Cognitive Neuroscience - Psychology - Oxford Bibliographies - Cognitive Neuroscience

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Cognitive Neuroscience -Psychology -Oxford Bibliographies - Cognitive

Neuroscience
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DOI: 10.1093/OBO/9780199828340-0015
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Cognitive Neuroscience
Teal S. Eich, Edward E. Smith
Introduction
In 1918, the American philosopher and psychologist William James wrote: “Nature in her unfathomable
designs had mixed us of clay and flame, of brain and mind, that the two things hang indubitably together and determine each other’s
being but how or why, no mortal may ever know” (Principles of Psychology, 1918, p. 200). The study of how the brain produces
thoughts and behaviors is referred to as cognitive neuroscience (CNS). CNS is defined as an interdisciplinary field that combines
neuroscience and cognitive psychology. Neuroscience is the scientific study of the central nervous system. Cognitive psychology is a
branch of psychology that explores human cognition (Latin cognitiōn-em, a getting to know, acquaintance, notion, knowledge [Oxford
English Dictionary]), or the internal mental processes, including learning, memory (including long term and short term), perception,
attention, cognitive control, language, motor control, decision making, and social cognition. CNS is devoted to understanding how the
human brain supports, through neural mechanisms, these cognitive processes. For example, the “primacy effect” in memory is a
cognitive phenomenon in which memory for items that appear at the beginning of a list will be better remembered than items that
appear toward the middle of the list. Cognitive psychology helps us to understand why and when this phenomenon occurs: the first
items are rehearsed more than the middle items because there are fewer interfering items at the beginning, and therefore the first items
are encoded more strongly into long-term memory. CNS would help us to understand what brain mechanisms contribute to this
phenomenon: The medial temporal lobe (an area long known to be involved in the formation of memories) is activated only for items
from the beginning of the list. Thus, rather than trying to simply understand how and when a memory is formed, CNS attempts to
discover how the brain allows for the formation of memories. The methods and technologies used to study these aspects of human
cognition are diverse. Cognitive neuroscientists perform behavioral tests on both animals and humans inside and outside of the
laboratory. Numerous types of structural brain imaging and functional brain-imaging technologies are used in CNS (for example, MRI,
fMRI, EEG, PET, CAT, MEG), and researchers also employ computational modeling, genetic and candidate gene studies, and
pharmacologic manipulations to better understand how the brain underlies cognitive processes. Research from numerous scientific
disciplines in addition to neuroscience and cognitive psychology are also integrated into the study of CNS, including social and affective
neuroscience, neurology, pharmacology, and computational neuroscience.
General Overviews
Readers wishing to gain a historical perspective on the formation of the field of CNS should look to Bennett and Hacker 2008 as well as
Moskowitz 2010. Both books provide broad historical overviews of the field, including major theoretical advances from the 20th century.
Crick 1995, Gazzaniga 2000, and Purves 2008 also provide excellent overviews of the major themes in CNS, and these works are
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intended for an introductory reader. LeDoux 2003 covers a more narrow field of study, including topics in emotion and motivation.
Bennett, M. R, and P. M. S. Hacker. 2008. History of Cognitive Neuroscience. Hoboken, NJ: Wiley-Blackwell.
This book explores the major neuroscientific experiments and theories undertaken and advanced during the last 150 years.
Crick, F. 1995. The astonishing hypothesis: The scientific search for the soul. New York: Scribner.
Aimed toward researchers and also lay persons who are interested in science in general, this book focuses on the cornerstone idea of
CNS, that all mental processes are due to neural function.
Gazzaniga, M. 2000. Cognitive neuroscience: A reader. Malden, MA: Blackwell.

An ideal source for students, this book provides a collection of seminal readings within the field of CNS.
LeDoux, J. 2003. Synaptic self: How our brains become who we are. New York: Penguin.
An excellent source for advanced undergraduates and scientists in the field, Ledoux covers topics including cognitive, emotional, and
motivational functions of the brain, with an emphasis on how synaptic connections allow for neural communication and eventually
enable individuality.
Moskowitz, M. 2010. Reading minds: A guide to the cognitive neuroscience revolution. London: Karnack.
A practical and accessible guide to the cognitive neuroscience revolution.
Purves, D. 2008. Principles of cognitive neuroscience. Sunderland, MA: Sinauer.

One of the best basic CNS books available, aimed toward advanced undergraduates, this book provides an overview of topics within
CNS that are well established as well as a set of issues that remain to be solved by future generations of scientists.
Textbooks
A number of textbooks are available that are suitable for both introductory readers and those who are more advanced. All provide
excellent references. Baars and Gage 2010, Gazzaniga 2009, and Smith and Kosslyn 2006 constitute excellent, comprehensive
introductory level textbooks. Banich 2004 takes evidence from clinical populations, while Bear, et al. 2007 emphasizes the biological
systems involved in cognition. Finally, Springer and Deutsch 2001 is a more advanced text, focusing upon specific aspects of brain
function.
Baars, B. J., and N. M. Gage. 2010. Cognition, brain, and consciousness: An introduction to cognitive neuroscience. 2d ed.
Burlington, MA: Academic Press.

An introductory undergraduate-level textbook that will introduce students to the main concepts and topics within CNS.
Banich, M. T. 2004. Cognitive neuroscience and neuropsychology. Florence, KY: Wadsworth.

