nonfundamental) can have significant causal power in terms
of system function, facilitating mutual manipulability between multiple levels
cluding both detailed modeling and large-scale theoretical of the system and multiple realizability of system function.
frameworks. Reductionism: in philosophy, a view that a complex system can be modeled
and understood simply by reducing the examination to a study of the system’s
constituent parts.
Complexity and multiscale organization Scaling: the term scaling indicates a similarity of some organizational structure
A first step in understanding mind–brain mechanisms is to of phenomenon across multiple scales of a system. Spatial scaling therefore
characterize what is known about the structure of the brain indicates that a principle or phenomenon is consistently displayed at multiple
spatial resolutions. Temporal scaling indicates that a principle or phenomenon
and its organizing principles. The brain is a complex is consistently displayed at multiple temporal resolutions.
temporally and spatially multiscale structure that Wiring diagram: a map of the hard-wired connectivity of a system. In
neuroscience, the term is usually used to refer to the map of connections
gives rise to elaborate molecular, cellular, and neuronal
(e.g. synapses) between neurons specifically but can also be used to refer to
the map of larger scale white matter connections between brain regions.
Corresponding author: Bassett, D.S. (dbassett@physics.ucsb.edu).

200 1364-6613/$ – see front matter ß 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.tics.2011.03.006 Trends in Cognitive Sciences, May 2011, Vol. 15, No. 5

Vol.Opinion Trends in Cognitive Sciences May 2011. become possible with the recent application of complex portantly. memory whereas in the spatial domain anatomical mod. These results are consistent with recent work character- connectivity maps are supplemented with additional information on izing the physical embedding of brain network organization into the 3- the characteristics of individual components. theta (2– In addition to enhancing robustness and specificity. heterogeneous within the cortex. gamma (4–8 Hz). Complex network theory Complex network theory draws from advances in statistical physics. dorsal attention and sensorimotor networks main. Although a limited conception [14]. gene–gene interactions. Im. such as that present in disease states. time that can be decomposed into subcomponents and the interac- mathematics. fault mode. cific groups of brain regions are more highly correlated ganization both functionally and structurally in both time with one another than to fluctuations of regions in other and space. the system’s connectivity. applied to neuroimaging data. and temporal and spatial scales. the more learning and memory processes (e. tions. connections of modularity of function has influenced neuroscientific between large swaths of cortex can also be altered by thought since the early years of phrenology [27]. modu- and relate to different cognitive capacities. groups of cortical and subcortical brain regions such as the tivity [18. themselves and include the control. functional algorithms and dimensional space of the skull. In addition to applicability. the framework is principled framework in which to examine complex systems that are generalizable across neuroimaging modalities and provides results composed of unique components and display nontrivial component. for example in disease states. supporting the view that One particular experimental approach has recently brain organization dynamically changes over multiple highlighted the modular nature of cortical function. example modules might be short. This framework has been applied to systems previous neuroscientific and theoretical work [79]. food webs. More formal descriptions of modular architecture have ules are present in cortical minicolumns or columns. Combined with the principle of hierarchy.23. (12–30 Hz). as varied as metabolic networks. IQ and working memory of mathematical graph theory to describe the statistical properties of accuracy [30]. graphical properties of human brain networks have been directly social networks and more recently the human brain. systems can be directly related to characteristics of the system’s Complementary avenues of inquiry have uncovered evidence that function and to external constraints that might have shaped the metabolic properties of the brain can be mapped to network system’s growth. longer-term aging are evident.22]. ing in a motor skill such as juggling) as indicated by white non-random. the entire brain system can be decom. structural predictors of altered function. These spatially specific modules have tempo- a nontrivial ‘nearly decomposable’ [20] or modular nature rally dynamic interconnections both within and between (Figure 3). which has 201 . An additional important avenue of inquiry is the construction of done in a cost-efficient manner characteristic of other highly generative models of network organization that can shed light on the constrained physical systems [7]. In summary. 