Neuroscience and Biobehavioral Reviews 124 (2021) 235–244

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Neuroscience and Biobehavioral Reviews

journal homepage: www.elsevier.com/locate/neubiorev

A new era for executive function research: On the transition from

centralized to distributed executive functioning
Nicolas Zink a, *, Agatha Lenartowicz a, Sebastian Markett b
a
Department of Psychiatry and Biobehavioral Sciences, University of California, Los Angeles, Los Angeles, United States
b
Department of Psychology, Humboldt University Berlin, Berlin, Germany

A R T I C L E I N F O A B S T R A C T

Keywords: “Executive functions” (EFs) is an umbrella term for higher cognitive control functions such as working memory,
Executive functions inhibition, and cognitive flexibility. One of the most challenging problems in this field of research has been to
Working memory explain how the wide range of cognitive processes subsumed as EFs are controlled without an all-powerful but ill-
Inhibition
defined central executive in the brain. Efforts to localize control mechanisms in circumscribed brain regions have
Flexibility
Central executive
not led to a breakthrough in understanding how the brain controls and regulates itself. We propose to re-
Brain networks conceptualize EFs as emergent consequences of highly distributed brain processes that communicate with a
Cognitive control pool of highly connected hub regions, thus precluding the need for a central executive. We further discuss how
Distributed networks graph-theory driven analysis of brain networks offers a unique lens on this problem by providing a reference
frame to study brain connectivity in EFs in a holistic way and helps to refine our understanding of the mecha­
nisms underlying EFs by providing new, testable hypotheses and resolves empirical and theoretical in­
consistencies in the EF literature.

1. Introduction which are stored in short-term memory (Baddeley et al., 1986; Baddeley
and Hitch, 1974; Cowan, 1999; Diamond, 2013; Miller, 1956; Smith and
The ability to exert control over actions is essential for personal Jonides, 1999). Inhibition has been defined as “being able to control
autonomy and one of the most impressive yet poorly understood ca­ one’s attention, behavior, thoughts, and/or emotions to override a
pacities our brains endow us with. The processes that control actions strong internal predisposition or external lure, and instead do what’s
cover a range of cognitive skills referred to as executive functions (EFs) more appropriate or needed” (Diamond, 2013). Cognitive flexibility
(Diamond, 2013; Powell and Voeller, 2004). While the precise definition reflects the ability to consider multiple conflicting representations of a
of EFs remains a topic of debate (Duncan and Owen, 2000), there is a single object or event simultaneously (Jacques and Zelazo, 2005) and
common consent in terms of the importance of EFs for adaptive behavior selectively switch between actions, perspectives, and strategies for
in an ever-changing environment (Jurado and Rosselli, 2007; Norman appropriate action in a changing environment (Dajani and Uddin, 2015;
and Shallice, 1986; Rabbitt, 1997). Numerous theoretical frameworks Diamond, 2013). In addition to flexibly switching between several lab­
exist to classify different EFs and to describe how they work together oratory tasks, every-day life as well as modern work environments often
(Burgess et al., 2000; Diamond, 2013; Duncan and Owen, 2000; Luria require performance on two or more tasks at the same time, e.g. driving
et al., 1966; Miyake et al., 2000; Norman and Shallice, 1986). According while having a telephone conversation or simultaneously controlling
to Miyake et al. (2000) and Diamond (2013), at least three core func­ several displays in air traffic control. In this multitasking context, effi­
tions of EF can be identified, which are likely to represent distinct cient EFs are essential to select relevant information from the environ­
cognitive subsystems that nevertheless share a functional overlap to ment and to adjust performance to environmental demands by flexibly
some degree: updating and monitoring of working memory represen­ shifting between more serial or more parallel processing strategies
tations, inhibition and interference control, as well as cognitive flexi­ (Fischer and Plessow, 2015).
bility and shifting. Working memory (WM) is a cognitive system that
enables to manipulate and maintain restricted chunks of information

* Corresponding author.
E-mail address: [email protected] (N. Zink).

https://doi.org/10.1016/j.neubiorev.2021.02.011
Received 22 September 2020; Accepted 4 February 2021
Available online 11 February 2021
0149-7634/© 2021 Elsevier Ltd. All rights reserved.
N. Zink et al. Neuroscience and Biobehavioral Reviews 124 (2021) 235–244

