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The Student’s Guide to
Cognitive Neuroscience
Reflecting recent changes in the way cognition and the brain are studied, this
thoroughly updated fourth edition of this bestselling textbook provides a
comprehensive and student-friendly guide to cognitive neuroscience. Jamie
Ward provides an easy-to-follow introduction to neural structure and function,
as well as all the key methods and procedures of cognitive neuroscience, with
a view to helping students understand how they can be used to shed light on
the neural basis of cognition.
The book presents a comprehensive overview of the latest theories and
findings in all the key topics in cognitive neuroscience, including vision,
hearing, attention, memory, speech and language, numeracy, executive
function, social and emotional behavior and developmental neuroscience.
Throughout, case studies, newspaper reports, everyday examples and student-
friendly pedagogy are used to help students understand the more challenging
ideas that underpin the subject.
www.routledge.com/cw/ward
For students:
For instructors:
•• Lecture guides
• Downloadable PowerPoint teaching slides
•• Additional quiz questions and answers.
THE STUDENT’S GUIDE
TO COGNITIVE
NEUROSCIENCE
Fourth Edition
JAMIE WARD
Fourth edition published 2020
by Routledge
2 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN
and by Routledge
52 Vanderbilt Avenue, New York, NY 10017
Routledge is an imprint of the Taylor & Francis Group, an informa business
© 2020 Jamie Ward
The right of Jamie Ward to be identified as author of this work
has been asserted by him in accordance with Sections 77 and 78
of the Copyright, Designs and Patents Act 1988.
All rights reserved. No part of this book may be reprinted or
reproduced or utilised in any form or by any electronic,
mechanical, or other means, now known or hereafter invented,
including photocopying and recording, or in any information
storage or retrieval system, without permission in writing from
the publishers.
Trademark notice: Product or corporate names may be
trademarks or registered trademarks, and are used only for
identification and explanation without intent to infringe.
First edition published by Psychology Press 2006
Third edition published by Psychology Press 2013
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data
A catalog record has been requested for this book
ISBN: 978-1-138-49052-9 (hbk)
ISBN: 978-1-138-49054-3 (pbk)
ISBN: 978-1-351-03518-7 (ebk)
Typeset in Times New Roman
by Swales & Willis Ltd, Exeter, Devon, UK
Visit the companion website: www.routledge.com/cw/ward
Contents
References451
Author index 515
Subject index 517
About the author
The motivation for writing this book came out of my experiences of teaching
cognitive neuroscience. When asked by students which book they should buy,
I felt that none of the existing books would satisfactorily meet their needs.
Other books in the market were variously too encyclopedic, too advanced or
not up to date, or gave short shrift to explaining the methods of the field. My
brief for writing this textbook was to provide a text that presents key ideas
and findings but is not too long, that is up to date and that considers both
method and theory. I hope that it will be useful to both lecturers and students.
In writing a book on cognitive neuroscience I had to make a decision as
to how much would be “cognitive” and how much would be “neuroscience.”
In my opinion, the theoretical underpinnings of cognitive neuroscience lie
within the cognitive psychology tradition. Some of the most elegant studies
using methods such as fMRI and TMS have been motivated by previous
research in cognitive psychology and neuropsychology. The ultimate aim of
cognitive neuroscience is to provide a brain-based account of cognition, and
so the methods of cognitive neuroscience must necessarily speak to some
aspect of brain function. However, I believe that cognitive neuroscience has
much to learn from cognitive psychology in terms of which theoretically
interesting questions to ask.
In Chapter 1, I discuss the current status of cognitive neuroscience
as I see it. Some of the topics raised in this chapter are directly aimed at
other researchers in the field who are skeptical about the merits of the newer
methodologies. I suspect that students who are new to the field will approach
the topic with open-mindedness rather than skepticism, but I hope that they
will nevertheless be able to gain something from this debate.
Chapter 2 is intended primarily as a reference source that can be referred
back to. It is deliberately pitched at a need-to-know level.
Chapters 3 to 5 describe in detail the methods of cognitive neuroscience.
The aim of an undergraduate course in cognitive neuroscience is presumably
to enable students to critically evaluate the field and, in my opinion, this
can only be achieved if the students fully understand the limitations of the
methods on which the field is based. I also hope that these chapters will be
xii PREFACE TO THE FOURTH EDITION
of use to researchers who are starting out in the field. This fourth edition has
been updated to include the latest research tools (such as tES, transcranial
electrical stimulation) and the latest research methodology (such as multi-
voxel pattern analysis, MVPA, in fMRI research).
Chapters 6 to 16 outline the main theories and findings in the field. I hope
that they convey something of the excitement and optimism that currently
exists. This fourth edition represents a substantial update. The order of the
chapters has been changed to bring development much earlier on (as it deals
with general issues relating to brain structure and function). These chapters
were also extensively updated to take into account the rapid changes in this
field, notably the links with genetic methods and connectomics. Vision and
hearing are now consecutive chapters (Chapters 7 and 8), which link well to
the following chapters on attention and action.
The following topics have either been added for the first time or
extensively updated: network versus modular approaches (Chapter 1),
magnetoencephalography (MEG) (Chapter 3), functional near-infrared
spectroscopy (fNIRS) (Chapter 4), visual imagery (Chapter 7), parietal
lobe mechanisms of sensorimotor transformation (Chapter 10), recent
neurobiological models of speech perception and production (Chapter
12), developmental dyslexia (Chapter 13) and the neuroscience of racial
biases (Chapter 16).
In addition, we have created a demonstration library of cognitive tests
(www.testable.org/ward) with thanks to Constantin Rezlescu. Within the
textbook, we provide more guidance to web resources via new feature boxes in
the text, as well as via our dedicated webpage (www.routledge.com/cw/ward).
