Emotion Explained Edmund T. Rolls

Introduction: the issues
University Press Scholarship Online
Oxford Scholarship Online

Emotion Explained
Edmund T. Rolls
Print publication date: 2005

Print ISBN-13: 9780198570035
Published to Oxford Scholarship Online: September 2009
DOI: 10.1093/acprof:oso/9780198570035.001.0001

Edmund T. Rolls
DOI:10.1093/acprof:oso/9780198570035.003.0001
Abstract and Keywords

This introductory chapter asks a number of questions. What
are emotions? Why do we have emotions? What is their
adaptive value? What brain processes make us have emotions?
Why does it feel like something to be in an emotional state?
The issue of consciousness is introduced here. Further
questions are asked. How is motivation controlled? What
factors influence hunger and appetite, and affect body weight
control? What brain processes lead to drug addiction? How is
sexual behaviour controlled, and what factors underlie
differences in sexual behavior? The aims of the book are to
explain emotions in terms of what produces them (reinforcing
stimuli); why we have them (their evolutionary adaptive value:
an efficient way for genes to influence our behaviour to
increase their success); how we have them (the brain
mechanisms that implement them); and why they feel like
something (the problem of consciousness).
Page 1 of 16
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Chinese University of Hong Kong; date: 10 April 2016
Keywords: emotion, genes, evolution, consciousness, motivation, hunger,

appetite
1.1 Introduction
What are emotions? Why do we have emotions? What is their
adaptive value? What are the brain mechanisms of emotion,
and how can disorders of emotion be understood? Why does it
feel like something to have an emotion? Why do emotions
sometimes feel so intense? This book aims to provide answers
to all these questions. When we know what emotions are, why
we have them, how they are produced by our brains, and why
it feels like something to have an emotion, we will have a
broad-ranging explanation of emotion. It is in this sense that
the title of this book is Emotion Explained.
We can similarly ask what motivates us: What is motivation?
How is motivation controlled? How is motivation produced and
regulated by the brain? What goes wrong in motivational
disorders, for example in appetite disorders which produce
overeating and obesity? How do these motivational control
systems operate to ensure that we eat approximately the
correct amount of food to maintain our body weight, or
drinkjust enough to replenish our thirst? What are some of the
underlying reasons for the different patterns of sexual
behaviour found in different animals and humans? Why (and
how) do we like some types of touch (e.g. a caress), and what
is the relation of this to motivation? What brain processes
underlie addiction? What is the relation between emotion, and
motivational states such as hunger, appetite, and sexual
behaviour? It turns out that the explanations for motivational
behaviour are in many ways similar to those for emotional
behaviour, and therefore I also treat motivation in this book.
The aims of the book are to explain emotions in terms of the
following: What produces emotions? (The general answer I
propose is reinforcing stimuli, that is rewards and punishers.)
Why do we have emotions? (The overall answer I propose is
that emotions are evolutionarily adaptive as they provide an
efficient way for genes to influence our behaviour to increase
their fitness.) How do we have emotions? (I answer this by
describing what is known about the brain mechanisms of
emotion.) Why do emotional states feel like something? This is
Page 2 of 16
part of the large problem of consciousness, which I address in

Chapter 10.
Emotion and motivation are linked by the property that both
involve rewards and punishers. Emotions can be thought of as
states elicited by rewards or punishers. A full definition of
emotion, and theory of emotion, with a starting point as the
relation to rewards and punishers is described in Chapter 2.
Motivation can be thought of as a state in which a reward is
being sought, or a punisher is being avoided or escaped from.
This is made clear in Chapters 2, 3, 5, 6 and 9. Because of the
importance of reward and punishment for emotion and
motivation, I define in Section 1.2 reward and punishment, and
describe some of the types of learning that involve rewards
and punishers. This is useful groundwork for what follows in
the rest of this book. However, for those who wish in a first
reading to skip the definitions in Section 1.2 (which are
provided to ensure that there is a firm foundation for
understanding emotion and motivation), it may be useful
simply to think of a reward as something for which an animal
(which includes humans) will work, and a punisher as
something that an animal will work to
(p.2)
escape from or
avoid.
Some stimuli are innately rewarding or punishing and are
called primary reinforcers (for example no learning is
necessary to respond to pain as aversive), while other stimuli
are learned or secondary reinforcers (for example the sight of
a chocolate cake is not innately rewarding, but may become a
learned reinforcer, for which we may work, by the process of
association learning between the sight of the cake and its
taste, where the taste is a primary reward or reinforcer). This
type of learning, which is important in emotion and
motivation, is called stimulusreinforcement association
learning. (A better term is stimulusreinforcer association
learning, where reinforcer is being used to mean a stimulus
that might be a reward or a punisher.)
Page 3 of 16
1.2 Rewards and punishers, and learning about