A clear and accessible introductory level textbook with an emphasis on both experimental and clinical perspectives of topics within
CNS.
Bear, M. F., B. W. Connors, and M. A. Paradiso. 2007. Neuroscience: Exploring the brain. 3d ed. Baltimore: Lippincott Williams
& Wilkins.
An introductory-level textbook with an emphasis on the biology of the brain and the neural systems that support behavior.
Gazzaniga, M. S., ed. 2009. The cognitive neurosciences. 4th ed. Cambridge, MA: MIT Press.
A broad and encompassing undergraduate-level textbook; includes topics that range from the molecular structure of neurons to
language and consciousness.
Smith, E. E., and S. M. Kosslyn. 2006. Cognitive psychology: Mind and brain. Upper Saddle River, NJ: Prentice Hall.
This is a survey of the major topics in CNS, with an emphasis on the cognitive processes involved. The book covers everything from
early perception to problem solving.
Springer, S. P., and G. Deutsch. 2001. Left brain, right brain: Perspectives from cognitive neuroscience. New York: W. H.
Freeman.
For more advanced readers, this book provides evidence from behavioral and neural imaging techniques of the laterality of mental
functions, and how specific cognitive abilities are divided between the left and the right sides of the brain.
Perception
The study of perception within CNS is concerned with how humans perceive and act upon the world. Subcategories of this field include
eye movements, object recognition, segmentation and recognition (including reading), face recognition, spatial cognition, perceptual
organization, mental imagery, and scene perception. How do we recognize everyday objects? How do we perceive depth and shape?
These are just some of the questions addressed by cognitive neuroscientists whose work focuses on perception. The study of
perception in CNS is vast, with research dedicated to diverse issues within the field. Different categories of objects are processed by
different neural mechanisms (see Downing, et al. 2006 and also Wolfe and Horowitz 2004, which is cited under Attention). Wang and
Klein 2010 and O’Craven, et al. 1997 show that the location and movement of objects affect the speed at which perception occurs.
Reynolds and Chelazzi 2004 (cited under Attention) and Levi 2008 address, through different experimental manipulations, how the
amount of perceptual information in a scene affects the ability to attend to the scene, while Grill-Spector and Kanwisher 2005
investigate the time course of perception. Chun 2003 reviews the literature on how visual scenes are learned and incorporated into
memory. Visual perception and auditory perception constitute two main aspects of perception. Farah 2000 provides an overview of the
first aspect, while Schnupp and King 2011 investigates the neural mechanisms underlying the second. Finally, Newsome, et al. 1989
compares the neural correlates of perceptual decision making in humans and monkeys.
Chun, M. 2003. Scene perception and memory. Psychology of Learning and Motivation 42:79–108.
This article reviews the literature on how information from visual scenes are learned and represented in the brain.
Downing, P. E., A. W. Chan, M. V. Peelen, C. M. Dodds, and N. Kanwisher. 2006. Domain specificity in visual cortex. Cerebral
Cortex 16.10: 1453–1461.
This article provides important evidence for regions in the visual cortex that are specialized for recognizing different categories of
objects, as is evident for faces, scenes, and bodies.
Farah, M. J. 2000. The cognitive neuroscience of vision. Malden, MA: Blackwell.

This book offers a comprehensive overview of the neural mechanism of vision perception, starting from the transformation of light to
neural images in the eye, and extending to our conscious awareness of the things that we see.
Grill-Spector, K., and N. Kanwisher. 2005. Visual recognition: As soon as you know it is there, you know what it is.
Psychological Science 16.2: 152–160.
This article addresses the timing of visual object recognition in perception, and provides unexpected evidence that, by the time a person
is aware that an object is present at all, even if they are unable to name the specific object (e.g., pigeon), they already know its category
(e.g., bird).
Levi, D. M. 2008. Crowding—an essential bottleneck for object recognition: A mini-review. Vision Research 48.5: 635–654.
Crowding refers to the phenomenon in which one’s ability to recognize stimuli is impaired when they occur in clusters. This article
provides an overview of the various theories of how crowding occurs in the brain.
Newsome, W. T., K. H. Britten, and J. A. Movshon. 1989. Neuronal correlates of a perceptual decision. Nature 341:52–54.
The behavioral performance of monkeys on a psychophysical task is compared to evidence derived from recordings from their visual
cortical neurons. Data recorded from the visual neurons of the monkeys were more sensitive and reliable that that derived from their
behavioral performance on the task.
O’Craven, K. M., B. R. Rosen, K. K. Kwong, A. Treisman, and R. L. Savoy. 1997. Voluntary attention modulates fMRI activation
in human MT/MST. Neuron 18:591–598.
This article focuses on the MT/MST (middle temporal/medial superior temporal) area in the brain, a complex that is important in
processing visual motion. MT-MST activation is higher when participants attend to moving dots versus stationary dots in a visual
stimulus.
Schnupp, J., I. Nelken, and A. King. 2011. Auditory neuroscience: Making sense of sound. Cambridge, MA: MIT Press.
The process of hearing requires an intricate perceptual process in order to make sense of sound. This book covers the neural
mechanisms in the auditory system that allows us to do this.
Wang, Z., and R. M. Klein. 2010. Searching for inhibition of return in visual search: A review. Vision Research 50.2: 220–228.
Inhibition of return (IOR) is a phenomenon in visual search in which the speed and accuracy with which an object is detected are first
briefly enhanced after the object is attended and then impaired. It is thought that the function of the IOR is to encourage orientation
toward novel items, thus aiding search. This article reviews the support for the IOR as a mechanism to aid in foraging tasks.
Attention
We are bombarded with countless bits of information in daily life. How does the brain allow us both to pay attention to the salient and
important information and to filter out unimportant, unnecessary, and distracting information? How are attentional resources used to
select visual or auditory information for perception, cognition, and the planning and production of actions? The study of attention in CNS
aims to understand the neural mechanism that underlies the cognitive process of selectively concentrating on one aspect of the
environment while ignoring others. Posner 2004 provides an excellent overview of the problems and topics within the field, as well as
in-depth discussions and explanations of various attentional processes. A number of thorough articles on the neural correlates of
attention have been published, including Corbetta and Shulman 2002, which provides a review of the evidence for different attentional
functions; Kastner and Ungerleider 2000, which provides a review of the systems involved in selective attention; Corbetta, et al. 2005
and Desimone and Duncan 1995, which provide evidence for the neural substrates of spatial and visual attention, respectively; Luck, et
al. 1989, which investigates attentional processes in split-brain patients; Wolfe and Horowitz 2004, which investigates how properties of
a stimulus affect attention; and finally Reynolds and Chelazzi 2004, which investigates the effects of visual perception on attention in
monkeys.
Corbetta, M., M. J. Kincade, C. Lewis, A. Z. Snyder, and A. Sapir. 2005. Neural basis and recovery of spatial attention deficits in
spatial neglect. Nature Neuroscience 8.11: 1603–1610.
This article investigates spatial neglect, a condition arising from damage to one hemisphere of the brain that results in a deficit in
attention to, and awareness of, one side of space; the article proposes a model of spontaneous recovery from these deficits.
Corbetta, M., and G. L. Shulman. 2002. Control of goal-directed and stimulus-driven attention in the brain. Nature Reviews
Neuroscience 3:201–215.
This article reviews the evidence for two systems of partially segregated brain areas that underlie different attentional functions.
Desimone, R., and J. Duncan. 1995. Neural mechanisms of selective visual attention. Annual Review of Neuroscience 18:193–
197.
This article reviews the two basic phenomena that define the problem of visual attention: capacity limits (at a given time, only a certain
amount of information can be processed) and selectivity (the ability to filter out unwanted information).
Kastner, S., and L. Ungerleider. 2000. Mechanisms of visual attention in the human cortex. Annual Review of Neuroscience
23:315–341.
This article provides a review of the systems involved in selective attention and biased competition between stimuli in the human visual
cortex.
Luck, S. J., S. A. Hillyard, G. R. Mangun, and M. S. Gazzaniga. 1989. Independent hemispheric attentional systems mediate
visual search in split-brain patients. Nature 342:543–545.
Split-brain patients’ left and right hemispheres are shown to have their own system for focus of attention. Split-brain patients are able to
detect bilateral stimulus arrays more quickly than normal control participants.
Posner, M. I. 2004. The cognitive neuroscience of attention. New York: Guilford.