Jamie Ward
jamiew@sussex.ac.uk
Brighton, UK, March 2019
CH A P TE R 1
Introducing cognitive
neuroscience
CONTENTS
Between 1928 and 1947, Wilder Penfield and colleagues carried out a series
of remarkable experiments on over 400 living human brains (Penfield &
Rasmussen, 1950). The patients in question were undergoing brain surgery
for epilepsy. To identify and spare regions of the brain involved in movement
and sensation, Penfield electrically stimulated regions of the cortex while
the patient was still conscious. The procedure was not painful (the surface
of the brain does not contain pain receptors), but the patients did report
some fascinating experiences. When stimulating the occipital lobe one patient
reported, “a star came down toward my nose.” Upon stimulating a region near
the central sulcus, another patient commented, “those fingers and my thumb
gave a jump.” After temporal lobe stimulation, another patient claimed, “I
heard the music again; it is like the radio.” She was later able to recall the tune
she heard and was absolutely convinced that there must have been a radio in
the operating theatre. Of course, the patients had no idea when the electrical
stimulation was being applied—they couldn’t physically feel it or see it. As
far as they were concerned, an electrical stimulation applied to the brain felt
pretty much like a mental/cognitive event.
This book tells the emerging story of how mental processes such as
thoughts, memories and perceptions are organized and implemented by the
2 THE STUDENT’S GUIDE TO COGNITIVE NEUROSCIENCE
in vivo blood flow measured in humans, enabling PET (Reivich et al., 1979)
1980
First study of TMS reported (Barker et al., 1985)
2000
BOLD response reported enabling fMRI development (Ogawa et al., 1990)
2010 Attempt to map all major connections of the brain (Human Connectome Project)
brain. It is also concerned with how it is possible to study the mind and brain,
and how we know what we know. The term cognition collectively refers to
a variety of higher mental processes such as thinking, perceiving, imagining,
ONLINE RESOURCES speaking, acting and planning. Cognitive neuroscience is a bridging
discipline between cognitive science and cognitive psychology, on the one
To discover more
about Wilder Penfield hand, and biology and neuroscience, on the other. It has emerged as a distinct
and his pioneering enterprise only recently and has been driven by methodological advances that
research, watch the enable the study of the human brain safely in the laboratory (see Figure 1.1).
videos found on the It is perhaps not too surprising that earlier methods, such as direct electrical
companion website stimulation of the brain, failed to enter into the mainstream of research.
(www.routledge.com/
cw/ward). This chapter begins by placing a number of philosophical and scientific
approaches to the mind and brain in a historical perspective. The coverage
is selective rather than exhaustive, and students with a particular interest
K EY TERMS in these issues might want to read more deeply elsewhere (Wickens, 2015).
The chapter then provides a basic overview of the current methods used
Cognition
A variety of higher in cognitive neuroscience. A more detailed analysis and comparison of the
mental processes such different methods is provided in Chapters 3 to 5. Finally, the chapter attempts
as thinking, perceiving, to address some of the criticisms of the cognitive neuroscience approach that
imagining, speaking, have been articulated and outlines how it can move forward.
acting and planning.
Cognitive neuroscience C O G N I T I V E N E UROSCIE NCE IN HISTORICAL
Aims to explain cognitive
processes in terms of
PER S PEC T I V E
brain-based mechanisms.
Philosophical approaches to mind and brain
Mind–body problem
The problem of how a Philosophers, as well as scientists, have long been interested in how the brain
physical substance (the can create our mental world. How is it that a physical substance can give
brain) can give rise to our rise to our sensations, thoughts and emotions? This has been termed the
sensations, thoughts and
mind–body problem, although it should more properly be called the mind–
emotions (our mind).
brain problem, because it is now agreed that the brain is the key part of
Introducing cognitive neuroscience 3
the body for cognition. One position is that the mind and brain are made KEY TERMS
up of different kinds of substance, even though they may interact. This is
known as dualism, and the most famous proponent of this idea was René Dualism
The belief that mind
Descartes (1596–1650). Descartes believed that the mind was non-physical
and brain are made up
and immortal whereas the body was physical and mortal. He suggested that of different kinds of
they interact in the pineal gland, which lies at the center of the brain and substance.
is now considered part of the endocrine system. According to Descartes,
Dual-aspect theory
stimulation of the sense organs would cause vibrations in the body/brain that The belief that mind and
would be picked up in the pineal gland, and this would create a non-physical brain are two levels of
sense of awareness. There is little hope for cognitive neuroscience if dualism is description of the same
true because the methods of physical and biological sciences cannot tap into thing.
the non-physical domain (if such a thing were to exist). Reductionism
Even in Descartes’ time, there were critics of his position. One can The belief that mind-
identify a number of broad approaches to the mind–body problem that still based concepts will
have a contemporary resonance. Spinoza (1632–1677) argued that mind and eventually be replaced by
brain were two different levels of explanation for the same thing, but not two neuroscientific concepts.
different kinds of thing. This has been termed dual-aspect theory and it
remains popular with some current researchers in the field (Velmans, 2000).
An analogy can be drawn to wave–particle duality in physics, in which the
same entity (e.g., an electron) can be described both as a wave and as a particle.
An alternative approach to the mind–body problem that is endorsed
by many contemporary thinkers is reductionism (Churchland, 1995; Crick,
1994). This position states that, although cognitive, mind-based concepts (e.g.,
emotions, memories, attention) are currently useful for scientific exploration,
they will eventually be replaced by purely biological constructs (e.g., patterns
of neuronal firings, neurotransmitter release). As such, psychology will
eventually reduce to biology as we learn more and more about the brain.
Advocates of this approach note that there are many historical precedents in
which scientific constructs are abandoned when a better explanation is found.
In the seventeenth century, scientists believed that flammable materials
contained a substance, called phlogiston, which was released when burned.