2. Centralized control in cognition and cognitive neuroscience 2017; Koechlin and Summerfield, 2007; Miller and Cohen, 2001; Mon­
sell and Driver, 2000b; Pezzulo et al., 2018; Shenhav et al., 2016, 2013),
Substantial theoretical and experimental progress has been made in now a “macroconstruct” in which the central executive consists of
the past 50 years to describe how humans control their actions by discrete subsystems with anatomically segregated and functionally
selecting, processing, and prioritizing task-relevant stimuli (often at the specialized modules. In a typical neuroimaging study, the cognitive
expense of simultaneously present yet less relevant signals), and flexibly process of interest is isolated via correlation to an external manipulation
adapt to changing demands from the environment (Broadbent, 2013; thought to elicit that process, and associated with a brain region of in­
Deutsch and Deutsch, 1963; Kahneman, 1973; Norman and Shallice, terest. If this association is statistically significant, it is assumed that this
1986; Schneider and Shiffrin, 1977; Shiffrin and Schneider, 1977; process is located within this brain area. This strategy has ascribed
Treisman and Gelade, 1980; Wickens, 1991). Early cognitive models different aspects of EFs with specific anatomically circumscribed brain
have introduced a hypothetical central executive to their cognitive ar­ areas. For instance, goal-directed behavior with the ventromedial pre­
chitectures (Baddeley and Hitch, 1974; Norman and Shallice, 1986). frontal cortex (vmPFC; Hare et al., 2014, 2009; Kahnt et al., 2011; Lim
One unifying premise underlying these theories is that some sort of et al., 2011; Sokol-Hessner et al., 2012; Steinbeis et al., 2016; Vassena
central executive system is required that governs exerted control by et al., 2014) and dorsolateral prefrontal cortex (dlPFC; Hare et al., 2014;
“dampening” irrelevant and prioritizing relevant salient information or Kahnt et al., 2011; Milham et al., 2001; Sokol-Hessner et al., 2012;
schemes. But while the necessity of higher-order control functions that Steinbeis et al., 2016), inhibition with right ventrolateral prefrontal
control lower-order functions is self-evident, it does not explain how this cortex (vlPFC; Aron, 2007; Aron et al., 2014, 2004; Hampshire et al.,
central executive operates, reducing instead to the question of who is in 2010), conflict monitoring, selective attention, and computation of the
charge of controlling the central executive. This problem of expected value of control with anterior cingulate cortex (ACC; Alex­
self-recurrency has been widely acknowledged and has led to the criti­ ander and Brown, 2011; Botvinick et al., 2004; Bush et al., 2002, 1999;
cism that the central executive resembles an all-powerful but ill-defined Carter and Veen, 2007; Egner and Hirsch, 2005; Kerns, 2004; Kondo
“homunculus” that directs all processes that are not “automatic” et al., 2004; MacDonald et al., 2000; Milham et al., 2001; Ridderinkhof
(Monsell and Driver, 2000a). Any theoretical framework of EFs must et al., 2004; Shenhav et al., 2016, 2013; van Veen et al., 2001; Vassena
therefore not only specify how control operates, how it is implemented et al., 2014; Weissman, 2004; Weissman et al., 2003), and cognitive
at the neurophysiological level, and how it integrates other cognitive flexibility and voluntary motor action selection with basal ganglia (BG;
processes and their interrelatedness, but also formally address the Aron et al., 2003; Cameron et al., 2010; Chakravarthy et al., 2010; Pauli
question of what is controlling the controller in order to avoid an infinite et al., 2016; Redgrave et al., 1999; Stocco et al., 2010). The problems
hierarchy of central executives or homunculi (Logie, 2016). with such an approach have been extensively discussed and include
One of the most influential theoretical frameworks on the neuro­ subsequent tendencies towards reverse inference and inherent bias in
physiological mechanisms of EFs that has pushed beyond the central interpretation (Poldrack, 2006; Poldrack and Wagner, 2004). Further­
executive view has been proposed by Miller and Cohen (Miller and more, a modular perspective fails to appreciate the activity of this region
Cohen, 2001). This framework emphasizes the importance of the pre­ in the context of the other activities within the brain. The systematic
frontal cortex (PFC) as a key region to implement a wide range of EFs association of a region with an EF such as inhibition could therefore