4 Hz) and delta (1–2 Hz) bands each modulate other dis. which can provide important insights into has also been correlated with underlying structure in clinical states as underlying organizational principles. structural [17] and connec. but recent evidence also that are more highly connected to other elements within suggests that so. computer science and the social sciences to provide a tions between them. denced by soft boundaries between coherently operating Furthermore. Graphical models can organization [81] suggesting energetic constraints on underlying be extended to create more complicated models in which simple architecture. Importantly. in fact.g. That is. 15. de- posed into subsystems or modules. nonhomogenous nature of the brain as evi- matter fiber structure measured by diffusion imaging [15]. alpha (8–12 Hz). without adversely perturbing the remainder of the system. In addition to synapses between neurons.a process that seems to have been so on. after long-term train. modularity facilitates behavioral adaptation [21] because tinct yet complementary functions [13]. In the temporal do. thereby providing a compartmentalization that reduces Temporal scaling is evident across the inherent the interdependence of modules and enhances robustness rhythms of brain activity [9–11] that vary in frequency [21.and long-term [29]. The graphical properties of diverse as schizophrenia and Alzheimer’s disease [80]. functional [16]. linked to system function through correlations with behavioral and The simplest application of the theory to these systems is in the use cognitive variables including verbal fluency. that is resting state functional magnetic resonance imag- ing (fMRI) where spontaneous low frequency fluctuations Modularity of the blood-oxygen-level dependent (BOLD) signal in spe- Although this hierarchy of scales characterizes brain or. the pattern of neuronal connections in The principle of modularity. auditory. each of these modules is composed of elements network theory to neuroimaging data [30]. visual.19] signatures of short-term development and motor network and the visual network. can uniquely the human brain . is the connectivity between those the same module than to elements in other modules. Altered function. development and operation.24]. by reducing constraints on the brain changes with learning and memory through the change [21. 5 Box 1. The highest larity allows for a formation of complex architectures frequency gamma band (>30 Hz) is thought to be necessary composed of subsystems within subsystems within sub- for the cognitive binding of information from disparate systems that facilitate a high degree of functional specific- sensory and cognitive modalities [12] whereas the beta ity.a complex system on multiple scales of space and facilitate neuroscientific inquiry. that can be intuitively interpreted in relation to large bodies of to-component relations. No. A broad range of each module can both function and change its function temporal scales also characterizes human cognitive func. For example. contemporary understanding admits the nonarbitrary. therefore forms the structural basis on process of synaptic plasticity on both short (seconds to which subsystems can evolve and adapt [25] in a highly minutes) and long (hours to days to months) timescales variable environment [26]. components [6–8]. organization within a single scale also displays groups [28]. there exists a wealth of emerging evidence that Complex network theory is particularly applicable to the study of complex network theory.

Vol. [()TD$FIG] Increasing spatial resolution Spatial scale 1 2 3 Number of ROIs 180 360 720 Cortical parcellation Hierarchical organization TRENDS in Cognitive Sciences Figure 2. Spatial scaling indicates that an organizational principle characterizes the structure of the human cortex over multiple spatial scales. are represented by edges between those nodes. 15. This structure facilitates global communication and is thought to play a role in the modular organization of connectivity in the cortex [33. No.80]. 5 Brain graph construction Nodes Anatomical connectivity Functional connectivity Brain graph Parietal Occipital Frontal Subcortical Temporal TRENDS in Cognitive Sciences Figure 1. whether anatomical (using diffusion imaging) or functional (using fMRI.()TD$FIG][ Opinion Trends in Cognitive Sciences May 2011. An example of spatial scaling is the mathematical principle of network hierarchy [8]. electroencephalography or magnetoelectroencephalography). brain regions are represented by nodes in a graph and connections between those regions.78]. 202 . In this way a graph can be constructed that characterizes the entire brain system according to its components (nodes) and their relations with one another (edges). Hierarchical network structure is consistently displayed at increasing spatial resolutions in which the brain is parcellated into more and more regions of interest (ROIs).79. the neighbors of a hub are not clustered together. Hierarchical structure is defined as a relation between network nodes whereby hubs (or highly connected nodes) are connected to nodes that are not otherwise connected to one another. In this process. Brain graph construction One of the recent applications of complex network theory in neuroscience has been in the creation of brain graphs from neuroimaging data [30. in other words.