This is similar to classical notions that fire was a basic element along with water,
air and earth. Eventually, this construct was replaced by an understanding
of how chemicals combine with oxygen. The process of burning became
just one example (along with rusting) of this particular chemical reaction.
Reductionists believe that mind-based concepts, and conscious experiences
in particular, will have the same status as phlogiston in a future theory of the
brain. Those who favor dual-aspect theory over reductionism point out that
an emotion would still feel like an emotion even if we were to fully understand
its neural basis and, as such, the usefulness of cognitive, mind-based concepts
will never be fully replaced.
the heart rather than the brain. He believed that the brain acted as a coolant
system: the higher the intellect, the larger the cooling system needed. In the
Roman age, Galen (circa ad 129–199) observed brain injury in gladiators and
noted that nerves project to and from the brain. Nonetheless, he believed that
mental experiences themselves resided in the ventricles of the brain. This idea
went essentially unchallenged for well over 1,500 years. For example, when
Vesalius (1514–1564), the father of modern anatomy, published his plates
of dissected brains, the ventricles were drawn in exacting detail, whereas the
cortex was drawn crudely and schematically (see Figure 1.2). Others followed
in this tradition, often drawing the surface of the brain like the intestines. This
situation probably reflected a lack of interest in the cortex rather than a lack of
penmanship. It is not until one looks at the drawings of Gall and Spurzheim
(1810) that the features of the brain become recognizable to modern eyes.
FIGURE 1.2: Drawings of the brain from Vesalius (1543) (top), de Viessens (1685) (bottom left) and Gall and Spurzheim
(1810) (bottom right). Note how the earlier two drawings emphasized the ventricles and/or misrepresented the cortical surface.
Introducing cognitive neuroscience 5
Gall (1758–1828) and Spurzheim (1776–1832) received a bad press, KEY TERMS
historically speaking, because of their invention and advocacy of phrenology.
Phrenology had two key assumptions: first, that different regions of the brain Phrenology
The failed idea that
perform different functions and are associated with different behaviors; and
individual differences in
second, that the size of these regions produces distortions of the skull and cognition can be mapped
correlates with individual differences in cognition and personality. Taking onto differences in skull
these two ideas in turn, the notion of functional specialization within the shape.
brain has effectively endured into modern cognitive neuroscience, having Functional specialization
seen off a number of challenges over the years (Flourens, 1824; Lashley, Different regions of the
1929). The observations of Penfield and co-workers on the electrically brain are specialized for
stimulated brain provide some striking examples of this principle. However, different functions.
the functional specializations of phrenology were not based on controlled
experiments and were not constrained by theories of cognition. For example,
Fowler’s famous phrenologist’s head had regions dedicated to “parental
love,” “destructiveness” and “firmness” (Figure 1.3). Moreover, skull shape
has nothing to do with cognitive function.
Although phrenology was fatally flawed, the basic idea of different FIGURE 1.3: The
parts of the brain serving different functions paved the way for future phrenologist’s head was
developments in the nineteenth century, the most notable of which are Broca’s used to represent the
(1861) reports of two brain-damaged patients. Broca documented two cases hypothetical functions of
different regions of the brain.
in which acquired brain damage had impaired the ability to speak but left
© Photos.com/Thinkstock
other aspects of cognition relatively intact. He
concluded that language could be localized to
a particular region of the brain. Subsequent
studies argued that language itself was not a
single entity but could be further subdivided
into speech recognition, speech production
and conceptual knowledge (Lichtheim, 1885;
Wernicke, 1874). This was motivated by the
observation that brain damage can lead either
to poor speech comprehension and good
production, or good speech comprehension
and poor production (see Chapter 12 for full
details). This suggests that there are at least
two speech faculties in the brain and that
each can be independently impaired by brain
damage. This body of work was a huge step
forward in terms of thinking about mind
and brain. First, empirical observations were
being used to determine the building blocks
of cognition (is language a single module?)
rather than listing them from first principles.
Second, and related, they were developing
models of cognition that did not make direct
reference to the brain. That is, one could infer
that speech recognition and production were
separable without necessarily knowing where
in the brain they were located, or how the
underlying neurons brought these processes
6 THE STUDENT’S GUIDE TO COGNITIVE NEUROSCIENCE
K EY TERMS about. The approach of using patients with acquired brain damage to inform
theories of normal cognition is called cognitive neuropsychology and
Cognitive remains influential today (Chapter 5 discusses the logic of this method in
neuropsychology
detail). Cognitive neuropsychology is now effectively subsumed within the
The study of brain-
damaged patients to term “cognitive neuroscience,” where the latter phrase is seen as being less
inform theories of normal restrictive in terms of methodology.
cognition. Whereas discoveries in the neurosciences continued apace throughout
Information processing
the nineteenth and twentieth centuries, the formation of psychology as a
An approach in which discipline at the end of the nineteenth century took the study of the mind
behavior is described in away from its biological underpinnings. This did not reflect a belief in
terms of a sequence of dualism. It was due, in part, to some pragmatic constraints. Early pioneers
cognitive stages. of psychology, such as William James and Sigmund Freud, were interested
in topics like consciousness, attention and personality. Neuroscience has had
virtually nothing to say about these issues until quite recently. Another reason
for the schism between psychology and biology lies in the notion that one can
develop coherent and testable theories of cognition that do not make claims
about the brain. The modern foundations of cognitive psychology lie in the
computer metaphor of the brain and the information-processing approach,
popular from the 1950s onwards. For example, Broadbent (1958) argued
that much of cognition consists of a sequence of processing stages. In his
simple model, perceptual processes occur, followed by attentional processes
that transfer information to short-term memory and thence to long-term
memory (see also Atkinson & Shiffrin, 1968). These were often drawn as
a series of box-and-arrow diagrams (e.g., Figure 1.4). The implication was
that one could understand the cognitive system in the same way as one
could understand the series of steps performed by a computer program, and KEY TERMS
without reference to the brain.
