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The amygdala, emotion and learning

The amygdala, emotion and learning

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Published by Markoff Chaney
For many years the amygdala has been implicated in emotional processing. More recently its importance to our everyday psychology has been highlighted in the popular science press and in books such as Descartes’ Error by Antonio Damasio (1994) and the Emotional Brain by Joe LeDoux (1996). Evidence for its importance comes from a wide variety of sources, including cognitive psychology, neuropsychology and behavioural neuroscience. In this article I discuss some recent studies combining contemporary animal learning theory with behavioural neuroscience. These complement studies in normal humans and patient populations to increase our understanding of this vital brain structure. But what exactly is the amygdala?
For many years the amygdala has been implicated in emotional processing. More recently its importance to our everyday psychology has been highlighted in the popular science press and in books such as Descartes’ Error by Antonio Damasio (1994) and the Emotional Brain by Joe LeDoux (1996). Evidence for its importance comes from a wide variety of sources, including cognitive psychology, neuropsychology and behavioural neuroscience. In this article I discuss some recent studies combining contemporary animal learning theory with behavioural neuroscience. These complement studies in normal humans and patient populations to increase our understanding of this vital brain structure. But what exactly is the amygdala?

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Published by: Markoff Chaney on Mar 04, 2011
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October 2000
Spearman Medal Lecture
The Psychologist
Vol 13 No 10
OR many years the amygdalahas been implicated in emotionalprocessing. More recently itsimportance to our everyday psychologyhas been highlighted in the popular sciencepress and in books such as
by Antonio Damasio (1994) andthe
Emotional Brain
by Joe LeDoux(1996). Evidence for its importance comesfrom a wide variety of sources, includingcognitive psychology, neuropsychologyand behavioural neuroscience.In this article I discuss some recentstudies combining contemporary animallearning theory with behaviouralneuroscience. These complement studiesin normal humans and patient populationsto increase our understanding of this vitalbrain structure.But what exactly is the amygdala?
The amygdala
The amygdala is a small subcorticalstructure lying deep within the temporalregion of the brain. The name amygdala,like many of the seemingly complicatedterms in neuroanatomy, is merelya descriptive term related to the appearanceof the structure  in this case it is derivedfrom a word meaning ‘almond’.However, this relatively smallstructure has a complicated neuroanatomy.It receives an enormous range of neuralinputs from a wide variety of areas of thebrain, both cortical areas and subcorticalstructures. Much of the information itreceives is of a highly processed nature by which I mean it is information that hasundergone a great degree of manipulation,combination and recombination in otherregions of the brain.It receives highly processed informationfrom the visual system, the auditory cortex,the olfactory and gustatory neocortex andthe somatosensory cortex. In short, it isinformed about each of our five senses sight, hearing, smell, taste and touch.The amygdala also receives projectionsfrom areas of so-called association cortex information of a polymodal nature suchas that represented in the frontal cortex.Subcortically it receives information fromthe thalamus (relaying basic, unprocessedsensory signals), hippocampus (givingmore highly processed information aboutthe relationship between different objectsand events in the world), and a range of structures important in representing internalbodily states, such as hunger and thirst.In addition to this vast array of inputsit also projects to a wide variety of areas.It has reciprocal connections to many of the cortical areas just mentioned, as wellas projections to subcortical areas involvedin motor output, and hypothalamic andbrainstem regions involved in theco-ordination of autonomic, endocrineand behavioural responses.
The amygdala and emotion
There is a substantial body of work examining the role of the amygdalathat makes use of patients with selectivedamage to this region. This may occurdue to surgery, for example to destroy theepileptic focus in the case of intractableepilepsy, or in some very rare cases dueto congenital defects that lead to theprogressive calcification of limbicstructures sometimes restricted to theamygdala  Urbech–Wiethe disorder.More recent studies have madeexcellent use of emerging imagingtechnology, employing positron emissiontomography (PET) and functional magneticresonance imaging (fMRI) of healthyparticipants to examine brain systemsunderpinning learning and emotion.One striking finding, which has beenreplicated a number of times, relatesamygdala damage to a deficit inrecognising facial expressions of emotion.This work is typified by studies conductedby Andy Young at the University of York and Andy Calder at the MRC Cognitionand Brain Sciences Unit, Cambridge (e.g.Calder
et al.
, 1996). Moreover, this deficitappears to be selective, relating toexpressions of fear, anger and disgust,but not to happiness, sadness and surprise.Similarly, although patients withamygdala damage show surprisinglyfew deficits across a wide range of neuropsychological tests and can functionwell in everyday life, recent studies havestarted to pick out specific problems withsocial and emotional judgement.For example, Adolphs
et al.