For example. 15. mentary avenue of research will be to study the inherent nectivity [33. an interesting comple- tural [7. Importantly. that are coherently active (functionally connected) with cal modularity is thought to be compatible with an evolu.humanconnectomeproject.70].org/) [43] parallels similar [7. two regions of the brain Anatomically. ment on the outside of the brain allows the complex wiring The mapping of neuronal connectomes from many spe- diagram of the brain to contain short-range energetically cies represents an extensive effort in data collection and efficient connections [40]. provided direct evidence for both functional [31] and struc. brain metric geometry of the brain’s connectivity (independent of regions perform distinct roles either as hubs of high con.8. No. In fact the large majority of the energy budget in efforts in the mouse [44. whereas nodes that do not share many links are likely to be assigned to different modules. sustaining Mounting evidence suggests that the physical constraints varying amounts of synapse development and redevelop. structural [6] and functional [35] connectivity networks and might have neurophysiological correlates: each region Relationship between structure and function of the brain is differentially energetically active. attempt to comprehensively map the wiring diagram of the The physicality of wiring constraints is also compatible human brain at increasing levels of spatial resolution with the inherent spatiality of the observed connectivity: Figure 4). on the anatomical organization of the human brain con- ment or plasticity [36]. this organizational principle of hierarchi. This cal network curvature [41]: a description of the inherently stratification of regional roles is in fact evident in both curved space in which a network’s organization exists. Within these modular structures. of the brain and its connections. Components that subserve a similar function are said to belong to a single module. Modularity The general concept of modularity is that the components of a system can be categorized according to their functions. The Human Connectome Project (http:// consumption in developing and maintaining wiring www. one another are often connected by a direct white matter tionary pressure for the minimization of energy pathway [6. In light of the physical geometry will require significant advances in data acquisition [49]. independent predictors of brain function [48].42]. physical constraints) as measured by for example topologi- nectivity or as provincial nodes of local processing.32].47] in its the human brain is used for the function of synapses [38].8.34]. 203 . whereas components that subserve a second function are said to belong to another module. 5 General concept of modularity System Functions Modules Mathematical concept of network modularity Graph Partitioning Modules TRENDS in Cognitive Sciences Figure 3.45].32] hierarchical modularity of human brain con. this and similar efforts in future regions close together in physical space interact strongly will need to be supplemented by investigations into neu- [39] whereas long-range anatomical connections or func. strain its function. Modularity can also be defined mathematically in terms of network organization [30. the folding of the cortical sheet as well as its place. This categorization procedure is an important current area of network science research. romodulatory networks acting in parallel with structural tional interactions connect disparate modules [7. The categorization of nodes into partitions is a process known as ‘community detection’ because – rather appropriately – it detects communities or modules composed of highly connected nodes and delineates the boundaries between those communities. In connectivity networks or wiring diagrams but which are fact.()TD$FIG][ Opinion Trends in Cognitive Sciences May 2011. Nodes that share many common links are said to belong to a module.37]. Vol.33. worm [39] and fly [46.

mental ture–function mappings are many-to-many and inherently properties seem to be fundamentally indivisible [4]. system phenomena are explained by breaking or reducing tems and provides the structural basis for another defining the system down into molecules. C. erful it is also fraught with caveats. Consider that you can blow the mill up in size the identical anatomical substrate. the mind and the physical brain. 5 Wiring diagram components = Neuron = Connection Wiring diagram C. such of function [52]. although a one-to-one rela. All that you find are mechanical components tures specific to vision in the seeing adult might develop that push against each other but there is little if any trace into auditory processing units in the early blind [54]. imagine that you are walking with Leibniz through a [53]: the brain performs many (degenerate) functions on mill [56].8]) or to test ness – qualia) are more than the sum of the system’s parts specific hypotheses about the structural underpinnings at any particular level or across levels [4]. although the functional inter. The worm is known to contain 302 neurons and here only a small fraction of the connections known to exist between these neurons are displayed (the full connectivity would be too dense to visualize clearly in this way). however. However. characteristics of the human brain suggest that anatomical This analogy