through active maintenance of goals and the means to achieve them. reflect an emergent property of a broader neural network (e.g., involved
Rooted in the biased-competition model of selective attention (Desi­ in detection of salient cues, such as for inhibition; Hampshire et al.,
mone and Duncan, 1995), it cast the problem of control as a case of 2010), and thus risks to falsely localize higher-level cognitive processes
prioritization of competing perceptual or response representations. The at a lower level of processing (Eisenreich et al., 2017), especially in
role of the PFC is to bias activity in regions responsible for those rep­ highly connected regions such as the ACC and basal ganglia.
resentation, in accordance with a behavioral goal, that it maintains in Although it is intuitive to assume that conceptually distinct EFs must
memory. Thus, the PFC indirectly acts as a central executive and in­ also have distinct anatomical representations (Botvinick and Cohen,
tervenes in a top-down fashion by selectively prioritizing the relevant 2014), attempts to identify neuroanatomical correlates of the central
process. According to this perspective, exerted control in EF is executive (Baddeley, 2012; Botvinick et al., 2004, 2001; Braver, 2012;
domain-general and operates on specific subordinated cognitive func­ Diamond, 2013; Engle and Kane, 2003; Friston, 1994; Goldman-Rakic
tions in service of a higher-order task goal. EFs such as inhibition, et al., 1996; Goschke and Bolte, 2014; Hazy et al., 2007, 2006; Hommel
working memory and cognitive flexibility emerge from the dynamics of and Wiers, 2017; Koechlin and Summerfield, 2007; Miller and Cohen,
the PFC biasing goal-relevant activity patterns in the brain. In the last 20 2001; Monsell and Driver, 2000b; Pezzulo et al., 2018; Shenhav et al.,
years, this theoretical model has provided a versatile framework that has 2016, 2013) have not yet led to breakthroughs to describe their
generated much empirical evidence supporting the pivotal role of the neurophysiological mechanisms. The idea of subsystems that provide
PFC in many EFs. However, despite progress in understanding the specialized, subsidiary central executive functions is challenged by
neurophysiological mechanisms of how the PFC represents and imple­ empirical evidence suggesting that the same putative brain correlates for
ments behavioral goals, the mechanism by which it accounts for EFs EFs such as the PFC and ACC have been associated with functionally
remains elusive. Although the PFC may participate to a greater extent different EFs (Alexander and Brown, 2011; Botvinick et al., 2004;
than other areas in EFs, increasing evidence suggests that it does not act Braver, 2012; Bush et al., 1999; Carter and Veen, 2007; Hare et al., 2009;
alone. Damage to the PFC, for instance, may or may not be accompanied Kahnt et al., 2011; Kerns, 2004; Kondo et al., 2004; Lim et al., 2011;
by deficits in EFs, and neither site nor size of a prefrontal lesion allow for MacDonald et al., 2000; Milham et al., 2001; Ridderinkhof et al., 2004;
a clear prediction about the nature of resulting problem in executive Shenhav et al., 2016, 2013; Steinbeis et al., 2016; van Veen et al., 2001;
functioning (Alvarez and Emory, 2006). Vassena et al., 2014; Weissman, 2004; Weissman et al., 2003). This
In parallel, the advance of non-invasive neuroimaging techniques raises the question how a plethora of diverse EFs can arise from a rather
such as fMRI, in combination with experimental designs rooted in sub­ static neuroanatomical architecture (Park and Friston, 2013). In com­
tractive logic (Friston et al., 1996; Newman et al., 2001), contributed to plement, different EFs such as inhibition, flexibility, and working
the conceptualization of a modular brain architecture (Fodor, 1983; memory have been shown to activate large proportions of the cerebrum
Houk and Wise, 1995; Kanwisher et al., 1997; Minsky, 1988). The results and converge within PFC, ACC, and posterior parietal cortex (Niendam
could be construed as a fractionalization of the central executive (Bad­ et al., 2012). Moreover, the putative brain areas underlying EFs also
deley, 2012; Botvinick et al., 2004, 2001; Braver, 2012; Diamond, 2013; show large functional overlap, such as in goal directed behavior (ACC;
Engle and Kane, 2003; Friston, 1994; Goldman-Rakic et al., 1996; Carter and Veen, 2007; dlPFC; Hare et al., 2014; Kahnt et al., 2011;
Goschke and Bolte, 2014; Hazy et al., 2007, 2006; Hommel and Wiers, Sokol-Hessner et al., 2012; Steinbeis et al., 2016); vmPFC; Hare et al.,