Suited for readers of all levels, this book highlights the progress in the scientific understanding of attention and discusses the anatomy,
neurotransmitters, and development of attentional processes.
Raz, A., and J. Buhle. 2006. Typologies of attentional networks. Nature Reviews Neuroscience 7.5: 367–379.
While attention has typically been investigated as a unitary concept, research indicates that, in fact, disparate modules of attentional
networks exist.
Reynolds, J. H., and L. Chelazzi. 2004. Attentional modulation of visual processing. Annual Review of Neuroscience 27:611–
647.
Single unit recording from monkeys provides neural evidence for the processes of attention. Attention produces different effects on the
firing rates of neurons depending on whether a stimulus appears alone or with distracters.
Wolfe, J. M., and T. S. Horowitz. 2004. What attributes guide the deployment of visual attention and how do they do it? Nature
Reviews Neuroscience 5.6: 495–501.
This article reviews the research on how various properties of visual stimuli (e.g., speed, location) control the deployment of attention.
Learning
Our ability to effectively synthesize different types of information from the environment in order to acquire new skills, knowledge,
behaviors, preferences, and values as well as how the brain underlies these processes is the focus of “learning” in CNS. Associative
learning involves acquiring knowledge about relations between events or ideas. This type of learning includes instrumental conditioning
—the modification of a voluntary behavior based on both past and expected consequences of the behavior—and classical conditioning,
or a type of learning that occurs through the temporal association of two events (for example, if something that is fear provoking, such
as getting a shock, always occurs with something neutral, such as smelling a particular perfume, eventually the neutral thing (the smell
of the perfume) will result in a fear response). Gluck, et al. 2008 provides an excellent overview of the neural systems implicated in
learning. Gluck, et al. 2008 focuses to a greater extent on real-world examples of learning and behavior, making the book more suitable
for introductory student readers, while Gluck and Myers 2001 presents a more technical, computationally and biologically based review
of the current state of the field. D’Ardenne, et al. 2008 and Schulz 1998 look at the relationship between the firing of dopaminergic
neurons in specific brain areas, such as the striatum, and reward prediction. Guitart-Masip, et al. 2010 investigates how novelty
modulates the reward response in the striatum. Hampton, et al. 2007 looks not only at the mid-brain and its relation to reward prediction
but also at prefrontal cortex activation and its role in guiding decision making and behavior. Valentin, et al. 2007 similarly investigates
the role of the orbitofrontal cortex in goal-directed learning. O’Doherty, et al. 2004 provides evidence for a neural model of
reinforcement learning. Finally, Seymour, et al. 2004 explores the neural correlates of aversive conditioning, while Bray, et al. 2008
explicates the neural mechanisms of classical conditioning and their relation to decision making.
Bray, S., A. Rangel, S. Shimojo, B. Balleine, and J. P. O’Doherty. 2008. The neural mechanisms underlying the influence of
Pavlovian cues on human decision making. Journal of Neuroscience 28.22: 5861–5866.
There are neural correlates of outcome-specific transfer in the ventrolateral putamen. This indicates that choosing an action
incompatible with Pavlovian conditioning may require the inhibition of the nonselected association.
D’Ardenne, K., S. M. McClure, L. E. Nystrom, and J. D. Cohen. 2008. BOLD responses in the dopaminergic ventral tegmental
area during a classical conditioning task. Science 319.5867:1264–1267.
A seminal article showing through fMRI that dopaminergic signals in the ventral tegmental area reflect the prediction of positive reward.
Gluck, M. A., E. Mercado, and C. E. Myers. 2008. Learning and memory: From brain to behavior. New York: Worth.
This book constitutes a review of the converging studies and behavioral approaches to learning and memory in animals and humans.
Gluck, M. A., and Myers, C. E. 2001. Gateway to memory: An introduction to neural network modeling of the hippocampus and
learning. Cambridge, MA: MIT Press.
A technical-, computational-, and biological-based review.
Guitart-Masip, M., N. Bunzeck, K. E. Stephan, R. J. Dolan, and E. Duzel. 2010. Contextual novelty changes reward
representations in the striatum. Journal of Neuroscience 30.5: 1721–1726.
Novelty enhances the representation of reward in the striatum. This effect occurs even when the reward and novelty are independent.
Hampton, A. N., R. Adolphs, M. J. Tyszka, J. P. O’Doherty. 2007. Contributions of the amygdala to reward expectancy and
choice signals in human prefrontal cortex. Neuron 16; 55.4: 545–555.
The prefrontal cortex plays a major role in guiding behavior in reward learning and decision making. However, the amygdala is crucial in
establishing expected reward in the prefrontal cortex, which is then used to guide behavior.
O’Doherty, J., P. Dayan, J. Schultz, R. Deichmann, K. Friston, and R. J. Dolan. 2004. Dissociable roles of ventral and dorsal
striatum in instrumental conditioning. Science 304:452–454.
Neural evidence is presented for a model of reinforcement learning in which a “critic” learns to predict reward and an “actor” maintains
information about the reward to enable better future choices. The critic is represented in the ventral striatum, whereas the actor is
represented in the dorsal striatum.
Schultz, W. 1998. Predictive reward signal of dopamine neurons. Journal of Neurophysiology 80.1: 1–27.
The dopaminergic system is important for reward learning, labeling stimuli with appetitive value, which helps guide behavior. This article
summarizes the research on the dopaminergic system, and how it works to direct reward learning.
Seymour, B., J. O’Doherty, P. Dayan, M. Koltzenburg, A. K. Jones, R. J. Dolan, K. J. Friston, and R. S. Frackowiak. 2004.
Temporal difference models describe higher-order learning in humans. Nature 426:664–667.
The ventral striatum serves a critical role in integrating complex predictions in higher-order aversive conditioning in order to make
predictions and coordinate behavior.
Valentin, V. V., A. Dickinson, and J. P. O’Doherty. 2007. Determining the neural substrates of goal-directed learning in the
human brain. Journal of Neuroscience 27.15: 4019–4026.
The orbitofrontal cortex is implicated in the mechanism for goal-directed learning in instrumental conditioning in humans.
Long-Term Memory
Long-term memory is divided into explicit (or declarative) memory, which permits the intentional recollection of previous experiences
(e.g., remembering what you had for breakfast yesterday), and implicit (or procedural) memory, which is largely unconscious (for
example, knowing how to ride a bike). Explicit memory can be further subdivided into semantic memory, which stores concept-based
knowledge that is independent of context and personal relevance (e.g., knowing who is the vice president), and episodic memory, which
stores personal episodes (e.g., autobiographical knowledge that is context and personally relevant, for example, knowing where you
went on vacation last year). Semantic memory is thought to be dependent on brain areas that include the left inferior prefrontal cortex,
the left posterior temporal areas, and the hippocampus (although this is controversial). Episodic memory is dependent primarily on the
prefrontal cortex and the medial temporal cortex, particularly the hippocampus. An excellent example of implicit memory is priming,
wherein prior exposure to a stimulus influences later behavior toward that stimulus (e.g., watching a DVD about cats will make you
more likely to notice cats immediately afterward). Another instance of implicit memory is the acquisition of a new skill, for example,
learning to play the guitar. Skill learning can depend on the striatum, motor cortex, and cerebellum, as well as other structures. The
study of long-term memory encompasses a vast field with many areas of research. Squire and Wixted 2010 and Squire 2009 provide