(1998) took three people with highly selective damageto the amygdala and tested them on theirability to make judgements about theapproachability of others fromphotographs. In comparison to controls
The amygdala,emotion and learning
In the 1999 Spearman Medal Lecture
discussed the complex role of thisinfluential part of the brain.
Requests for reprints of this article should be addressed to:
Dr Simon KillcrossSchool of PsychologyCardiff UniversityTower BuildingPark PlacePO Box 901Cardiff CF11 3YG.Tel: 029 20875393E-mail: KillcrossAS@cf.ac.uk.
Amygdala damage can reduce recognition of fear,anger and disgust in people’s faces
they were far more likely to rate individualcharacters as approachable. This findingprovides solid empirical evidence to back up anecdotal observations made byfamilies, clinicians, and others workingwith these groups that their ability to makepersonal judgements can be abnormal.Finally, some excellent work has beencarried out examining the role of theamygdala in responding to emotionallysignificant events. These studies haveutilised PET and fMRI during Pavlovianconditioning in humans.In one such study individuals receivedpairings of pictures of angry faces with anintense and aversive burst of white noise(Morris
et al.
, 1998). Following thesepairings, participants were presented withjust the pictures of angry faces, whilstundergoing PET analysis of their neuralactivity. Very brief presentations of anangry face were immediately followedby presentation of a neutral face. Thisprocedure served to ‘mask’awareness of the angry face, so that participants werenot even conscious they had seen it.Two important findings emerged fromthe study. Firstly, participants showeda change in skin conductance responseduring the subliminal presentations of theangry face, demonstrating that it couldgenerate an emotional response by virtueof previous Pavlovian conditioning, evenwhen subjects were unaware of it.Secondly, and just as striking, theyfound a selective activation of the rightamygdala following such presentations.This again demonstrates the central role of the amygdala in responding to emotionallysignificant situations. Morris
et al.
(1998)also found a selective activation of just theleft amygdala when the angry face was notmasked and participants were aware of thepresentations, although discussion of thisintriguing result lies beyond the scope of this article.In summary, there is much evidence toassociate the amygdala with emotion, andwith learning about emotionally significantevents. It is involved in a wide variety of emotional responses, including respondingto linguistic threat, facial and vocalexpressions of emotion, memory foremotional events, and even simpleresponding to pleasant and aversive stimuli,including music.The evidence for the primaryinvolvement of the amygdala in emotion isoverwhelming. However, it should be notedthat the deficits observed followingselective damage to the amygdala are rathersubtle, and perhaps not as dramatic as onemight expect following destruction of a structure that appears to be so criticallyconnected to such a wide variety of otherbrain regions. Many of the deficits are onlyfully apparent when a patient is in real-lifesituations, with all their complex social andemotional interactions, rather than in themore controlled environment of thepsychological laboratory.Nevertheless, the amygdala is implicatedin a huge array of mental disorders. Theseinclude schizophrenia, depression,generalised anxiety disorders (it has oneof the densest collections of benzodiazepinereceptors in the brain, receptors thatrespond selectively to anti-anxiety drugssuch as diazepam), obsessive compulsivedisorder, and post-traumatic stress disorder.It is also thought to be vitally important inthe development and maintenance of drugabuse. All this, and yet deficits followingdiscrete amygdala damage appear to berelatively mild.In fact these findings reflect earlierwork in animals that similarly revealed theinvolvement of the amygdala in emotion.Perhaps most well-known are the classicstudies by Kluver and Bucy (1939). Thesestudies demonstrated that damage to thetemporal lobe in monkeys led to a widevariety of symptoms, which theycharacterised as disruptions in normalemotional responding.A classic example of a monkey withKluver–Bucy syndrome might be an animalthat shows little fear of its handlers, agreater propensity to explore new placesthat are frightening, a decreased abilityto recognise objects in the world (visualagnosia), a compulsion to place objects inits mouth to see if they are edible, and anincreased tendency to mate inappropriatelywith cagemates and other monkeys.Monkeys with damage to the temporallobes may show all or some of thesesymptoms.Similar findings have since beenreported in humans with gross damageto the temporal lobe (Aggleton, 1992).However, it is now widely recognised thatseveral of the components of this syndromeare not due to damage to the amygdala, butrather to overlying cortical areas, or due todamage to nerve fibre projections throughthe damaged region.More recent and more careful studiesin primates have revealed that deficitsfollowing discrete damage to the amygdalaare both subtle and mild (Málková
et al.,
1997).Finally, work in laboratory rats hasalso supported the findings from researchexamining humans and non-humanprimates. Deficits following discreteamygdala damage (in many cases toa subnucleus of the amygdala known asthe basolateral nucleus) are mild. Thereis a clear deficit in learning about someaversive events, such as a simplePavlovian conditioning situation wherea signal such as a tone is paired with anaversive event such as mild footshock (Davis, 1992; LeDoux, 1996). But thereappears to be little change in the ability
October 2000
The Psychologist
Vol 13 No 10
Spearman Medal Lecture
Acquisition of conditioned responding followingselective damage to the amygdala.During a conditioned stimulus thatpredicts the delivery of food thereis no difference in rate of approachto the site of food delivery relativeto baseline (per cent) betweenexperimental and control animals
of animals to learn about pleasant events,such as learning that a signal such asa tone predicts the delivery of food (seeFigure 1).Overall we can see that damage tothe amygdala produces some deficits inlearning about the emotional significanceof events in the world, although theseeffects are quite subtle and rather varied.We have yet to produce a clear andaccurate characterisation of its function ineveryday life; given the importance of theamygdala in mental disorders, achievingthis would be a valuable goal.