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2014, 2009; Kahnt et al., 2011; Sokol-Hessner et al., 2012; Steinbeis

et al., 2016), prediction and evaluation of an action (ACC; Alexander
and Brown, 2011; Bush et al., 2002; Ridderinkhof et al., 2004; Shenhav
et al., 2016, 2013; Vassena et al., 2014); vmPFC; Vassena et al., 2014),
and selective attention (ACC; Bush et al., 1999; Milham et al., 2001;
Weissman, 2004; Weissman et al., 2003); vmPFC; Lim et al., 2011;
dlPFC; Milham et al., 2001), which challenges the functional specificity
of these regions. Arguably, the focus on activity of specific regions of
interests or single event-related components has therefore substantially
limited the possibility to identifying the neural correlates of EFs from a
whole brain perspective.
Increasing evidence suggests that these putative brain areas under­
lying EFs seem to interact. The ACC, for instance, has been found to
support PFC in maintaining and updating task-relevant representation in
working memory (Banich et al., 2000; D’Esposito and Postle, 2015;
Egner and Hirsch, 2005) and signal the need for control (Botvinick et al.,
2004; Hazy et al., 2007). Furthermore, studies indicate that cognitive
flexibility is orchestrated within BG (Aron et al., 2003), but also sup­
ported by PFC (Chakravarthy et al., 2010; Stocco et al., 2010) and ACC
(Nicolle and Baxter, 2003). Similarly, close interrelation between PFC
and BG were found to support WM processes (Ashby et al., 2005; Baier
et al., 2010; Chang et al., 2007; Frank et al., 2001; McNab and Klingberg,
2007; Schroll et al., 2012; Schroll and Hamker, 2013; Voytek and
Knight, 2010). Thus, once again, the data suggest that pivotal brain
areas that contribute significantly to EFs, such as PFC, ACC, and basal
ganglia, act within spatially distributed networks and include other
structures that may also be relevant to explain EFs. This point is echoed
in studies of neuropsychiatric disorders that exhibit impairments in EFs,
and that reveal aberrant development and functional connectivity in
spatially distributed brain networks (Assaf et al., 2010; Baker et al.,
2014; Bassett et al., 2012; Cerliani et al., 2015; Chai et al., 2011;
Cherkassky et al., 2006; dos Santos Siqueira et al., 2014; Du et al., 2016;
Fan et al., 2012; Fassbender et al., 2009; Franzen et al., 2013; Garrity
et al., 2007; Hull et al., 2017; Itahashi et al., 2014; Jafri et al., 2008; Jang
et al., 2011; Liddle et al., 2011; Lin et al., 2015; Manoliu et al., 2014; Fig. 1. Organization of a modular executive control network vs. a distributed
Meda et al., 2014; Murias et al., 2007; Öngür et al., 2010; Orliac et al., control network, in which two stimuli (stimulus A and stimulus B) lead to an
2013; Paakki et al., 2010; Pomarol-Clotet et al., 2008; Roiser et al., action outcome (action A vs. action B). A. Top-down controlled, hierarchically
organized control-modules govern automatized processing. In this system,
2013; Rotarska-Jagiela et al., 2010; Salgado-Pineda et al., 2011; Sripada
control and processing elements are distinct and localized to specific areas. B.
et al., 2014; Sun et al., 2014; Swanson et al., 2011; Tian et al., 2008; Tu
Distributed control networks, in which control and processing elements are
et al., 2013; Uddin et al., 2008; van Buuren et al., 2012; Wang et al.,