rewards and punishers: instrumental learning
and stimulusreinforcer association learning
A reward is something for which an animal (including of
course a human) will work. A punisher is something that an
animal will work to escape or avoid (or that will decrease the
probability of actions on which it is contingent). In order to
exclude simple reflex-like behaviour, the concept invoked here
by the term work is to perform an arbitrary behaviour (called
an operant response) in order to obtain the reward or avoid
the punisher. An example of an operant response might be
putting money in a vending machine to obtain food, or for a rat
pressing a lever to obtain food. In these cases, the food is the
reward. Another example of an operant response might be
moving from one place to another in order to escape from or
avoid an aversive (punishing) stimulus such as a cold draught.
If the aversive stimulus starts and then the response is made,
this is referred to as escape from the punisher. If a warning
stimulus (such as a flashing light) indicates that the punisher
will be delivered unless the operant response is made, then
the animal may learn to perform the operant response when
the warning stimulus is given in order to avoid the punisher.
Because the definitions of reward and punisher make it a
requirement that it must be at least possible to demonstrate
learning of an arbitrary operant response (made to obtain the
reward or to escape from or avoid the punisher), we see that
learning is implicit in the definition of reward and punisher.
(Merely swimming up a chemical gradient towards a source of
food as occurs in single cell organisms is called a taxis as
described in Chapter 3; it does not require learning, and does
not make the food qualify as a reward under the definition.) In
that rewards and punishers do imply the ability to learn what
to do to obtain the reward or escape from or avoid the
punisher, we call rewards and punishers reinforcers.
This introduction leads to the definition of reinforcers as
stimuli that if their occurrence, termination, or omission is
made contingent upon the making of a response, alter the
probability of the future emission of that response (as a result
of the contingency (i.e. dependency) on the response). The
Page 4 of 16
alteration of the probability of a response (or action) is the

measure that learning has taken place. A positive reinforcer
(such as food) increases the probability of emission of a
response on which it is contingent; the process is termed
positive reinforcement, and the outcome is a reward (such
as food). A negative reinforcer (such as a painful stimulus)
increases the probability of emission of a response which
causes the negative rein-forcer to be omitted (as in active
avoidance) or terminated (as in escape), and the procedure
(p.3)
is termed negative reinforcement. In contrast,
punishment refers to procedures in which the probability of

an action is decreased. Punishment thus describes procedures
in which an action decreases in probability if it is followed by a
painful stimulus, as in passive avoidance. Punishment can also
be used to refer to a procedure involving the omission or
termination of a reward (extinction and time out
respectively), both of which decrease the probability of
responses (Gray 1975, Mackintosh 1983, Dickinson 1980,
Lieberman 2000). My argument is that an affectively positive
or appetitive stimulus (which produces a state of pleasure)
acts operationally as a reward, which when delivered acts
instrumentally as a positive reinforcer, or when not delivered
(omitted or terminated) acts to decrease the probability of
responses on which it is contingent. Conversely I argue that an
affectively negative or aversive stimulus (which produces an
unpleasant state) acts operationally as a punisher, which
when delivered acts instrumentally to decrease the probability
of responses on which it is contingent, or when not delivered
(escaped from or avoided) acts as a negative reinforcer in that
it then increases the probability of the action on which its nondelivery is contingent1.
Reinforcers, that is rewards or punishers, may be unlearned or
primary reinforcers, or learned or secondary reinforcers. An
example of a primary reinforcer is pain, which is innately a
punisher. The first time a painful stimulus is ever delivered, it
will be escaped from, and no learning that it is aversive is
needed. Similarly, the first time a sweet taste is delivered, it
can act as a positive reinforcer, so it is a primary positive
reinforcer or reward. Other stimuli become reinforcing by
learning, because of their association with primary
Page 5 of 16
reinforcers, thereby becoming secondary reinforcers. For