points to an important disconnect in the connectivity alone does not uniquely specify the global mind–brain interface: although the material components functionality of a circuit or brain system. the human brain system occurring between multiple phys- a more complicated emergence of function from multiscale ical and functional levels [57]. Rather struc. fundamental paradigm shift away from the so-called re- ductionism perspective in which the strongest explanatory Emergence power lies at the lowest level of investigation: that is. Furthermore. of the physical brain might be highly decomposable. function and/or other properties of the system atory analysis to characterize the organizational principles (e. [6. atoms. It is plausible that Perhaps most simply. Efforts to characterizing the interaction between two broad levels: this end are complicated by an inherent functional degen. this is the concept of emergence in which the wiring diagrams. can be very useful for explor. The analysis of wiring diagrams can be used to assess important organizational principles of biological system structure and might provide insight into system function [46].()TD$FIG][ Opinion Trends in Cognitive Sciences May 2011. No. emergence – of consciousness or structural connectivity might enable us to predict function otherwise – in the human brain can be thought of as but it is not yet clear how to make that prediction. elegans Tail Body Head TRENDS in Cognitive Sciences Figure 4. consciousness or the subjective features of conscious- underlying structural connectivity (e. The resulting phenomenon. system properties can emerge from complex patterns of pretation of the connectome is potentially immensely pow. Therefore. ly as occurring between the two levels of the mind and the tionship between structure and function might be brain. processing [50] and neuroinformatics [51]. The color of each node represents the number of connections emanating from that node (red indicating many and blue indicating few). Wiring diagram of Caenorhabditis elegans The wiring diagram of the nematode worm. particles and then 204 . elegans. To visualize this dichoto- eracy that might facilitate robustness and adaptability my. Multiscale organization is one hallmark of complex sys. In fact. degenerate because they depend on both network interac. Although emergence can be conceptualized most broad- tions and context. such that all components are magnified and you can walk tex is characterized by multipotentiality: anatomical struc. Vol. is composed of nodes (neurons) and connections between those nodes (electrical and chemical synapses). underlying subsystems. These of the function of the whole mill represented at this level. among them. the cor. emergence might be a more fundamental property of inconsistent with our current understanding of the brain. This idea represents a structure is plausible [55]. 15. behavior.g.g.

The mind–brain emergence therefore on its inherent organization. However. i. Furthermore. Furthermore. It also provides a framework in change. The mind–brain interface might be human brains display a critical level of complexity consis. Indeed. or – like genotypes and phenotypes – is one-to- results support the view that intervention at the phenom. that is. The strike of a match does often light a match. characterized by complex conditional causation in which tent with their emergent functions. The most effective thera. In addition to emergence and multiscale Emergence is characterized by a higher-level phenomenon structure. Finally. No. 15. Interest. mental intervention). However.42]. the match was wet) and neither characterizes the brain system? Are different types of is it a necessary cause (e. however. This nonreducibility of the brain might be predicated types of emergence. how a human traverses mind-space as it passes through cal abnormality. there is no way to know appropriate co-solution of an inherently neurophysiologi. it is not necessarily a sufficient cause for the standing its emergent properties. unlike correlation or determinism [60]. a machine might generate a machine as complex as The combination of upward emergence and downward itself. What type of emergence lighting of the match (e. causes are neither necessary nor sufficient under all cir- cumstances. suggesting that might not be simple.g.Opinion Trends in Cognitive Sciences May 2011. several different categories causal phenomenon can be realized in many different are often used including substance (a baby emerges from ways. for example in the information about correlations between mental and physi- treatment of clinical depression. which is not a simple sum of requires a tailored definition. sation seems to occur both upwards and downwards structural (three lines make a triangle). The mind emerges from the constraining environment of mental and physical functions brain in a way that is arguably unlike any of these weak [59]. complex systems often display so. Vol. However. Mental states emerge from the parts with which it is organized. serotonin reuptake inhibitors (SSRIs) (pharmacotherapy. however. molecular intervention) and cognitive behavioral ther. many (one brain state corresponds to many mental states enological level of thought processes and beliefs is an or vice versa). it is important to describe cause and scale system? When is the interaction between levels effect at the right level of generality and to determine simply correlational and when is it causal? which levels of data are pertinent. a process Measurement of emergence that poses significant difficulties in the context of deter. might be beneficial in characterizing mind–brain emer- The study of altered minds is perhaps one of the greatest gence. do not exist or change unless physical properties exist or ability of system function. evidence suggests that neither intervention alone is states is one-to-one (one mind state corresponds to one as effective as the two interventions together [63]. stemming from a lower system level. plexity. there are other ways to light a emergence present between different levels of the multi. in most cases. These brain state). 