excellent broad reviews of the neural systems involved. Rugg and Curran 2007 investigates the neural systems responsible for
familiarity versus recollection. Poldrack, et al. 2001 provides evidence for the interaction between two types of long-term memory,
namely, declarative and procedural memory. Uncapher and Rugg 2005 focuses on the relationship between the amount of brain
activation while a memory is being encoded and the strength of the ensuing memory. Finally, two articles provide evidence for the
effects of emotion on memory (Kroes, et al. 2010 and Schiller, et al. 2010).
Kroes, M. C., B. A. Strange, and R. J. Dolan. 2010. Beta-adrenergic blockade during memory retrieval in humans evokes a
sustained reduction of declarative emotional memory enhancement. Journal of Neuroscience 30.11: 3959–3963.
Emotional events are often better remembered than nonemotional memories due to noradrenergic modulation and amygdala activation
and this article reports evidence that blocking adrenergic receptors with propranolol during the retrieval of emotional memories results
in an inability to retrieve the memories at later times. Broader implications of this finding include the possibility for clinical interventions
after an emotional or traumatic event.
Poldrack, R. A., J. Clark, E. J. Pare-Blagoev, D. Shohamy, J. Creso-Moyano, C. Myers, and M. A. Gluck. 2001. Interactive
memory systems in the human brain. Nature 414:546–550.
This article provides evidence that a simple variation in behavioral task can alter the memory system required to perform the task from
explicit (declarative) to implicit (procedural).
Rugg, M. D., and T. Curran. 2007. Event-related potentials and recognition memory. Trends in Cognitive Sciences 11.6: 251–
257.
This article evaluates the efficacy of using event-related potentials to offer support for dual process models of memory (recognition
memory is supported by distinct retrieval processes known as familiarity and recollection).
Schiller, D., M. Monfils, C. M. Raio, D. Johnson, J. E. LeDoux, and E. A. Phelps. 2010. Preventing the return of fear in humans
using reconsolidation update mechanisms. Nature 463:49–53.
This article presents a new way to eliminate fear memories that is noninvasive and that does not require pharmacological intervention.
Old fear memories are updated with non-fearful information provided during the reconsolidation stage of memory, such that fear
responses are no longer expressed.
Squire, L. R. 2009. Memory and brain systems: 1969–2009. Journal of Neuroscience 29:12711–12716.
This article highlights the structure and organization of memory and the brain systems that support memory.
Squire, L. R., and J. Wixted. 2010. The cognitive neuroscience of human memory since H.M. Annual Review of Neuroscience
34.
A broad review of the literature since the 1950s on the neuroanatomical structures involved in memory with an emphasis on medial
temporal structures, the stages of memory processing, memory systems in the brain, short-term memory, and long-term memory
storage.
Uncapher, M. R., and M. D. Rugg. 2005. Encoding and the durability of episodic memory: A functional magnetic resonance
imaging study. Journal of Neuroscience 25.31: 7260–7267.
This study used fMRI to determine that the neural activity elicited by an event as it is encoded is related to the strength of the resulting
memory, even when all other factors are kept constant.
Short-Term Memory
Working memory is the ability to hold information in an active state (for around 20–30 seconds), as in when you hear a phone number
and need to hold the number in mind until you record it. It is thought to be dependent on the dorsolateral prefrontal cortex and the
parietal lobe. The frontal lobes are widely believed to underpin the ability to hold information in short-term, or working, memory. Osaka,
et al. 2007 provides a broad overview of the current state of knowledge regarding the neural basis of working memory. Johnson, et al.
2005; Jonides, et al. 2008; Smith and Jonides 1999; and Curtis and D’Esposito 2003 all provide excellent reviews and data supporting
the role of the prefrontal cortex in working memory. Cohen, et al. 1997 provides evidence for different brain processes involved in verbal
and visual working memory tasks, respectively. Armstrong, et al. 2009 elucidates the role of the frontal eye fields in the selection and
maintenance of spatial information in working memory. Lewis-Peacock and Postle 2008 provides evidence for the interaction between
long-term memory and working memory. Finally, Luck and Vogel 1997 investigates the capacity limits of working memory.
Armstrong, K. M., M. H. Chang, and T. Moore. 2009. Selection and maintenance of spatial information by frontal eye field
neurons. Journal of Neuroscience 29:15621–15629.
This article provides evidence from single cell recordings from frontal eye field neurons in monkeys that information that is held in
working memory can serve to override reflexive, “bottom-up” attentional capture from stimulus-driven factors.
Cohen, J. D., W. M. Perlstein, T. S. Braver, L. E. Nystrom, D. C. Noll, J. Jonides, and E. E. Smith. 1997. Temporal dynamics of
brain activation during a working memory task. Nature 386:604–608.
One of the first fMRI experiments to isolate delay period activity in a working memory study of verbal materials.
Curtis, C. E., and M. D’Esposito. 2003. Persistent activity in the prefrontal cortex during working memory. Trends in Cognitive
Sciences 7.9: 415–423.
A systematic review of how information is maintained in working memory.
Johnson, M. K., C. L. Raye, K. J. Mitchell, E. J. Greene, W. A. Cunningham, and C. A. Sanislow. 2005. Using fMRI to investigate
a component process of reflection: Prefrontal correlates of refreshing a just-activated representation. Cognitive, Affective, &
Behavioral Neuroscience 5:339–361.
This study used fMRI to investigate the functional organization of the prefrontal cortex (PFC) during a working memory task. Areas in
the left dorsolateral, anterior, and ventrolateral PFC were identified as being related to varying types of information, including visual and
auditory words, drawings, patterns, people, places, and locations.
Jonides, J., R. L. Lewis, D. E. Nee, C. A. Lustig, M. G. Berman, and K. S. Moore. 2008. The mind and brain of short-term
memory. Annual Review of Psychology 59:193–224.
A thorough review of the literature on the neural basis of working memory.
Lewis-Peacock, J., and B. R. Postle. 2008. Temporary activation of long-term memory supports working memory. Journal of
This article provides fMRI evidence that the short-term retention of information can be supported by the temporary reactivation of LTM
representations in working memory.
Luck, S. J., and E. K. Vogel. 1997. The capacity of visual working memory for features and conjunctions. Nature 390:279–281
This article provides striking behavioral and neural evidence that the capacity of working memory is three to four items (rather than the
classic seven plus/minus two).
Osaka, N., R. Logie, and M. D’Esposito. 2007. The cognitive neuroscience of working memory. New York: Oxford Univ. Press.
This book provides an excellent and broad summary of the current literature and theories regarding the neural underpinnings of working
memory, discussing numerous investigational approaches (single cell recording, neuroimaging, computational modeling).
Smith, E. E., and J. Jonides. 1999. Storage and executive processes in the frontal lobes. Science 283:1657–1661.
A review of working memory studies that focuses on the extent to which frontal lobe function is organized by the type of information and
type of processing required.
Cognitive Control
This area of CNS focuses on the neurobiological mechanisms underlying selective attention, inhibition, and other processes that allow
for the ability to flexibly adapt behavior to current demands rather than remaining rigid and inflexible. The prefrontal cortex, and in
particular the dorsolateral PFC, is thought to play a key role in supporting cognitive control in the brain. Badre and D’Esposito 2007;
Koechlin, et al. 2003; and Miller and Cohen 2001 provide evidence for the role of the prefrontal cortex in cognitive control. Depoe, et al.