combined in individual agents. These agents form individual clusters. Adapted
2005; Weng et al., 2010; Whitfield-Gabrieli et al., 2009; Wilson et al., and modified from Eisenreich et al. (2017).
2011; Woodward et al., 2011).
Thus, mounting evidence suggest that there is no single brain region
Vandierendonck (2016) and Eisenreich et al. (2017), have argued that
that sits at the control apex, and, furthermore, efforts to define EF-
EFs can also be a result of a conceptually and anatomically distributed
dedicated neuroanatomical substrates have not been conclusive. The
control system (DCS). One such DCS view is inspired from early con­
question of how EFs are generated by the brain persists. In the following
nectionist models (Hopfield, 1982; James and Rumelhart, 1986), in
sections we consider an alternate view to centralized or localized EFs.
which control and controlled processes are co-localized within large
We argue based on suggestions from the cognitive and computational
numbers of dispersed elements (Fig. 1B). In contrast to a hierarchical
literature(Barnard and Bowman, 2004; Eisenreich et al., 2017; Van­
architecture where core brain regions are assumed to solely exert control
dierendonck, 2016), and in line with the recognized significance of
on more basic processes, in a DCS, each element can be a controller and
functional connectivity patterns to neural organization and function
an element that is being controlled. In computational models of
(Collin et al., 2014; Margulies et al., 2016; van den Heuvel et al., 2012;
distributed control, the state of the overall system is often independent
van den Heuvel and Sporns, 2013), that EFs are neither a strictly
from any constituting element (Barnard and Bowman, 2004). In other
top-down generated process nor one that can be localized in specific
words, in such systems, behavioral control is analogous to the state of
brain correlates, but rather that EFs are the emerging consequence of
the overall system, and is therefore an emergent function arising from
communication within a broad network of spatially and functionally
the interaction of all elements (McClelland et al., 2010; Mitchell, 2009).
dispersed brain areas that integrate different aspects of EFs.
Recent evidence of distributed control in nonbrain biological systems
such as in schools of fish, flocks of birds, swarms of insects (Couzin,
3. Distributed control and executive functions 2009; Eisenreich et al., 2017; Passino et al., 2008), and herds of baboons
(Couzin and Krause, 2003; Strandburg-Peshkin et al., 2015) and in
A core assumption underlying many theoretical frameworks on EFs is distributed deep learning algorithms (LeCun et al., 2015) highlight the
of hierarchical architecture, namely, that brain areas or circuits under­ strength and plausibility of a DCS view. Apparent control in these bio­
lying EFs regulate, but do not participate in the more basic cognitive logical distributed systems is often realized via a couple of simple rules
processes that they control (Botvinick et al., 2001; Miller and Cohen, that guide the indirect coordination between agents or actions. In
2001; Fig. 1A). This top-down view, by design, relies on a centrally schools of fish or flocks of birds, the control over the shape of the swarm
operating executive. However, Barnard and Bowman (2004);