example, a (previously neutral) sound that regularly precedes
an electric shock can become a secondary reinforcer. Animals
will learn operant responses reinforced by the secondary
reinforcer, for example jumping to a place where the
secondary reinforcer is not present or terminates. Secondary
reinforcers are thus important in enabling animals to avoid
primary punishers such as pain.
There is a close relation of all these processes to emotion, for
as we will see in Chapter 2, fear is an emotional state that
might be produced by a sound that has previously been
associated with an electric shock. Shock in this example is the
primary punisher, and fear is the emotional state that occurs
to the tone stimulus as a result of the learning of the stimulus
(i.e. tone)reinforcer (i.e. shock) association. Another example
of a secondary reinforcer is a visual stimulus associated with
the taste of a food. For example, the first time we see a new
type of food we do not treat the sight of the new visual
stimulus as reinforcing, but if the stimulus has a good taste,
the sight of the object becomes a positive secondary
reinforcer, and we may choose the food when we see it in
future by virtue of its association with a primary reinforcer.
This type of learning is thus called stimulusreinforcer
association learning. (The operation is often referred to as
stimulusreinforcement association learning.) This type of
learning is very important in many emotions, because it is as a
result of this type of learning that many previously neutral
stimuli come to elicit emotional responses, as in the example
of fear above.
Unconditioned reinforcing stimuli often elicit autonomic
responses. (Autonomic responses are those mediated through
the autonomic nervous system, via the vagus and sympathetic
nerves, which affect smooth muscle.) Examples include
alterations of heart rate
(p.4)
and of blood pressure which
might be produced by a painful stimulus; and salivation which

might be produced by the taste of food. Many endocrine
(hormonal) responses are also mediated through the
autonomic nervous system and so are autonomic responses,
for example the release of adrenaline (epinephrine) from the
adrenal gland during emotional excitement. Previously neutral
Page 6 of 16
stimuli, such as the sound in our previous example, can by

pairing with unconditioned stimuli, such as shock in the
previous example, come by learning the association, to
produce learned autonomic responses. In the example the tone
might by pairing with shock come to elicit a change in heart
rate, and sweating. This type of learning is called classical
conditioning, and also Pavlovian conditioning after Ivan
Pavlov who performed many of the original studies of this type
of learning, including learned salivation to the sound of a bell
that predicted the taste of food. It is a type of learning that is
very similar to stimulusreinforcer association learning, except
that in the case of classical conditioning the responses
involved are autonomic and endocrine responses.
In the case of stimulusreinforcer association learning, the
effects of the learning are mediated through the skeletal
motor system, in that actions are performed that are
instrumental in enabling animals to obtain rewards or avoid
punishers, and are described as voluntary in humans. A key
difference between instrumental learning and classical
conditioning apart from the response systems involved lies in
the contingencies that operate. In classical conditioning the
animal has no control over whether the unconditioned
stimulus is delivered (as in the experiments of Pavlov just
described). In contrast, the whole notion of instrumental
learning is that what the animal does is instrumental in
determining whether the reinforcer (the goal) is obtained, or
escaped from or avoided. Both types of learning are important
in emotions because (as we will see in Chapter 2) instrumental
reinforcers produce emotional responses, but also typically
produce autonomic responses that therefore typically occur
during emotional states, and indeed mediate important effects
of emotions such as preparing the body for action by
increasing heart rate etc.
A more detailed description of the nature of classical
(Pavlovian) conditioning and instrumental learning, and how
both are related to emotion, is provided in Section 4.6.1.
Motivation refers to the state an animal is in when it is
willing to work for a reward or to escape from or avoid a
punisher. So for example we say that an animal is motivated to
Page 7 of 16
work for the taste of food, and in this case the motivational
state is called hunger. The definition of motivation thus
implies the capacity to perform any, arbitrary, operant
response in order to obtain the reward or escape from or avoid
the punisher. By implying an operant response, we exclude
simple behaviours such as reflexes and taxes (such as
swimming up a chemical gradient), as described above and in
Chapter 2. By implying learning of any response to obtain a
reward (or avoid a punisher), motivation thus focuses on
behaviours in which a goal is defined. Motivation is one of the
states that are involved in the large area of brain design
related to the fundamental issue of how goals for behaviour
are defined, and an appropriate behaviour is selected, as
described in this book and brought together into a theory in
Chapter 2.
1.3 The approaches taken to emotion and