The notion that the brain contains different regions of functional Modularity
The notion that certain
specialization has been around in various guises for 200 years. However,
cognitive processes (or
one particular variation on this theme has attracted particular attention regions of the brain) are
and controversy—namely Fodor’s (1983, 1998) theory of modularity. restricted in the type of
First, Fodor makes a distinction between two different classes of cognitive information they process.
process: central systems and modules. The key difference between them Domain specificity
relates to the types of information they can process. Modules are held to The idea that a cognitive
demonstrate domain specificity in that they process only one particular process (or brain region)
type of information (e.g., color, shape, words, faces), whereas central is dedicated solely to
systems are held to be domain independent in that the type of information one particular type of
processed is non-specific (candidates would be memory, attention, executive information (e.g., colors,
faces, words).
functions). According to Fodor, one advantage of modular systems is that,
by processing only a limited type of information, they can operate rapidly, Interactivity
efficiently and in isolation from other cognitive systems. An additional Later stages of processing
can begin before earlier
claim is that modules may be innately specified in the genetic code. Many
stages are complete.
of these ideas have been criticized on empirical and theoretical grounds. For
example, it has been suggested that domain specificity is not innate, although Top-down processing
The influence of later stag-
the means of acquiring it could be (Karmiloff-Smith, 1992). Moreover,
es on the processing of
systems like reading appear modular in some respects but cannot be innate earlier ones (e.g., memory
because they are recent in evolution. Others have argued that evidence for influences on perception).
interactivity suggests that modules are not isolated from other cognitive
Bottom-up processing
processes (Farah, 1994).
The passage of informa-
The idea of the mind as a computer program has advanced over the tion from simpler (e.g.,
years along with advances in computational science. For example, many edges) to more complex
cognitive models contain some element of interactivity and parallel (e.g., objects).
processing. Interactivity refers to the fact that stages in processing may Parallel processing
not be strictly separate and that later stages can begin before earlier Different information is
stages are complete. Moreover, later stages can influence the outcome of processed at the same
early ones (top-down processing, in contrast to bottom-up processing). time (i.e., in parallel).
Parallel processing refers to the fact that lots of different information Neural network models
can be processed simultaneously (by contrast, serial computers process Computational models
each piece of information one at a time). Although these computationally in which information
explicit models are more sophisticated than earlier box-and-arrow processing occurs using
diagrams, they, like their predecessors, do not always make contact with many interconnected
nodes.
the neuroscience literature.
In the 1980s, powerful computers became widely accessible as never before. This enabled cognitive
psychologists to develop computationally explicit models of cognition (that literally calculate a set of
outputs given a set of inputs) rather than the computationally inspired, but underspecified, box-and-
arrow approach. One particular way of implementing computational models has been very influential;
namely the neural network, connectionist or parallel distributed processing (PDP) approach
(McClelland et al., 1986). These models are considered in a number of places throughout this book,
notably in the chapters dealing with memory, speaking and literacy.
8 THE STUDENT’S GUIDE TO COGNITIVE NEUROSCIENCE
Connectionist models have a number of architectural features. First, they are composed of arrays
of simple information-carrying units called nodes. Nodes are information-carrying in the sense
that they respond to a particular set of inputs (e.g., certain letters, certain sounds) and produce a
restricted set of outputs. The responsiveness of a node depends on how strongly it is connected
to other nodes in the network (the “weight” of the connection) and how active the other nodes
are. It is possible to calculate, mathematically, what the output of any node would be, given a set
of input activations and a set of weights. There are a number of advantages to this type of model.
For example, by adjusting the weights over time as a result of experience, the model can develop
and learn. The parallel processing enables large amounts of data to be processed simultaneously.
A more controversial claim is that they have “neural plausibility.” Nodes, activation and weights
are in many ways analogous to neurons, firing rates and neural connectivity, respectively. However,
these models have been criticized for being too powerful in that they can learn many things that
real brains cannot (Pinker & Prince, 1988). A more moderate view is that connectionist models
provide examples of ways in which the brain might implement a given cognitive function, and
they generate new predictions that can then be tested. Whether or not the brain actually does
implement cognition in that particular way will ultimately be a question for empirical research in
cognitive neuroscience.
4 Naturally
MEG & ERP Functional MRI occurring
3 PET
lesions
Brain
2
Spatial resolution (log mm)
Map 1 TMS
Column 0
Multi-unit
Layer –1 recording
Neuron
–2
Dendrite
–3 Single-cell
recording
Synapse
–4
–3 –2 –1 0 1 2 3 4 5 6 7
Millisecond Second Minute Hour Day
Temporal resolution (log seconds)
FIGURE 1.5: The methods of cognitive neuroscience can be categorized according to their
spatial and temporal resolution.
Adapted from Churchland and Sejnowski, 1988.
physiological changes associated with blood supply to the brain, which evolve
more slowly over time. These are called hemodynamic methods and are
considered in Chapter 4.
The methods of cognitive neuroscience can be placed on a number of
dimensions (see Figure 1.5):
KEY TERM
•• The temporal resolution refers to the accuracy with which one can
measure when an event is occurring. The effects of brain damage are Temporal resolution
permanent and so this has no temporal resolution as such. Methods The accuracy with which
one can measure when
such as EEG, MEG, TMS and single-cell recording have millisecond
an event (e.g., a physio-
resolution. fMRI has a temporal resolution of several seconds that logical change) occurs.
reflects the slower hemodynamic response.