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and the moving direction is a consequence of the distance kept between disease (Berman et al., 2016; Lebedev et al., 2014). As noted previously,
adjacent agents and the rule about when to change direction (i.e. follow damage to the PFC, as well, may or may not be accompanied by deficits
the group average; Couzin and Krause, 2003). These rules enable fast in EFs, and neither site nor size of a prefrontal lesion allow for a clear
communication of information (e.g. about the direction of movement) prediction about the nature of resulting problem in executive func­
across all agents, without reliance on a top-down controller. tioning (Alvarez and Emory, 2006). Thus, EFs do exemplify some
Can a DCS framework explain EFs? EFs can be conceptualized as a robustness in the human brain. One way to investigate the robustness of
consequence of the interaction between distributed elements, thus EFs as a DCS is using ‘lesioned’ networks in computational models,
avoiding the problem of any central executive or central executive which investigate how an observed profile of anatomical or functional
subsystem. Increasingly evidence suggests that, indeed, properties of EF dysconnectivity in a mature network might have been generated by
generation in the brain are consistent with properties of a DCS. One such earlier developmental abnormalities (Achard, 2006; Honey and Sporns,
principle is that all elements can be controlled or be a controller, or, in 2008; Kaiser et al., 2007; Ravasz and Barabási, 2003; Sporns, 2006). In
other words, the emergent property can be observed in any subset of the order to explore the effect of acute damage on overall performance in
system. Several studies have highlighted that prefrontal regions, a pu­ EFs, nodes or connections are deleted. The vulnerability of a network to
tative apex of the central executive, as well as posterior regions seem to damage is then assessed by comparing its efficiency after the lesioning to
exhibit both basic and control processing (Awh and Jonides, 2001; Cisek its intact behavior. In an anatomically informed computational model,
and Kalaska, 2010; Postle, 2006; Sleezer and Hayden, 2016). In line with in line with the DCS view, deletion of nodes did not impair EFs.
the DCS view, it has been suggested that working memory (WM), a core
EF, is a general property of the cortex, not limited to specific regions (Lee 4. A new era for executive functions
and Baker, 2016; Postle, 2006). This proposal shifts perspective of WM
from localized to a highly distributed process wherein information can Despite the appeal of the DCS view as an elegant theoretical solution
be maintained in any system engaged in the initial perceptual process­ for the aforementioned problems with centralized EF frameworks, and
ing, including any region contributing to the initial percept (Lee and despite indirect evidence for the viability of DCS as a putative alterna­
Baker, 2016). The second key principle of a DCS is the existence of local tive, direct empirical studies of the neurophysiological mechanisms of a
rules capable of generating the emergent phenomenon, such as the DCS governing EFs are scarce. This may be attributed in part to limited
distance kept between adjacent agents in a swarm. In the brain, from awareness of this framework in the field and in part to inherent diffi­
single neuron firing to neuron populations and between brain area culties in translating DCS concepts, such as the existence of organizing
communications, each level is a complex function of its lower level rules, to testable hypotheses. A key challenge is to derive descriptive
constituents and embedded in a larger-scale organization (Buzsáki, measures that could provide testable hypothesis of the organizational
2006). In this context, EFs such as the initiation vs. the inhibition of an rules that drive the network towards emergence of EFs. However, one
action or switching to another action might be the emergent behavioral promising approach to investigate EFs from a DCS perspective is the
outcome from simpler local neurophysiological mechanisms of the application of methods from network science, primarily graph theory, to
participating brain areas, for instance via neuronal interactions resulting neuroimaging data. A DCS conceptualizes executive functioning as an
in local gain modulation (Abbott, 1997; Chance et al., 2002; Donner and eminent property of multiple interacting elements of the brain. This is