5 subparticles. some of the direct) measurements of physical and mental states strongest lines of evidence for nonfundamental causality in [28. However. level changes through what is called downward causation. Because a high-level In describing emergence. functional (letters between multiple levels (either neighboring or distant) form words) and real (a cell is alive unlike the molecules of of the system [58] creating a complementarity or mutually which it is made) emergence. The interpretation of results from this experimental line i. opposed to some other complex systems. which to place human responsibility [61] and relate neu- roscientific advances to ethics and law [59.62]. an important property of the brain. not a property of a molecule but of a group of molecules). The lighting of a match can serve as an Types of emergence example. at a given threshold of com. allows ducible and highly dependent manner: mental properties for mutual manipulability over levels and multiple realiz. cal states but do not necessarily provide insight into the py for depression combines the administration of Selective underlying emergent phenomena. biological systems such as the brain a mother).g. a machine might causation suggests a simple bidirectionality or more generate another machine more complex than itself. can hear more sounds and can Defining causation for the mind–brain system. Understanding the brain depends significantly on under. conjunction (two parts can perform a different are fundamentally nonreducible in the sense that nonfun. match). that is in a nonre- causality.e.e. brain states. Without such a map. upward. Neuroimaging tools such as fMRI provide coarse- tools currently available to assess the mind–brain interface grained statistical evidence for relationships between (in- and the characteristics of emergence. The nuanced mutual complementarity [59] that adds to the human brain is an example of a machine that can generate complexity of the system. property (wetness is damental components have significant causal power: cau. compute more swiftly than their makers. these properties of emergence and causation are inherent to complex systems that show multiscale Bidirectional causation and complementarity organization. and underscores the fact that the a plethora of other machines with functions other (and emergence of mental properties cannot be understood possibly more complicated) than its own. Empirical measurement of the mind–brain continuum ministic reductionism. machines can see farther. However. understood whether the mapping from mind states to brain ingly. emergence is called ‘threshold behavior’: a phenomenon studied for ex. of inquiry is complicated by the fact that it is not yet apy (psychotherapeutic. as ample in the context of the behavior of finite automata [64]. Human-made using fundamental reductionism. is that emergent Von Neumann noted that simple machines might generate phenomena can feedback to lower levels. Nonfundamental physical states by strong emergence. above that threshold. function than either part separately). the range of realizers that can lead to the same or 205 . these studies provide the human brain come from psychiatry. causing lower simpler machines.

to establish more than a metaphor might re. structure and function. and feedback loops to allow learning to occur. Vol. Box 2. complexity theory attempts to provide of a computer whose connections mimic those of the brain a compact representation of the dynamics of an aggregate (e. tional organization [30].65]. engineering of a brain-inspired robot with a continuous work in which to study the organizational structure of sensory motor flow experience highlighted several funda- complex systems made up of many interacting parts and mental organizational principles necessary for complex can therefore be thought of as an extension of statistical motor behaviors [67]. specific hypothesis testing of neurophysiological mechan- cept of phase transitions: thresholds above which the isms of brain and behavior. gence.88] for precise motor control [89]. recent work in this area has suggested that the brain functions as an optimal controller [86] that Fields uses feedforward modeling [87. the question of how the mind and the brain relate to and consistent organizational principles can be identified. systems as well as those that are unique to a given system. the development of more The concept that emergence of complex behaviors might nuanced null models in future will provide the basis for occur through the interaction of multiple temporal scales is 206 . retraction concepts with which it is familiar [68]. (thought) states of aggregate systems. 