motivation: their causes, functions, adaptive
value, and brain mechanisms
(p.5)
To explain emotion, and motivation, a number of different

approaches are taken, and some of these need some
introduction. To examine the causes of emotion, the
environmental stimuli and situations that elicit emotions are
identified. This is part of the subject of Chapter 2. It is shown
how the different environmental stimulus conditions that
produce emotions provide the basis for a classification of
different emotions. Understanding the functions of emotion
also provides part of the explanation of why we have emotions,
and many of the functions of emotion are described in Chapter
3. These functions of emotion explain in part the adaptive
value of emotion, and give part of an explanation about why
emotion has evolved. However, it turns out that emotions
provide a fundamental solution to the issue of how genes
design brains to produce behaviour that is advantageous to
the genes, and this deep understanding of the adaptive value
of emotion, and in a sense the cause of emotion, is elaborated
in Chapter 3. When considering the adaptive value of emotion
in the context of evolution, we must remember that animals
are generally social, and that evolution may have led to the
development of special reward and punishment systems to
Page 8 of 16
help to produce emotional behaviour that is adaptive in social

situations. This area, of understanding and explaining aspects
of social behaviour in terms of its evolutionary adaptive value,
is the field of sociobiology, and this approach is introduced
especially in the context of sexual behaviour in Chapter 9.
Another major approach taken to explain emotion and
motivation, and their underlying reward and punishment
systems, is in terms of the brain mechanisms that implement
them. Understanding the brain processing and mechanisms of
behaviour is one way to ensure that we have the correct
explanation for how the behaviour is produced. Another
important reason for investigating the actual brain
mechanisms that underlie emotion and motivation, and reward
and punishment, is not only to understand how our own brains
work, but also to have the basis for understanding and
treating medical disorders of these systems. Now it turns out
that many of the brain systems that are involved in emotion
and motivation have undergone considerable development in
primates (e.g. monkeys and humans) compared to nonprimates (for example rats and mice)2. It is because of the
intended relevance to understanding human emotion and its
disorders that emphasis is placed in this book on findings from
research in non-human primates, including monkeys. Another
reason for focusing interest on the primate brain is that there
has been great development of the visual system in primates,
and this itself has had important implications for the types of
sensory stimuli that are processed by brain systems involved
in emotion and motivation. One example is the importance of
face identity and face expression decoding, which are both
important in primate emotional behaviour, and indeed provide
an important part of the foundation for much primate social
behaviour. These are among the reasons why emphasis is
placed on brain systems in primates, including humans,
(p.6)
in the approach taken here. The overall medically relevant aim

of the research described in this book is to provide a
foundation for understanding the brain mechanisms of
emotion and motivation, and thus their disorders, including
depression, anxiety, addiction, sociopathy, and borderline
personality disorder, in humans.
Page 9 of 16
When considering brain mechanisms involved in emotion in