2007 provides data showing that the prefrontal cortex is implemented in the suppression of emotional information. Braver, et al. 2009
provides evidence that different cognitive control mechanisms can be implemented flexibly within the same brain regions. Finally,
Carter, et al. 1998; Kerns, et al. 2004; and MacDonald, et al. 2000 show that the anterior cingulate cortex is involved in monitoring
cognitive conflict, while Young, et al. 2004 elucidates how error-related negativity relates to cognitive conflict.
Badre, D., and M. D’Esposito. 2007. Functional magnetic resonance imaging evidence for a hierarchical organization of the
prefrontal cortex. Journal of Cognitive Neuroscience 19.12: 2082–2099.
An fMRI study that supports a model in which the prefrontal cortex reflects a hierarchical order of control, based on the level of
abstraction.
Braver, T. S., J. L. Paxton, H. S. Locke, and D. M. Barch. 2009. Flexible neural mechanisms of cognitive control within human
prefrontal cortex. Proceedings of the National Academy of Sciences USA 106.18: 7351–7356.
This study provides evidence for a theory in which different cognitive control mechanisms can be implemented flexibly within the same
brain regions, rather than a model in which dissociable anatomical mechanisms underlie each process.
Carter, C. S., T. S. Braver, D. M. Barch, M. M. Botvinick, D. C. Noll, and J. D. Cohen. 1998. Anterior cingulate cortex, error
detection and the on-line monitoring of performance. Science 280:747–749.
One of the first studies to provide neural evidence that the anterior cingulate cortex (ACC) monitors cognitive conflict.
Depoe, B., T. Curran, and M. T. Banish. 2007. Prefrontal regions orchestrate suppression of emotional memories via a two-
phase process. Science 317:215–219.
This article provides some of the strongest evidence to date for the neural mechanisms behind the intentional inhibition of information.
Kerns, J. G., J. D. Cohen, A. W. MacDonald III, R. Y. Cho, V. A. Stinger, and C. S. Carter. 2004. Anterior cingulate conflict
monitoring and adjustments in control. Science 303:1023–1026.
The ACC plays a crucial role in cognitive control, as is evidenced by neural and behavioral changes.
Koechlin, E., C. Ody, and F. Kouneiher. 2003. The architecture of cognitive control in the human prefrontal cortex. Science
302.5648: 1181–1185.
An fMRI study that supports a unified modular model of cognitive control, in which the lateral prefrontal cortex is organized according to
the type of stimuli, the present perceptual context, and the temporal episode in which the stimuli occur.
MacDonald, A. W., J. D. Cohen, V. A. Stinger, and C. S. Carter. 2000. Dissociating the role of dorsolateral prefrontal cortex and
anterior cingulate cortex in cognitive control. Science 288:1835–1837.
This study shows a neural double dissociation between mechanisms that monitor conflict and mechanisms that resolve conflict.
Miller, E. K., and J. D. Cohen. 2001. An integrative theory of prefrontal cortex function. Annual Review of Neuroscience
24:167–202.
This article proposes that cognitive control stems from the maintenance of patterns of activity in the prefrontal cortex that guide the
representations and means of achieving internal goals.
Young, N., M. M. Botvinick, and J. D. Cohen. 2004. The neural basis of error detection: Conflict monitoring and the error-
related negativity. Psychological Review 111.4: 931–959.
This article develops the theory that error-related negativity (ERN), a component of the event-related potential that exhibits after an
error is made in an attentional task, can be explained by a response conflict mediated by the anterior cingulate cortex.
Decision Making
Although decision making has been an object of study for centuries, and is of great importance to a wide range of specialists
(economists, lawyers, clinicians, policymakers, etc.), how this process is mediated by the brain has only recently begun to be
understood and investigated. The prefrontal cortex, including both lateral and ventromedial areas, is thought to be important in certain
as well as uncertain decision making in humans. For readers interested in the brain structures that underlie and support decision-
making abilities, Vartanian and Mandel 2011 provides a current and broad review of the relatively nascent literature of this field. Fellows
2004 and Nieuwenhuis, et al. 2005 provide excellent reviews of the neural substrates of decision making. Research presented in Daw,
et al. 2006 and Yu and Dayan 2005 demonstrates the brain’s response to decisions made under conditions of uncertainty. De Drue, et
al. 2010 explores the role of oxytocin in the regulation of group conflict. Chib, et al. 2009 explores the brain’s role in decisions that carry
differing values. Wunderlich, et al. 2009 provides data regarding decisions made with different reward outcomes, while Symmonds, et
al. 2010 provides data about how people evaluate the potential risks and rewards of multiple sequential decisions.
Chib, V., A. Rangel, S. Shimojo, and J. P. O’Doherty. 2009. Evidence for a common representation of decision values for
dissimilar goods in human ventromedial prefrontal cortex. Journal of Neuroscience 29.39: 12315–12320.
This article addresses the issue of whether different brain areas correlate with different types of value representations (e.g., food,
nonfood consumables, and monetary gambles). fMRI results implicated the ventromedial prefrontal cortex (vmPFC) in the valuation of
all categories of goods.
Daw, N., J. P. O’Doherty, P. Dayan, B. Seymour, and R. J. Dolan. 2006. Cortical substrates for exploratory decisions in humans.
Nature 441:876–879.
This article addresses the neural substrates of decision making under uncertainty. A model of action-selection under uncertainty that
involves switching between exploratory and exploitative behavioral modes is proposed. The frontopolar cortex and intraparietal sulcus
are preferentially active during exploratory decisions, while regions of the striatum and ventromedial prefrontal cortex are active in
value-based exploitative decision making.
De Dreu, C. K. W., L. L. Greer, M. J. J. Handgraaf, S. Shalvi, G. A. Van Kleef, M. Baas, F. S. Ten Velden, E. Van Dijk, and S. W. W.
Feith. 2010. The neuropeptide oxytocin regulates parochial altruism in intergroup conflict among humans. Science 328.5984:
1408–1411.
This article provides empirical evidence for the relationship between oxytocin and the regulation of intergroup conflict, which is
imperative for survival. Oxytocin promoted in-group trust and cooperation, and aggression toward competing out-groups.
Fellows, L. K. 2004. The cognitive neuroscience of human decision making: A review and conceptual framework. Behavioral
and Cognitive Neuroscience Reviews 3.3: 159–172.
This article provides an excellent review of the current neuroscientific literature of human decision making, focusing on the role of the
frontal lobes.
Nieuwenhuis, S., G. Aston-Jones, and J. D. Cohen. 2005. Decision-making, the P3, and the locus coeruleus-norepinephrine
system. Psychological Bulletin 131.4: 510–532.
This review integrates knowledge regarding the neural basis of the P3 (an event-related potential elicited by infrequent, task-relevant
stimuli, which is thought to be an index for novelty detection) and elucidates its functional role in internal decision-making processes.
Symmonds, M., P. Bossaerts, and R. J. Dolan. 2010. A behavioral and neural evaluation of prospective decision making under
risk. Journal of Neuroscience 30.43: 14380–14389.
This article addresses the question of how people evaluate the potential risks and rewards of multiple sequential decisions, and what
brain areas underlie such decisions.
Vartanian, O., and D. R. Mandel. 2011. Neuroscience of decision making: Contemporary topics in cognitive neuroscience
series. London: Psychology Press.
An extremely useful book for readers at all levels, it provides a summary of work in the field focusing on the intersection between
decision making and cognitive neuroscience. Topics covered include the role of emotions, dual systems, reward/loss processing,
planning, and creativity.