15. Complementary For example. large-scale physical models can be con- advances in the newer fields of complexity science and structed that attempt to mimic brain organization or network theory might also shed light on the emergence structure. A crucial advance in this area will require an assessment of dynamic tion of physical models. in which each module operated at a different time scale. has served as a scientific metaphor is a need to identify principles that are shared across for the mind–brain phenomenon. pruning. efforts have touched on the theoretical have focused on representing the brain system as a static network. system displays a new characteristic behavior.69] and the relationship between the anatomical organization and cognitive function [42. these and absorption. including the interactions between for testing such inter-level causal relations. the construc. Such efforts might range from the construction debate. couplings of neuronal oscillations [69]. No. Network theory.g. provides a new frame. empiri.g. Initial work has capitalized on principles derived from several branches of classical physics in which a common quantitative definition of emergence is related to the con. Clarifying these vari. For example. Recent applications of network theory have focused the system required a hierarchically modular architecture on descriptive statistics of the brain’s structural and func. one another is one for which there are few quantitative Statics to dynamics Although recent applications of complex tools and researchers often turn to simple intuitive meta. However. To date. to the construction of a adaptive system by collapsing the system’s many degrees of robot whose behavior mimics that of a human. where the robot can respond to commands tion that can be linked to underlying cellular mechanisms it has not heard before if they are made up of isolated such as synaptogenesis. application of physics and complexity theory. by contrast. For example. 5 similar effects must be determined. see [83] for technical and social metaphors. systems provided by complex network theory. A classic example from statistical physics is the Ising model that Metaphors although having little to do with neurons and everything to In addition to the development of theoretical models. and the development of both changes in that architecture with cognitive function (e. include the analysis of evolutionary pressures on brain Dynamical systems theory To complement a focus on an under- development and the construction of detailed multiscale standing of the underlying topological architecture of cortical multidisciplinary theories and models that coalesce cur. we often turn to both biological and nonbi- quire inherently new physics to deal with functional ological metaphors of the brain. and necessary to characterize relations between multiple levels of the potentially might facilitate the formulation of experiments multiscale human brain system. In human brain structure and the role of cross-frequency both exploratory and model-based investigations. and therefore cal results are necessarily compared to stated null models support current hypotheses regarding brain–behavior that are often based on purely random organizational mechanisms. Importantly. behaviors. proliferation. which can provide insight into the sparsity of connections between neurons and between fundamental principles of mind-brain phenomena. These network models could vary from the properties are consistent with current understanding of simple and stylized to the multiscale and complex [65]. the instantiation of current hypotheses regarding principal dynamics. However. neuromorphic computing [66]). mathematically compared to connectivity at the functional level [82]. Importantly. network theory to neuroimaging data have provided unique insights into brain structure and function. it is important to rent distributed patterns of knowledge. In both freedom into only those necessary for a description of the cases. Complex network theory provides one important context in which to address this Investigative tools question: connectivity at the anatomical level can be directly and Although it is important to philosophically define emer. Crucial frontiers ables might enable a better understanding of causation Structure–function To understand mind–brain mechanisms it is and complementarity between the mind and the body. However.Opinion Trends in Cognitive Sciences May 2011. there do with electron spins. characterize the function of the brain on this architecture as a dynamical system [65. Complex systems that can be distilled mind–brain mechanisms can be directly tested for realistic in this way are relatively robust to environmental perturba. the recent dynamical systems tions. generalizability. Additional insights (Box 2) will need to be provided by The distributed architecture also allows for impressive dynamic network models of brain development and func. In this context. modules. physics.85]. relatively a potential mathematical framework to assess dynamic changes in modular organization and [84] for a recent application in the context neglected areas of inquiry that might prove highly fruitful of human learning). temporal scales [55. the majority of studies to date phors. To produce behavioral complexity.