primates, recent findings with the human brain imaging
approaches are described. These approaches include
functional magnetic resonance imaging (fMRI) to measure
changes in brain oxygenation level locally (using a signal from
deoxyhaemoglobin) to provide an index of local brain activity,
as well as positron emission tomography (PET) studies to
estimate local regional cerebral blood flow, again to provide
an index of local brain activity. It is, however, important to
note that these functional neuroimaging approaches provide
rather coarse approaches to brain function, in that the spatial
resolution is seldom better than 2 mm, so that the picture
given is one of blobs on the brain, which give some indication
of what is happening where in the brain, and what types of
dissociation of functions are possible.
However, because there are millions of neurons in each of the
areas that can be resolved with functional neuroimaging, such
imaging techniques give rather little evidence on how the
brain works. For this, one needs to know what information is
represented in each brain area at the level at which
information is exchanged between the computing elements of
the brain, the neurons (brain cells). One also needs to know
how the representation of information (for example about
stimuli or events in the world) changes from stage to stage of
the processing in the brain, to understand how the brain
works as a system. It turns out that one can read this
information from the brain by recording the activity of single
neurons, or groups of single neurons. The reason that this is
an effective procedure for understanding what is represented
is that each neuron has one information output channel, the
firing of its action potentials, so that one can measure the full
richness of the information being represented in a region by
measuring the firing of its neurons. This can reveal
fundamental evidence crucial for understanding how the brain
operates. For example, neuronal recording can reveal all the
information represented in an area even if parts of it are
encoded by relatively small numbers, perhaps a few percent,
of its neurons. (This is impossible with brain-imaging
techniques, which also are susceptible to the interpretation
problem that whatever causes the largest activation is
interpreted as what is being encoded in a region).
Page 10 of 16
Neuronal recording also provides evidence for the level at

which it is appropriate to build computational models of brain
function, the neuronal network level. Such neuronal network
computational models consider how populations of neurons
with the connections found in a given brain area, and with
biologically plausible properties such as learning rules for
altering the strengths of synaptic connections between
neurons, actually could perform useful computation to
implement the functions being performed by that brain area.
This approach should not really be considered as a metaphor
for brain operation, but as a theory of how each part of the
brain operates. The neuronal network computational theory,
and any model or simulation based on it, may of course be
simplified to some extent to make it tractable, but
nevertheless the point is that the neuron-level approach,
coupled with neuronal network models, together provide some
of the fundamental elements for understanding how the brain
actually works. For this reason, emphasis is also placed in this
book on what is known about what is being processed in each
brain area as shown by recordings from neurons. Such
evidence, in terms of building theories and models of how the
brain functions, can never be replaced by brain imaging
evidence, although these approaches do complement each
(p.7)
other very effectively. The approach to brain function in
terms of computations performed by neuronal networks in

different brain areas is the subject of the books Neural
Networks and Brain Function by Rolls and Treves (1998) and
Computational Neuroscience of Vision by Rolls and Deco
(2002). The reader is referred to these books for more
comprehensive accounts of this biologically plausible approach
to brain function. In this book, some of the neurophysiological
evidence and its computational implications for understanding
how our brains work to produce emotion and motivation are
described.
One of the main points made in this section is that rapid
progress is being made now in understanding emotion and
motivation, and part of this advance is related to the fact that
we are just starting to be able to understand how the brain
actually works, in terms of how its neuronal networks
transform inputs into behaviour. Given that this basis for
Page 11 of 16
understanding how our own brains work does depend very

much on understanding in detail how the brains of relatively
close relatives, non-human primates such as monkeys, work,
research on non-human primate brain information processing
is quite crucial. We are really at the start of understanding in
full detail how this works, but one aim of this book is to show
how coming from that research new understanding into how
our own brains work is being produced to the extent that even
such complex processes as emotion and motivation are now
seen to be at the heart of brain design. Moreover, that
understanding is important for understanding the rich variety
of human emotional behaviour, and also for understanding and
treating disorders in emotion and motivation.
1.4 Reward, punishment, emotion, and