K EY TERM •• The spatial resolution refers to the accuracy with which one can measure
where an event is occurring. Lesion and functional imaging methods
Spatial resolution have comparable resolution at the millimeter level, whereas single-cell
The accuracy with
recordings have spatial resolution at the level of the neuron.
which one can measure
where an event (e.g., a •• The invasiveness of a method refers to whether the equipment is located
physiological change) is internally or externally. PET is invasive because it requires an injection of
occurring. a radio-labeled isotope. Single-cell recordings are performed on the brain
itself and are normally only carried out in non-human animals.
measure is to present the same word twice in the same case (e.g., RADIO-
RADIO) or different case (e.g., radio-RADIO) and compare this with
situations in which the word differs (e.g., mouse-RADIO, MOUSE-RADIO).
One general finding in reaction-time studies is that it is faster to process
a stimulus if the same stimulus has recently been presented. For example,
if asked to make a speeded decision about RADIO (e.g., is it animate or
inanimate?), performance will be faster if it has been previously encountered.
Dehaene et al. (2001) investigated this mechanism by comparing reaction-time
measures with functional imaging (fMRI) measures. In this task, the first word
in each pair was presented very briefly and was followed by visual noise. This
prevented the participants from consciously perceiving it and, hence, one can
be sure that they are not saying the word. The second word was consciously
seen and requires a response. Dehaene et al. found that reaction times were FIGURE 1.7: Both reaction
faster to the second word when it follows the same word, irrespective of case. times and fMRI activation
Importantly, there was a region in the left fusiform cortex that shows the in the left fusiform region
same effect (although in terms of “activation” rather than response time). demonstrate more efficient
This is shown in Figure 1.7. In this concrete example, it is meaningless to processing of words if they
are preceded by subliminal
argue that one type of measure is “better” for informing cognitive theory
presentation of the same
(to return to Coltheart’s question) given that both are measuring different word, irrespective of case.
aspects of the same event. One could explore the nature of this effect further Adapted from Dehaene et al.,
by, for instance, presenting the same word in different languages (in bilingual 2001.
12 THE STUDENT’S GUIDE TO COGNITIVE NEUROSCIENCE
speakers), presenting the words in different locations on the screen and so on.
This would provide further insights into the nature of this mechanism (e.g.,
what aspects of vision does it entail? Does it depend on word meaning?).
However, both reaction-time measures and brain-based measures could be
potentially informative. It is not the case that functional imaging is merely
telling us where cognition is happening and not how it is happening.
Another distinction that has been used to contrast cognitive psychology
and cognitive neuroscience is that between software and hardware,
respectively (Coltheart, 2004b; Harley, 2004). This derives from the familiar
computer analogy in which one can, supposedly, learn about information
processing (software) without knowing about the brain (hardware). As has
been shown, to some extent this is true. But the computer analogy is a little
misleading. Computer software is written by computer programmers (who,
incidentally, have human brains). However, information processing is not
written by some third person and then inscribed into the brain. Rather, the
brain provides causal constraints on the nature of information processing.
This is not analogous to the computer domain in which the link between
software and hardware is arbitrarily determined by a computer programmer.
To give a simple example, one model of visual word recognition suggests
that words are recognized by searching words in a mental dictionary one
by one until a match is found (Forster, 1976). The weight of evidence from
cognitive psychology argues against this serial search, and in favor of words
being searched in parallel (i.e., all candidate words are considered at the same
time). But why does human cognition work like this? Computer programs can
be made to recognize words adequately with both serial search and parallel
search. The reason why human information processing uses a parallel search
and not a serial search probably lies in the relatively slow neural response
time (acting against serial search). This constraint does not apply to the fast
processing of computers. Thus, cognitive psychology may be sufficient to
tell us the structure of information processing but it may not answer deeper
questions about why information processing should be configured in that
particular way. The biological constraints imposed by the brain shape the
nature and limitations of cognition.
F RO M M O D ULES T O N ET WO R KS
What does the future of cognitive neuroscience look like? Although nobody
knows for sure, much current research is centered on understanding the mind
and brain as a network. A network is a dynamically changing pattern of
activity over several brain regions. Rather than thinking of the brain as a
single network, there might be a multitude of different networks which are,
themselves, switched on or off depending on the kind of thought or behavior KEY TERM
that is needed. Thus, not only do brain regions have a degree of functional Connectome
specialization, but entire networks may also have some specializations. A comprehensive map
This network approach is exemplified by current efforts to map the human of neural connections
in the brain that may be
connectome (Sporns, 2011). The Human Connectome Project was launched
thought of as its “wiring
in 2010 at a cost of over $40M. The aim is to try to map out the pattern diagram.”
of connectivity in the human brain at a macro, i.e., millimeter, scale (rather
14 THE STUDENT’S GUIDE TO COGNITIVE NEUROSCIENCE
than the micro level of individual synapses). The project is based on MRI
techniques that measure structural connectivity (essentially white matter
fibers) and functional connectivity (essentially correlated patterns of brain
activity between regions). By scanning and testing thousands of people it will
ONLINE RESOURCES be possible to identify differences in the connectome that are linked to disease
Delve deeper and also, of particular relevance here, to understand how these networks
into the Human support cognitive function. This can be done by, for instance, linking
Connectome Project individual differences in the connectome to individual differences in specific
(humanconnectome.
cognitive abilities (Barch et al., 2013). Thus, it is not just an enterprise for
org) and watch TEDx
talks on connectomics biologists and neuroimagers—it also requires the input of psychologists who
by Jeff Lichtman and understand cognition. A complementary approach is to map the connectome
David van Essen. Visit at the micro scale of individual synapses. This is a daunting prospect as there
the companion website are 1010 neurons linked by 1014 synaptic connections (Azevedo et al., 2009).
at www.routledge.
By comparison, the size of a human genome is far smaller (3×109). Aside
com/cw/ward.
from the sheer scale of this challenge, there is no obvious way of interpreting
the connectome “code” (unlike the genome where there is a simple mapping
between the code and the proteins they create).