Nieuwenhuis, 2013; Saalmann and Kastner, 2009; Salinas and Thier, analogous to the network science framework, which aims to summarize,
2000) and the collective state of each contributing neuron (Buzsáki, via a family of derived metrics, the organizing principles of a set of
2006). In particular, evidence supporting biased competition among connected nodes. Hence, the graph theory framework is naturally
neuronal populations in generating initiation or inhibition in selective aligned with a distributed perspective on EFs, with derived metrics
attention (Desimone, 1998; Desimone and Duncan, 1995), suggests that potentially capturing the organizing principles or local rules that guide
such competition is a general property of neurons across the brain, not the behavior of the system. The task then becomes to model the brain as
limited to any central controller. From this perspective, the emergence a network in which brain areas or constellations act as nodes (i.e. agents)
of executive control patterns (e.g. initiating or inhibiting an action or and functional communication among these nodes are represented as
switching to another action) observed in putative control structures like edges, and, critically, to relate the static and dynamic features of the
the PFC, ACC, and basal ganglia may reflect the overall state of the resulting network to EFs. In contrast to putting emphasis on the iden­
system, in the same way a single fish in a swarm or a single bird in a flock tification of specialized subsystems of EFs that are assumed to exert
reflects the state of the overall system (i.e. the trajectory of the move­ control over basic processes, a network analytical approach puts focus
ment), without the requirement to be on top of the hierarchy in the on how these brain regions or sub-systems (i.e. network nodes)
control system. communicate (i.e. are connected) with each other.
Another key property of a DCS is its robustness to perturbations. In From a network perspective, this means that functionally different
contrast to centralized systems, in which a nonbrain biological systems EFs such as inhibition, and cognitive flexibility can recruit a set of
such as swarm would be vulnerable to the loss of its leading agent, a similar or overlapping brain structures (i.e. PFC, ACC, and other re­
swarm organized as a DCS has been shown to be robust to degradation gions), but the connectivity patterns within these networks between
(Sumpter, 2006). Similarly, decentralized (i.e. distributed) networks perceptual input and motor output are functionally different. The con­
have been shown to be resilient systems which are capable of absorbing nectivity patterns between the involved agents can, in principle, either
large external perturbations without undergoing functional breakdown result in refraining from an action (i.e. inhibition) or shifting to another
(Achard, 2006; Bassett and Bullmore, 2006; Bullmore and Sporns, 2009; action (i.e. cognitive flexibility). In other words, we hypothesize that the
Buzsáki, 2006). A DCS network organization in the brain may therefore emergence of a given EF may lie in a given network state, as determined
explain how EFs can be preserved to some extent in the face of patho­ by organizing rules.
logical attack by lesion and substance-related disorders (Ahmadlou One important characteristic of networks, and putative organizing
et al., 2013; Wang et al., 2015; Yuan et al., 2010), neuropsychological rule, is the structural and functional efficiency of its architecture. From
disorders like ADHD (Ahmadlou et al., 2012; Liu et al., 2015; Wang an evolutionary perspective, brain networks have been evolved to
et al., 2009; Xia et al., 2014), schizophrenia (Liu et al., 2008; Michel­ maximize the diversity of possible functional circuits (i.e. enabling
oyannis et al., 2006; Rubinov et al., 2009; Shim et al., 2014), and restless functionally different EFs), while minimizing the diversity of structural
legs syndrome (Choi et al., 2017), aberrant development such as in circuits (i.e. enabling efficient assembly and encoding) (Sporns and
autism (Barttfeld et al., 2011; Itahashi et al., 2014), and cognitive Kötter, 2004). Thus, the brain seeks towards minimal complexity of
decline such as Alzheimer’s (Frantzidis et al., 2014; Stam et al., 2006; neural architecture with maximal richness of function. This may explain
Vecchio et al., 2016; Wei et al., 2015; Zeng et al., 2015) and Parkinson’s why putative brain correlates of specific EFs (i.e. PFC and ACC) have