motivation: the plan of the book
It may be useful to make it clear why the brain mechanisms of
both emotion and motivation (with the examples of motivated
behaviour considered being hunger, thirst, addiction, and
sexual behaviour) are being considered together in this book.
The reason is that for both emotion and motivation, rewards
and punishers are assessed in order to provide the goals for
behaviour. Operation of the brain to evaluate rewards and
punishers is the fundamental solution of the brain to
interfacing sensory systems to action selection and execution
systems. Computing the reward and punisher value of sensory
stimuli, and then using selection between different rewards
and avoidance of punishers in a common reward-based
currency appears to be the fundamental design that brains use
in order to produce appropriate behaviour (see Chapter 2).
The behaviour selected can be thought of as appropriate in the
sense that it is based on the sensory systems and reward
decoding that our genes specify (through the process of
natural selection) in order to maximize their fitness
(reproductive potential). Having reward and punishment
systems is the solution that evolution has developed to
produce appropriate behaviour. It happens that motivational
and emotional behaviour are the types of behaviour in which
rewards and punishers operate.
Page 12 of 16
Considering both emotional and motivational behaviour in this

book means that we can describe many of the principles that
underlie the decoding of many types of rewarding and
punishing stimuli, which have in common that they produce
affective states. We can also see to some extent how the
common currency of reward works to enable different rewards
to be compared, and in particular how the reward value of all
the different potential rewards that our genes specify is kept
within a comparable range, so that we select different
behaviours as appropriate. That is, we can examine many of
the different ways in which the reward value
(p.8)
of different
sensory stimuli is modulated, both by internal signals as

physiological needs are satisfied, and in addition to some
extent by sensory-specific satiety (the mechanism by which
repeating one reward causes it gradually to decrease its
reward value somewhat, assisting the selection of other
rewards in the environment).
However, perhaps the most important reason for treating
reward and punishment systems, and the brain systems
dealing with rewards and punishers, together is that we can
develop an overall theory of how this set of issues, which
might sometimes appear mysterious, is actually at the heart of
brain design. Much of sensory processing, at least through the
brain systems that are concerned with object identification
(whether by sight, sound, smell, taste, or touch), can be seen
to have the goal of enabling the correct reward value to be
decoded and represented after the object has been identified.
This means for example that in vision, representations of
objects that can be accessed regardless of the view of the
object shown, the size of the object on the retina, etc, must be
formed. Moreover, these invariant representations of objects
must be encoded in an appropriate way for the brain to
associate in simple neuronal networks the object with primary
(unlearned) reinforcers, such as the taste associated with the
object, or the pain produced by the object. The actual
motivational and emotional parts of the processing, the parts
where the reward or punisher value is made explicit in the
representation, should indeed no longer be seen as mysterious
or perhaps superfluous aspects of brain processing. Instead,
they are at the heart of which behavioural actions are
Page 13 of 16
selected, and how they are selected. Moreover, a large part of

the brain's action and motor systems can be seen as having
the goal in systems-level design of producing behaviour that
will obtain the rewards decoded from sensory (and memory)
inputs by the motivational and emotional systems of the brain.
In particular, the implication is that the action systems for
implicit (unconscious) behaviour have as part of their design
principle the property that they will perform actions to
optimize the output of the reward and punishment systems
involved in motivation and emotion. Put another way, the brain
systems involved in motivation and emotion must pass reward
or punisher signals to the action systems, which must be built
to attempt to obtain and maximize the reward signals being
received; to switch behaviour from one reward to another as
the reward values being received alter; and to switch
behaviour also if signals indicating possible punishers are
received (see Chapter 4).
This book is thus intended to uncover some of the most
important and fundamental aspects and principles of brain
function and design. The book is also intended to show that
the way in which the brain works in motivation and emotion
can be seen to be the result of natural selection operating to
select genes that optimize our behaviour by building into us
the appropriate reward and punisher systems, and the
appropriate rules for the operation of these systems.
The plan of the book is that we consider in Chapters 2 and 3
the major issue of emotion, and its functions. These Chapters
address the explanation of emotion by defining emotion, and
elucidating its functions. Another part of the explanation of
emotion is how it actually works, that is how it is implemented
in the brain, which is described in Chapter 4. By
understanding its mechanisms, we not only understand better
the different processes that contribute to emotion, but also we
provide a fundamental basis for starting to understand many
disorders of emotion, including for example depression, and
how they can be treated. Affective (emotional) states, and
rewards, are involved in motivated behaviour such as eating,
drinking, addiction, and sexual behaviour, and we consider
these in Chapters 5, 6 and 9. These topics provide many clear
examples of how the pleasantness or reward value of stimuli
Page 14 of 16
reflect a fundamental aspect
(p.9)
of the design of both the
brain and behaviour, and help to show the rewards, and