less lofty levels of been simply likened to the relation between software and description. One area of evolutionary theory that might be particu- ment of aggregates is a direct consequence of temporally larly useful in understanding the human brain is multi- dependent system uncertainty which. No. relationship to an information-rich environment. stand brains it might be necessary to understand the For example in the study of visual plasticity a software evolutionary forces of natural selection and the environ. In this case. 15. Striking similarities are evident. suggesting compatibility of these data is often tenuous. they provide are not well understood. neural of this size it might not be clear at the outset what the Darwinism). investigators and scientific communities. a tage of fissioning into independent replicators. modalities [76]. systems has its limitations and confounds. or a different phenomenon efficiency in information processing systems. whereas when higher order associa. by contrast. is not confined to neuro. therefore. Models and theory Historically. specifically very large-scale integrated animals models to support their own results. date the possible convergence between these two disparate systems. spatial resolutions. The fact that this signal ture and then transmit information that grows nonlinearly occurs over longer time scales allows for the decision to in the number of individual components. but with different tools. better understanding of the sequence of selective pressures tions in BOLD signal. issues of neuroinformatics [51]. It is difficult to that both technological innovation and natural selection parse whether it is the same phenomenon that is mea- have discovered similar solutions to the problems of wiring sured. whereas selection itself is a form of develop. The field of neuroscience is flooded with highly hardware in a computer system. These are per- puter and the Internet [72]. The social system in to members of a fused aggregate can overcome the advan- these animals is composed of an interaction network. robots. expected BOLD response. the large-scale wiring diagram ment techniques all measuring the same system. animal societies suggests that emergence or the develop. The mind–brain dualism has haps most troubling at the more detailed. vastly outweigh the information available to individual tion areas are recruited responses might drop to 5 Hz. Increasingly inclusive ‘levels’ are able to cap- adaptation of the system [71]. cells having to accumulate information independently. modeling environment could be developed in which infor- ment [73]. the drawing of links between disparate perhaps the most recent examples of which are the com. A canonical fight network and a submissive–dominant signal network example is the evolution of multicellular life and the that change on short-. biology – is in terms of a nonlinear increase in the efficiency Some neurophysiological evidence for the emergence of of information processing. many non-neurophysiological systems Despite the usefulness of theoretical and physical meta- have been used as explanatory metaphors for the brain. perhaps unsurprisingly. Recalling Von Neumann’s observations [64] (see mation from single unit physiology. decisions based It is reasonable. storage. The amount of adaptive infor- slow time scales from faster time scales does exist: for mation available to cells in which subpopulations special- example when V1 is recruited. further work is necessary to eluci. In this way. it is possible that some erbated by the lack of detailed neurophysiological models important insights can only be gleaned by examining the that combine data and results from many experimental system that is doing the constructing. However. niche construction) [74]. It has been animal to other animals reduces uncertainty in the social hypothesized that major transitions of this sort are associ- system. it is plausible that these forces are at least as pathways and anatomical connectivity derived from tract complicated as the brain itself. given the implications of low-frequency oscilla. altogether. essence as the selection of construction rules. mounting evidence tracing or diffusion imaging. to under.g. the empirical circuits [7]. The submissive–dominant signal given by an which have forfeited the ability to replicate. 5 one that. This predicament stems from the fact that the Although the majority of metaphors have been drawn relations between present technologies or the data that from systems that are constructed (e. In fact. of animal–animal interactions. that one way to understand the on faster variables might only increase system uncertainty brain – one of the most complex multicellular structures in and thereby decrease both predictive and adaptive power.and long-temporal scales subsequent emergence of specialized cell types many of respectively. could be used to predict the supports the combined notions that the brain’s develop. The complexity of these is that connections exist between levels to provide a means evolutionary forces can be synthesized with the intuitive of quantifying whether results are consistent with stan- simplicity of natural selection by viewing evolution in dards of the field and collective understanding. in social systems can level selection theory [75] that seeks to explain how payoffs be based on misaligned interests [70]. Vol. integration and mobility is expected to might occur at 40 Hz.g. subsystems. what is imperative ment (e. and is further exac- the Internet and social systems). Recent work characterizing power structures in the products of these rules.g. researchers often point to studies using other measure- tional structure has been directly compared to that of ment streams. and therefore allows for logical tuning and information. retinal response times ize in sensing. That is. informed by molecular above).Opinion Trends in Cognitive Sciences May 2011. species or computer chips. phors of the brain. lowest relevant level of data should be. it is plausible that slower time scales leading to these advantages will lead to an improved are important for the understanding of cognitive function understanding of the functional role of a brain and its and control. A Further. In a multidimensional endeavor ment is in part a form of selection dynamic (e. Similarly. Due to recent advances in detailed data produced by a plethora of different measure- diffusion weighted imaging. computers. computational 207 . Although of the human brain has been estimated and its organiza. medium. defines that animal’s place in the emergent power ated with novel mechanisms for storing and transmitting structure. selection must be complex enough to discriminate among science. create that signal to be based on a long history or memory higher levels can offset the replicative cost of social living.

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