K EY TERM Of course, networks are nothing new. Networks were there from the start
Graph theory in the form of black-box-and-arrow diagrams. However, the contemporary
A mathematical tech- and emerging landscape looks very different from this. Firstly, the network
nique for computing the architecture that supports cognition is derived from biologically based
pattern of connectivity (or observations of the structural and functional connectome. This is supported
“wiring diagram”) from a by advanced mathematical tools such as graph theory (Bullmore & Sporns,
set of correlations.
2009). This essentially creates a wiring diagram, rather like a subway map,
in which some brain regions act as central hubs within the network (where
several subway lines cross, in that analogy) and other regions are less
connected (the suburbs, in that analogy). Secondly, there has been a shift
away from conceptualizing the hubs in the network as highly specialized
units. Instead, brain regions might perform a range of different functions
depending on which other parts of the brain they are communicating with.
A good example is Broca’s region itself which, whilst everyone agrees it is
important for language, seems to also be involved in other cognitive processes
such as detecting musical violations (Koelsch et al., 2006).
Does this mean that the era of functional specialization, stretching from
phrenology through to Broca and Penfield, is now over? This is certainly
not the case. It has even been argued on first principles that if the brain is a
network then the different hubs in the network must have different functional
specializations (Sporns & Betzel, 2016), except in the hypothetical scenario
that all regions in the network connect equally strongly to each other (in which
case each hub is identical). However, the function assigned to a region may
be harder to map onto simple cognitive concepts in this new framework. For
instance, the function of a brain region may be something like “integrating
vision and speech” rather than “a store of words.”
Thus, the central challenge for cognitive neuroscience for the future is
to develop new ways of describing the relationship between brain structure
(notably connectomics) and function (i.e., cognition and behavior). Barrett
and Satpute (2013) offer a useful summary of three different approaches as
shown in Figure 1.9. In the first scenario (a), there is a very simple one-to-one
mapping between different brain regions and different cognitive functions.
Introducing cognitive neuroscience 15
FIGURE 1.9: Three different ways in which different brain structures might be mapped to
different functions (tasks and processes). In (a) there is a one-to-one association between
brain structure and function whereas in both (b) and (c) a network of regions may make
different contributions to a given function. In (b) the network consists of specialized units
that interact, but in (c) the network consists of interactions between nonspecialized units.
From Barrett and Satpute (2013).
16 THE STUDENT’S GUIDE TO COGNITIVE NEUROSCIENCE
Few researchers would endorse such a view. In the second scenario (b), there
is still a high degree of specialization of brain regions but multiple regions
need to interact to generate a cognitive function. In the third scenario (c)
there is far less specialization of regions and cognitive functions are generated
by the interaction of multiple networks (with each network having some
specialization). Barrett and Satpute (2013) favor this third option, although
others argue that the cognitive architecture of the brain is more like the
second option (Vytal & Hamann, 2010).
III.
I.
Atticus, the friend of everybody, brought them together. It was
about the year 700, a short time after Cicero’s return from exile, and
in the midst of the troubles stirred up by Clodius, one of those vulgar
agitators like Catiline, by whose means Caesar exhausted the
strength of the Roman aristocracy that he might one day overcome it
more easily. Cicero and Brutus occupied at that time very different
positions in the republic. Cicero had filled the highest offices, and in
them had rendered eminent services. His talents and his probity
made him a valuable auxiliary for the aristocratical party to which he
was attached; he was not without influence with the people whom his
eloquence charmed; the provinces loved him, as they had seen him
more than once defend their interests against greedy governors, and
still more recently Italy had shown her affection by carrying him in
triumph from Brundusium to Rome. Brutus was only thirty-one; a
great part of his life had been passed away from Rome, at Athens,
where we know that he devoted himself earnestly to the study of
Greek philosophy, in Cyprus, and in the East, where he had followed
Cato. He had not yet filled any of those offices which gave political
importance, and he had to wait more than ten years before thinking
of the consulship. Nevertheless Brutus was already an important
person. In his early relations with Cicero, notwithstanding the
distance that age and official position set between them, it is Cicero
who makes the advances, who treats Brutus with consideration, and
who seeks his friendship. One would say that this young man had
given rise to singular expectations, and that it was already vaguely
felt that he was destined for great things. While Cicero was in Cilicia,
Atticus, pressing him to do justice to certain claims of Brutus, said:
“It would be something if you only brought back his friendship from
that province.”[321] And Cicero wrote of him at the same period: “He
is already the first among the young men; he will soon be, I hope, the
first in the city.”[322]
Everything in fact seemed to promise a splendid future for Brutus.
A descendant of one of the most illustrious families of Rome, the
nephew of Cato, the brother-in-law of Cassius and Lepidus, he had
just married one of the daughters of Appius Claudius; another was
already married to Pompey’s eldest son. By these alliances he was
connected on all sides with the most influential families; but his
character and manners distinguished him even more than his birth.
His youth had been austere: he had studied philosophy, not merely
as a dilettante and as being a most useful discipline for the mind, but
like a wise man who wishes to apply the lessons that it gives. He had
returned from Athens with a great reputation for wisdom, which his
virtuous and regular life confirmed. The admiration that his virtue
excited was redoubled when his surroundings and the detestable
examples he had resisted were considered. His mother Servilia had
been the object of one of the most violent passions of Caesar,
perhaps his first love. She always held a great sway over him, and
took advantage of it to enrich herself after Pharsalia by getting the
property of the conquered awarded to her. When she became old,
and felt the powerful dictator slipping from her, in order to continue
to rule him, she favoured, it is said, his amours with one of her
daughters, the wife of Cassius. The daughter who had married
Lepidus, had no better reputation, and Cicero tells a merry tale about
her. A young Roman fop, C. Vedius, going through Cilicia with a
great train, found it convenient to leave part of his baggage with one
of his hosts. Unfortunately this host died; seals were put on the
traveller’s baggage along with the rest, and to begin with, the
portraits of five great ladies were found in it, and among them that of
Brutus’ sister. “It must be admitted,” said Cicero, who did not lose an
opportunity for a joke, “that the brother and the husband well
deserve their names. The brother is very stupid (brutus) who
perceives nothing, and the husband very easy-going (lepidus) who
endures all without complaint.”[323] Such was the family of Brutus. As
to his friends, there is no need to say much about them. We know
how the rich young men of Rome lived at that time, and what
Caelius, Curio, and Dolabella were. Among all this dissipation the
rigid integrity of Brutus, his application to business, that disdain for
pleasures, that taste for study to which his pale and serious
countenance bore witness, stood out in higher relief by the contrast.