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also been associated with other functionally different EFs (Alexander small-world properties in the involved functional networks (Wolff et al.,
and Brown, 2011; Botvinick et al., 2004; Braver, 2012; Bush et al., 1999; 2017), suggesting that EFs emerge when the brain is tipped away from a
Carter and Veen, 2007; Hare et al., 2009; Kahnt et al., 2011; Kerns, small-world state. This would predict that training EF capacities should
2004; Kondo et al., 2004; Lim et al., 2011; MacDonald et al., 2000; lead to more efficient (i.e. more ‘small-worldish’) functional network
Milham et al., 2001; Ridderinkhof et al., 2004; Shenhav et al., 2016, communication. Consistent with this prediction, training in EFs such as
2013; Steinbeis et al., 2016; van Veen et al., 2001; Vassena et al., 2014; WM capacity has shown to improve the efficiency of functional networks
Weissman, 2004; Weissman et al., 2003). From a graph theoretical in the brain (Langer et al., 2013).
perspective, the efficiency of large-scale communication between Of particular interest, small-world architecture results in a system
different brain areas is defined by the number of connections necessary that is neither exclusively top-down in information flow, or exclusively
to connect all neurons, a measure that is referred to as average synaptic distributed. The system necessitates the existence of so-called hub
path length (Fig. 2A). Additionally, metabolic efficiency in brain net­ nodes, that are highly connected and serve a key role in facilitating the
works is realized by predominantly local connectivity (Yoshimura et al., balance between high-local clustering and low average path length (van
2005) or local clustering (Fig. 2B), which avoids metabolically expen­ den Heuvel and Sporns, 2013; Fig. 2C). However, such hub-nodes, due to
sive long-range axonal connections (Kalisman et al., 2005). However, a their high-connectivity, can, in a way, exert control over less-strongly
purely local clustering in the network would dramatically increase the connected nodes along the action cascade and thus may play a key
average number of connections necessary to connect all neurons (i.e. the role in driving the functional connectivity state of the system and the
average path length). Thus, two network organization principles—local emergence of behavior (Fig. 2C). Supporting this notion, in a simulated
clustering and average path length —compete with each other to model, deletion of well-connected nodes produced disruptions of func­
providing network communication that is metabolically efficient (i.e. tional connectivity (Falcon et al., 2015; Honey and Sporns, 2008; Jirsa
high local clustering), but also enables effective large-scale communi­ et al., 2010) consistent with reported effects of focal human brain lesions
cation (i.e. low average path length). Networks, which enable efficient on EFs (He et al., 2007a). These facts have also led to the proposal that
large-scale traffic while maintaining mainly local connectivity are called there exists a connective core network, that is a metabolically costly yet
‘small-world’ networks (Watts and Strogatz, 1998; Fig. 2E). Despite highly integratory collective of brain regions, whose anatomical wiring
considerable heterogeneity in the methodological approaches, there is and topological position within the brain network supports brain-wide
an encouraging degree of convergence between small-world properties communicability and which exerts control over brain dynamics and
on the structural (Chen et al., 2008; Hagmann et al., 2007; He et al., cortical states (Collin et al., 2014; de Reus and van den Heuvel, 2014;
2007b; Iturria-Medina et al., 2008, 2007; Sporns and Kötter, 2004) as Senden et al., 2017; Shine et al., 2018; van den Heuvel et al., 2012; van
well as on the functional brain network level (Ferrarini et al., 2009; Liu den Heuvel and Sporns, 2013). Here we additionally suggest that such a
et al., 2008; Meunier et al., 2009; Salvador, 2004; Schwarz et al., 2008; network drives brain states in observance of or striding towards a
Wang et al., 2009; Xia et al., 2014). Thus, the small-world properties of small-world state, with no single region acting as a core executive or
neural networks may be a key organizing principle that provides a solely responsible solely for the maintenance of behavioral goals (e.g.,
functional link between brain structure and EFs (Beste et al., 2019). PFC). It is sensible to suppose that regions previously associated with
Namely, we suggest that EFs are the result of the brain’s need to be EFs (i.e. PFC, ACC, basal ganglia) contribute to this network, though the
structurally and functionally efficient. Recent findings highlight that the identity of this network, or networks, remains an open question. A large
efficiency of how brain networks communicate (i.e. the small-world scale meta-analysis of fMRI literature has shown that a superordinate
network characteristic) is strongly related to the demand on EFs. Dif­ fronto-cingulo-parietal network underlies a range of different EFs such
ferences in small-world properties have been shown for different de­ as inhibition, flexibility, and working memory, consistent with expec­
mands on EFs (Beste et al., 2019; Wolff et al., 2017; Zink et al., 2018). tation, but also integrating the inherent biases in published literature
Higher demand on inhibition (Hong et al., 2016), cognitive flexibility (Niendam et al., 2012). In contrast, in an original research study, Dos­
and WM capacity, for instance, have been found to lead to decreased enbach et al. (2007), used network analyses on a set of 39 predefined

Fig. 2. Graph theoretical characteristics of distributed control

networks in the brain. Functional integration capacity in neu­
ral networks can be quantified by the average path length (A),
which describes the average number of edges (marked in blue)
to connect each node to any other node in the network (marked
in blue). Functional separation capacity in neural networks can
be quantified by the clustering coefficient (B), which describes
the interconnectivity of neighboring nodes. The clustering co­
efficient is exemplified in (B), where connections for one node
(marked in red) to its immediate neighbors (marked in green)
are colored in red and the connections between these neigh­
bors are marked in blue. All real networks lie on a spectrum
between very regularly and very randomly connected. Regular
networks (D) exhibit a high clustering coefficient and a high
average path length. Random networks (F) show a low clus­
tering coefficient and a low average path length. According to
Watts and Strogatz (1998), a network shows small-world
network properties (E) when it demonstrates a high clus­
tering coefficient as in regular networks and a low average
path length as in very random networks. In case a network
shows small-world properties, highly connected nodes form
hubs that integrate high local connections with long range
connections (C).

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tion and another network including the ACC, that was more strongly Baker, J.T., Holmes, A.J., Masters, G.A., Yeo, B.T.T., Krienen, F., Buckner, R.L.,
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