Wunderlich, K., A. Rangel, and J. P. O’Doherty. 2009. Neural computations underlying action-based decision making in the
human brain. Proceedings of the National Academy of Sciences 106.40: 17199–17204.
Choices between different physical actions to obtain reward activate brain areas, including the supplementary motor cortex. Further, the
ventromedial prefrontal cortex was found to be involved in encoding the expected value of the action that was ultimately taken.
Yu, A. J., and P. Dayan. 2005. Uncertainty, neuromodulation, and attention. Neuron 46.4: 681–692.
This article proposes that the neuromodulators acetylcholine and norepinephrine play a major role in the brain’s response to conditions
of uncertainty.
Social Cognition
The field of social cognition within cognitive neuroscience aims to infuse the study of the relations between people and groups with
brain science methodology. Social cognitive neuroscience attempts to determine the neural pathways and mechanisms responsible for
social phenomena such as stereotyping, attitudes, self-control, prejudice, empathy, perspective-taking, theory of mind, moral reasoning,
and emotion regulation. Two of the main books in social neuroscience are Cacioppo and Berntson 2005 and Decety and Cacioppo
2011. Each provides a thorough review of the existing literature and also speaks to future directions in this rapidly advancing area of
study. A number of excellent reviews of the neural systems involved in social cognition have been published to date. These include
Amodio and Frith 2006; Cacioppo, et al. 2010; and Ochsner 2004. Siegal and Varley 2002 provides a review of the literature in the area
of theory of mind, while Saxe and Kanwisher 2003 provides evidence for the difference between the recognition of another’s mind and
the simple recognition of another. Johnson, et al. 2009 shows medial cortex activity differences between healthy and depressed
individuals as they self-reflect. Mitchell, et al. 2006 provides neural evidence for the dissociation between mentalizing about the self
versus another person. Singer, et al. 2004 explores how empathy for someone else’s pain relates to feeling the pain oneself.
Amodio, D. M., and C. D. Frith. 2006. Meeting of minds: The medial frontal cortex and social cognition. Nature Reviews
This article provides a comprehensive review of the neural mechanisms underlying social interaction, emphasizing the role of the
medial frontal cortex.
Cacioppo, J. T., and G. G. Berntson. 2005. Social neuroscience. London: Psychology Press.
This book is intended for more advanced readers. It provides a thorough, comprehensive collection of articles and reviews of the
current literature within social neuroscience.
Cacioppo, J. T., G. G. Berntson, and J. Decety. 2010. Social neuroscience and its relation to social psychology. Social
Cognition 28:675–684.
This article reviews the current literature on social interactions and behavior, with an emphasis on evidence relating to the neural,
hormonal, cellular, and genetic mechanisms underlying such abilities.
Decety, J., and J. T. Cacioppo. 2011. Handbook of social neuroscience. New York: Oxford Univ. Press.
A thorough handbook with an emphasis on the neural, hormonal, cellular, and genetic mechanisms of social cognition.
Johnson, M. K., S. Nolen-Hoeksema, K. J. Mitchell, and Y. Levin. 2009. Medial cortex activity, self-reflection, and depression.
Social Cognitive and Affective Neuroscience 4:313–327.
An fMRI study investigating medial cortex activity in depressed and healthy individuals as they self-reflect.
Mitchell, J. P., C. N. Macrae, and M. R. Banaji. 2006. Dissociable medial prefrontal contributions to judgments of similar and
dissimilar others. Neuron 50:655–663.
This article provides fMRI evidence for a double dissociation between acts of mentalizing about a similar other, which recruits an area in
ventral mPFC, and acts of mentalizing about a dissimilar other, which recruits an area in the dorsal subregion of mPFC.
Ochsner, K. N. 2004. Current directions in social cognitive neuroscience. Current Opinion in Neurobiology 14:254–258.
An excellent, encompassing review of the current psychological and neural literature in the field, including topics that are well
developed and subjects that are emerging.
Saxe, R., and N. Kanwisher. 2003. People thinking about people: The role of the temporo-parietal junction in “theory of mind.”
NeuroImage 19:1835–1842.
This study provides fMRI evidence for the dissociation between theory of mind, which represents another person’s mental states, and a
representation of the simple presence of another person per se.
Siegal, M., and R. Varley. 2002. Neural systems involved in theory of mind. Nature Reviews Neuroscience 3:462–471.
Theory of mind is the ability to understand and reason about the beliefs of others. This article provides a review from imaging and lesion
studies that implicate the involvement of a widely distributed neural system.
Singer, T., B. Seymour, J. O’Doherty, H. Kaube, R. J. Dolan, and C. D. Frith. 2004. Empathy for pain involves the affective but
not sensory components of pain. Science 303.5661: 1157–1162.
This study provides fMRI evidence that empathy for another’s pain (the ability to have an experience of another’s pain) activates
specific areas in the brain that are also activated when one experiences pain.
Language
Language appears to be a uniquely human ability, one that is profoundly important to our species. In 1861 Paul Broca localized a
language-related function in a specific area in the human brain. Since then, the attempt to uncover the neural bases and genetic
underpinnings of the acquisition and use of language, including comprehension and production, has been the subject of intense study.
Brain areas thought to be critical for these abilities include Broca’s area in the left frontal cortex (responsible for language production),
Wernike’s area in the posterior part of the temporal lobe (responsible for language processing), and areas surrounding the Sylvian
fissure and the inferior frontal gyrus (IFG) including pars opercularis, pars triangularis, pars orbitalis, and the cortex along the inferior
frontal sulcus. Pinker 1994 is essential reading for anyone interested in the question of why we possess the ability for language. Pinker
is an expert in the field, and his clear and fascinating book provides a comprehensive overview of the important questions and topics
within this field of study. Hickok and Bellugi 2001 provides evidence for the neural organization of language. The research presented in
Mechelli 2004; Sakai 2005; and Sakai, et al. 2008 deals with investigation of the brain’s role in the acquisition of a second language.
Hickok, G., and U. Bellugi. 2010. Neural organization of language: Clues from sign language aphasia. In The handbook of
psycholinguistic & cognitive processes: Perspectives in communication disorders. Edited by J. Guendouzi, F. Loncke, and M.
Williams, 685–706. London: Taylor & Francis.
This article reviews the similarities between the lateralization of the neural mechanisms for normal speakers and for aphasics (those
who are impaired in producing or comprehending spoken or written language) in both spoken and signed language. This sort of
comparison gives evidence for the importance of the sensory and motor modalities through which language is perceived and produced.
Mechelli, A., J. T. Crinion, U. Noppeney, J. O’Doherty, J. Ashburner, R. S. Frackowiak, and C. J. Price. 2004. Neurolinguistics:
Structural plasticity in the bilingual brain. Nature 431.7010: 757.
Second-language ability is accompanied by an increase in gray matter in the left inferior parietal cortex. The amount of increase
depends on the level of proficiency in that language and the age at which the language is acquired. This demonstrates the level of
plasticity in the brain in relation to language ability.
Pinker, S. 1994. The language instinct: How the mind creates language. Cambridge, MA: William Morrow.
Pinker argues that humans are born with an innate ability for language. In this book he describes how language works, how children
learn it, how it evolved, and how it is constantly changing.
Sakai, K. L. 2005. Language acquisition and brain development. Science 310:815–819.