Accordingly all eyes were fixed on this grave young man who
resembled the others so little. In approaching him men could not
help a feeling that seemed ill suited to his age: he inspired respect.
Even those who were his elders and his superiors, Cicero and Caesar,
notwithstanding their glory, Antony who resembled him so little, his
opponents and his enemies, could not escape this impression in his
presence. What is most surprising is, that it has survived him. It has
been felt in presence of his memory as it was before his person; living
and dead he has commanded respect. The official historians of the
empire, Dio, who has so roughly handled Cicero, Velleius, the
flatterer of Tiberius, all have respected Brutus. It seems that political
rancour, the wish to flatter, and the violence of party have felt
themselves disarmed before this austere figure.
While respecting him, they loved him. These are sentiments which
do not always go together. Aristotle forbids us to represent heroes
perfect in all points in the drama, lest they should not interest the
public. Things go in ordinary life very much as they do on the stage; a
sort of instinctive dread holds us aloof from irreproachable
characters, and as it is usually by our common failings that we are
drawn together, we feel very little attraction towards a man who has
no failings, and are content to respect perfection at a distance. Yet it
was not so in the case of Brutus, and Cicero could say of him with
truth in one of the works that he addresses to him: “Who was ever
more respected and loved than you?”[324] And yet this man without
weaknesses was weak for those he loved. His mother and sisters had
great influence over him, and made him commit more than one fault.
He had many friends, and Cicero blames him for listening too readily
to their advice; they were worthy men who understood nothing of
public affairs; but Brutus was so much attached to them that he
could not protect himself against them. His last sorrow at Philippi
was to learn of the death of Flavius, his overseer of works, and that of
Labeo, his lieutenant; he forgot his own self to weep for them. His
last words before his death were to congratulate himself that none of
his friends had betrayed him: this fidelity, so rare at that time,
consoled his last moments. His legions also, although they were
partly composed of old soldiers of Caesar, and he kept them tightly in
hand, punishing plunderers and marauders, his very legions loved
him and remained faithful to him. Even the people of Rome
themselves, who were in general enemies of the cause he defended,
showed their sympathy for him more than once. When Octavius
proclaimed the assassins of Caesar public enemies, every one sadly
bowed their heads when they heard the name of Brutus pronounced
from the rostrum, and in the midst of the terrified senate which
foresaw the proscriptions, a voice dared to declare that it would
never condemn Brutus.
Cicero fell under the charm like the rest, but not without
resistance. His friendship with Brutus was troubled and stormy, and,
notwithstanding the general agreement of their opinions, violent
dissensions arose between them more than once. Their
disagreements are explained by the diversity of their characters.
Never did two friends resemble each other less. There never was a
man who seemed made for society more than Cicero; he brought into
it all the qualities that are necessary for success, great flexibility of
opinion, much toleration for others, allowance enough for himself,
the talent of steering with ease between all parties, and a certain
natural indulgence that made him understand and almost accept
everything. Although he made bad verses, he had the temperament
of a poet, a strange mobility of impression, an irritable sensitiveness,
a supple, broad, and quick intellect which conceived promptly, but
quickly abandoned its ideas, and passed from one extreme to another
at a bound. He did not make a single serious resolution of which he
did not repent the next day. Whenever he joined a party he was only
quick and decided at the beginning, and gradually cooled down.
Brutus, on the contrary, had not a quick intelligence: he usually
hesitated at the commencement of an enterprise and did not decide
at once. Slow and serious, he advanced step by step in everything,
but, once resolved, he was so absorbed in his conviction that nothing
could divert him: he isolated and concentrated himself in it, he
excited and inflamed himself for it by reflection, and at last listened
only to that inflexible logic that drove him to realize his purpose. He
was one of those minds of which Saint-Simon says that they have an
almost ferocious consistency. His obstinacy was the real source of his
strength, and Caesar well understood it when he said of him: “All
that he wills he means.”[325]
Two friends who resembled each other so little must naturally
have clashed at every opportunity. Their first differences were
literary. It was a custom then at the bar to divide an important case
among several orators; each took the part best suited to his talents.
Cicero, obliged to appear often before the judges, went with his
friends and pupils, and gave out to them part of his work in order to
be able to get through it. He was often satisfied with keeping the
peroration for himself, in which his copious and impassioned
eloquence was at home, and left them the rest. It was thus that
Brutus pleaded at his side and under his direction. Brutus, however,
was not of his school: a fanatical admirer of Demosthenes, whose
statue he had placed among those of his ancestors, and nurtured on
the study of the Attic masters, he sought to reproduce their graceful
severity and vigorous strength. Tacitus says that his efforts were not
always happy: by dint of avoiding ornament and pathos he was dull
and cold, and by too eagerly seeking precision and strength he
became dry and stiff. These faults were repugnant to Cicero, who
always saw in this type of eloquence, which was founding a school, a
criticism of his own, and tried by every means to convert Brutus; but
he did not succeed, and on this point they never agreed. After the
death of Caesar, and when something else than literary discussions
was in question, Brutus sent his friend the speech he had just
delivered in the Capitol, and begged him to correct it. Cicero took
good care not to do so: he knew too well by experience the self-
esteem of the literary man to run the risk of offending Brutus by
trying to do better than he. Besides, the speech seemed to him very
fine, and he wrote to Atticus that nothing could have been more
graceful or better written. “Yet,” added he, “if I had had to make it I
should have put more passion in it.”[326] Assuredly Brutus did not
lack passion, but it was in him a secret and repressed flame that only
touched the nearest, and he disliked to give the rein to those
powerful emotions and that fiery pathos without which one cannot
carry away the multitude.