Associative learning
Associative learning theory characteriseslearning as the formation of an associationbetween mental representations of eventsin the world. In many cases (but by nomeans all), this learning is a result of theco-occurrence of the two events. In thepsychological laboratory these eventsmight be a simple cue such as a tone,and an emotionally significant event suchas presentation of food  a commondemonstration of Pavlovian, or classical,conditioning.Alternatively, the events might be somebehavioural response, such as a rat pressinga lever, and an emotionally or motivationallysignificant event such as presentation of foodor water. In this case learning is referred toas instrumental, or operant, conditioning.It is important to emphasise at this pointthat we are not merely talking about theformation of stimulus–response habits thatare frequently associated with behaviouristtheories of learning. We are talking aboutthe formation of associations betweenmental representations of events, andcontemporary animal learning theorythat deals with cognitive issues such asattention and short- and long-term memory,as well as with more traditional topics suchas motivation and incentive.It is from this background that I wouldlike to introduce some studies examiningthe role of the amygdala in associativelearning. Each of these studies makes useof findings drawn from current animallearning theory to try to throw light onthe precise function of the amygdala inlearning about emotionally significantevents. In each case, similar procedureswere used, based on simple associationssuch as those outlined above  a tonepredicting presentation of food, and a leverpress leading to presentation of food.
Representations for eventoutcomes and goals
A number of researchers have notedthat although animals with damage tothe amygdala appear to learn well thata particular signal or response predicts theimminent arrival of rewarding food, theydo not seem to retain as much knowledgeabout this food as normal animals.Take a specific example of this fromrecent work (Hatfield
et al.
, 1996). Whennormal animals have learnt the relationshipbetween a response or stimulus and areward, researchers can make changes inthe value of that reward. For example, theymight reduce its value by prefeeding theanimal on that reward  a phenomenonknown as sensory-specific satiety. Thiswill produce predictable changes in theanimal’s responses to the predictor of thatreward: in this example, a reduction in thelevel of lever pressing for that reward.This is most clearly demonstrated whenanimals have a choice of responses tomake, one response leading to one type of reward (say, a pellet of food) and a secondresponse leading to another type of reward(delivery of a small amount of sucrosesolution). Prefeeding an animal with foodpellets preserves responses that lead tosucrose solution, but practically eliminatesresponses that lead to food pellets. Theyhave already had their fill of food pellets.This type of result is central toassociative learning theory, anddemonstrates that responding is governedby an association between a signal orresponse and a representation of the rewardthat is predicted. It is generally acceptedthat satiation is effective as it reduces thevalue of internal representations of the goalor predicted outcome, and thus reduces thevigour of responding (see Dickinson, 1994).But animals with amygdala damage failto show sensitivity to these changes in thevalue of reward. Figure 2 shows the rate of lever pressing per 10 seconds on two leversduring a test of outcome value. Animalshad learned that pressing one lever led tofood pellets and pressing the other leverled to sucrose solution. Prior to this test,animals were given free access to eitherfood pellets or sucrose solution andallowed to feed to satiety. Responding issplit into presses on the lever that led to thefood with which the animals had been sated(devalued outcome) and presses on the leverthat led to the alternative food (non-devaluedoutcome). Although control animals assign
October 2000
Spearman Medal Lecture
The Psychologist
Vol 13 No 10
Increase in presses on a rewarded lever over a non-rewarded lever.In control animals this discrimination is acquired more rapidly if the two leversgive different rewards,an effect not found in animals with amygdala damage
Lack of sensitivity tochanges in reward value followingamygdala damage

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