punishers, that actually influence many aspects of our
behaviour. In Chapter 10 the issue of emotional feelings,
which is part of the big issue of consciousness, is considered,
together with the brain processing involved in conscious
feelings.
The aims of this book are thus to explain emotions in terms of:
1. What produces emotions? The answer developed in
Chapter 2 is that emotions are produced by reinforcing
stimuli.
2. Why do we have emotions? The most fundamental,
and thoroughly Darwinian, explanation, developed in
Chapter 3, is that the evolutionary adaptive value of
emotions is that they provide an efficient way for genes
to influence our behaviour to increase their (the genes)
success.
3. How do we have emotions, that is, what are the brain
and body processes that implement emotions, and
motivational states such as hunger? These processes
are described in Chapters 49.
4. Why do emotional states feel like something, which is
part of the very large problem of consciousness. This is
considered in Chapter 10.
Notes:
(1) Note that my definition of a punisher, which is similar to

that of an aversive stimulus, is of a stimulus or event that can
either decrease the probability of actions on which it is
contingent, or increase the probability of actions on which its
non-delivery is contingent. The term punishment is restricted
to situations where the probability of an action is being
decreased.
(2) For example, the temporal lobe has undergone great
development in primates, and several systems in the temporal
lobe are either involved in emotion (e.g. the amygdala), or
provide some of the main sensory inputs to brain systems
involved in emotion and motivation. The prefrontal cortex has
Page 15 of 16
also undergone great development in primates, and one part

of it, the orbitofrontal cortex, is very little developed in
rodents, yet is one of the major brain areas involved in
emotion and motivation in primates including humans. The
development of some of these brain areas has been so great in
primates that even evolutionarily old systems such as the taste
system appear to have been rewired, compared with that of
rodents, to place much more emphasis on cortical processing,
taking place in areas such as the orbitofrontal cortex (see
Rolls (1999a) and Rolls and Scott (2003)).
Page 16 of 16

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Introduction: the issues

University Press Scholarship Online

Oxford Scholarship Online

Print publication date: 2005

Introduction: the issues

Abstract and Keywords

Introduction: the issues

Keywords: emotion, genes, evolution, consciousness, motivation, hunger,

Introduction: the issues

part of the large problem of consciousness, which I address in

Introduction: the issues

1.2 Rewards and punishers, and learning about

Introduction: the issues

alteration of the probability of a response (or action) is the

is termed negative reinforcement. In contrast,

punishment refers to procedures in which the probability of

Introduction: the issues

reinforcers, thereby becoming secondary reinforcers. For

and of blood pressure which

might be produced by a painful stimulus; and salivation which

Introduction: the issues

stimuli, such as the sound in our previous example, can by

Introduction: the issues

1.3 The approaches taken to emotion and

To explain emotion, and motivation, a number of different

Introduction: the issues

help to produce emotional behaviour that is adaptive in social

in the approach taken here. The overall medically relevant aim

Introduction: the issues

When considering brain mechanisms involved in emotion in

Introduction: the issues

Neuronal recording also provides evidence for the level at

other very effectively. The approach to brain function in

terms of computations performed by neuronal networks in

Introduction: the issues

understanding how our own brains work does depend very

1.4 Reward, punishment, emotion, and

Introduction: the issues

Considering both emotional and motivational behaviour in this

sensory stimuli is modulated, both by internal signals as

Introduction: the issues

selected, and how they are selected. Moreover, a large part of

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reflect a fundamental aspect

of the design of both the

brain and behaviour, and help to show the rewards, and

(1) Note that my definition of a punisher, which is similar to

Introduction: the issues

also undergone great development in primates, and one part

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