Research using fMRI helps to show the nature that language takes in the mature brain, and how cortical plasticity allows humans to
acquire second languages.
Sakai, K. L., A. Nauchi, Y. Tatsuno, K. Hirano, Y. Muraishi, M. Kimura, M. Bostwick, and N. Yusa. 2008. Distinct roles of left
inferior frontal regions that explain individual differences in second language acquisition. Human Brain Mapping 30:2440–
2452.
A study that shows there are individual differences in second language learning; the differential acquisition of the second language is
associated with specific regions in the F3t and F3O.
Motor Control
In everyday life, our brains issue thousands of motor commands that allow our bodies to be put through a diverse range of movements,
enabling us to navigate through the world. How does the brain combine sensory information about the environment and one’s own body
movement in order to achieve adaptive sensory-motor control? The study of motor control within CNS investigates the behavioral,
neural, and mechanical mechanisms underlying the selection, planning, learning, initiation, and execution of movements ranging from
blinking to running. Bernštejn 1967, Jeannerod 1997, and Rosenbaum 2010 all provide overviews of the central themes in the cognitive
neuroscience of action and motor control. All are geared toward more advanced readers, with Rosenbaum 2010 the most suitable for
introductory students. Chen, et al. 2010 provides evidence for the key neural networks in the human motor system. The brain areas
responsible for speed versus accuracy in motor control are reviewed in Elliot, et al. 2001. Rizzolatti and Craighero 2004 elucidates how
the brain’s mirror neuron system allows for learning through imitation.
Bernštejn, N. 1967. The coordination and regulation of movements. London: Pergamon.