He was not then a docile follower of Cicero, and we may add that
neither was he an accommodating friend. He lacked pliancy in his
relations with others, and his tone was always rough and abrupt. At
the commencement of their intercourse, Cicero, accustomed to be
treated with great respect even by the highest personages, thought
the letters of this young man were curt and haughty, and felt hurt.
This was not the only complaint he had to make of him. We know the
great consular’s irritable, suspicious and exacting vanity; we know to
what a degree he loved praise; he gave it to himself liberally and he
expected it from others, and if they were slow in giving it he was not
ashamed to ask for it. His friends were generally indulgent to this
harmless failing, and did not wait to be invited by him to praise him.
Brutus alone resisted; he prided himself on his candour, and spoke
out what he thought. Accordingly Cicero often complains that he was
chary of his praises; one day indeed he was seriously angry with him.
It was a question of the great consulship, and of the discussion in
consequence of which Lentulus and the accomplices of Catiline were
executed. This was the most vigorous action of Cicero’s life, and he
had a right to be proud of it, since he had paid for it with exile. In the
narrative that Brutus gave of the events of this day he depreciated
the part that Cicero had played in them to the advantage of his uncle
Cato. He only praised him for having punished the conspiracy,
without saying that he had discovered it, and contented himself with
calling him an excellent consul. “Poor praise!” said Cicero angrily;
“one would think it came from an enemy.”[327] But those were only
small differences arising from wounded self-esteem, which might
easily be made up; we must now mention a graver disagreement that
deserves to be dwelt upon, for it suggests some serious reflections on
the Roman society of that period.
In 702, that is to say, a short time after the commencement of his
relations with Brutus, Cicero went out as proconsul to Cilicia. He had
not sought this office, for he knew what difficulties he should find in
it. He set out decided to do his duty, and he could not do it without
bringing on his hands at the same time the patricians, his protectors,
and the knights, his protégés and clients. In fact, patricians and
knights, usually enemies, agreed, with a singular unanimity, in
plundering the provinces. The knights, farmers of the public revenue,
had only one thought: to make a fortune in five years, the usual
duration of their contract. Consequently they exacted without mercy
the tax of a tenth on the productions of the soil, a twentieth on
merchandise at the ports, the harbour dues, the tax on pasture-lands
in the interior; in fact, all the tribute that Rome had imposed on the
conquered nations. Their greed respected nothing. Livy wrote this
terrible sentence about them: “Wherever the ‘publican’ penetrates,
there is no more justice or liberty for any one.”[328] It was very
difficult for the wretched cities to satisfy these insatiable financiers;
almost everywhere the municipal coffers, ill administered by
incompetent, or pillaged by dishonest magistrates, were empty.
Money, however, had to be found at any price. Now, of whom could
they borrow it, except of the bankers of Rome, who had been for a
century the bankers of the whole world? It was to them therefore that
they applied. Some were rich enough to draw from their private
fortune money to lend to foreign cities, or sovereigns, like that
Rabirius Postumus, for whom Cicero had pleaded, who furnished the
king of Egypt with the money necessary to reconquer his kingdom.
Others, in order to run less risk, formed financial companies, in
which the most illustrious Romans invested their funds. Thus,
Pompey had a share for a considerable sum in one of those joint-
stock companies founded by Cluvius of Puteoli. All these money-
lenders, whether private individuals or companies, knights or
patricians, were very unscrupulous, and only advanced their money
at enormous interest, generally 4 or 5 per cent. per month. Their
difficulty consisted in getting paid. As it is only men who are quite
ruined who accept these hard terms, the money lent on such high
interest is always subject to risk. When the date of payment arrived
the poor city was less than ever in a position to pay: it employed a
thousand pettifogging tricks, spoke of complaining to the senate, and
began by appealing to the proconsul. Unfortunately for it, the
proconsul was usually an accomplice of its enemies, and took his
share of their profits. The creditors who had secured his co-operation
by paying him well, had then only to send into the province some
freedman or agent who represented them; the proconsul placed the
public forces at the service of private interests, gave this agent the
title of his lieutenant, some soldiers, and full powers, and if the
insolvent town did not quickly come to some satisfactory
arrangement, it suffered the horrors of a siege and of official pillage
in time of peace. The proconsul who refused to lend himself to these
abuses, and who intended, according to Cicero’s expression, to
prevent the provinces perishing, naturally aroused the anger of all
those who lived by the ruin of the provinces. The knights, the nobles,
who no longer got their money, became his deadly enemies. It is true
he had the gratitude of his province, but this did not amount to
much. It had been remarked that, in those Eastern countries “trained
by a long servitude to loathsome flattery,”[329] the governors who
received most adulation, and to whom they raised most statues were
precisely those who had robbed the most, because they were the
most dreaded. Cicero’s predecessor had completely ruined Cilicia:
consequently they thought of building him a temple. These were
some of the difficulties to which an upright governor exposed
himself. Cicero extricated himself with honour. Seldom was a
province so well administered as his under the Roman republic; but
he only brought back from it some gratitude, little money, and many
enemies, and very nearly quarrelled with Brutus.
Brutus, though we can scarcely believe it, had a hand in this traffic.
He had lent money to Ariobarzanes, king of Armenia, one of those
small princes that Rome charitably allowed to live, and to the town of