This book illuminates the main problems in understanding the regulation and control of human motor acts.
Chen, C. C., J. M. Kilner, K. J. Friston, S. J. Kiebel, R. K. Jolly, and N. S. Ward. 2010. Nonlinear coupling in the human motor
system. Journal of Neuroscience 30.25: 8393–8399.
This study used magnetoencephalography in humans to look for evidence for nonlinear (between-frequency) coupling among neuronal
sources thought to facilitate communication between areas within distributed networks in the motor system.
Elliot, D., W. F. Helsen, and R. Chua. 2001. A century later: Woodworth’s (1899) two component model of goal-directed aiming.
Psychological Bulletin 127:342–357.
Woodworth’s 1899 two-component model of speed-accuracy relations in the control of upper-limb movements is evaluated with regard
to current empirical and theoretical knowledge.
Jeannerod, M. 1997. The cognitive neuroscience of action. Hoboken, NJ: Wiley-Blackwell.

This book addresses the nature and role of different representations in the planning and execution of movements. The book is geared
toward advanced undergraduates or those working in the area.
Rizzolatti, G., and L. Craighero. 2004. The mirror-neuron system. Annual Review of Neuroscience 27:169–192.
Humans can learn by imitating others (observing the actions of others and copying them). This review presents data on a
neurophysiological mechanism—the mirror-neuron system—that appears to play a fundamental role in both action understanding and
imitation.
Rosenbaum, D. A. 2010. Human motor control. 2d ed. San Diego, CA: Academic Press.
This book provides a broad neurological, psychological, and theoretical background on human motor control both for an introductory
audience and for more advanced readers.
LAST MODIFIED: 11/29/2011
DOI: 10.1093/OBO/9780199828340-0015
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Gazzaniga, M. 2000. Cognitive neuroscience: A reader. Malden, MA: Blackwell.

Purves, D. 2008. Principles of cognitive neuroscience. Sunderland, MA: Sinauer.

Burlington, MA: Academic Press.

Banich, M. T. 2004. Cognitive neuroscience and neuropsychology. Florence, KY: Wadsworth.

Farah, M. J. 2000. The cognitive neuroscience of vision. Malden, MA: Blackwell.

Posner, M. I. 2004. The cognitive neuroscience of attention. New York: Guilford.

Sakai, K. L. 2005. Language acquisition and brain development. Science 310:815–819.

Bernštejn, N. 1967. The coordination and regulation of movements. London: Pergamon.

Jeannerod, M. 1997. The cognitive neuroscience of action. Hoboken, NJ: Wiley-Blackwell.

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