Brain, Vol. 125, No.

10, 2152-2190, October 2002

2002 Oxford University ress
Review Article
Disorders of memory
!ic"ael #. $o%el&an
Ne'ro%syc"iatry and !e&ory #isorders (linic, University #e%art&ent of syc"iatry and syc"olo)y,
*'y+s, $in)+s and ,t -"o&as+ !edical ,c"ool .$(/0, /ondon, U$
(orres%ondence to1 !ic"ael #. $o%el&an, Ne'ro%syc"iatry and !e&ory #isorders (linic, University
#e%art&ent of syc"iatry and syc"olo)y, *'y+s, $in)+s and ,t -"o&as+ !edical ,c"ool .$(/0, ,t
-"o&as+ (a&%'s, /ondon, ,21 324, U$ 2-&ail1 &ic"ael.5o%el&an65cl.ac.'5
7eceived October 80, 2001. 7evised 9%ril 2, 2002. ,econd revision 9%ril 1:, 2002. 9cce%ted 9%ril 20,
2002.
Summary

-"is %a%er revie;s disorders of &e&ory. 9fter a
brief s'rvey

of t"e clinical varieties of t"e a&nesic
syndro&e, transient

and %ersistent, selected
t"eoretical iss'es ;ill be considered

by %osin) a
series of <'estions. .i0 ="at is i&%aired and ;"at

is
s%ared in antero)rade a&nesia> .ii0 #o te&%oral
lobe, dience%"alic

and frontal lobe a&nesias differ>
.iii0 4o; inde%endently se&antic

is se&antic
&e&ory> .iv0 ="at deter&ines t"e %attern and
extent

of retro)rade &e&ory loss> .v0 (an retro)rade
a&nesia ever be

?isolated+> .vi0 #oes %syc"o)enic
a&nesia involve

t"e sa&e &ec"anis&s as or)anic a&nesia> .vii0 4o; and ;"en do

false
&e&ories arise> (o&&onalities as ;ell as differences across

se%arate literat'res ;ill be
e&%"asi@ed, and t"e case for a

&ore ?dyna&ic+ .interactionist0 a%%roac" to t"e

investi)ation of a&nesia ;ill be advocated.

Keywords1 a&nesiaA antero)radeA confab'lationA %syc"o)enicA retro)rade
Top
Summary
Introduction
Varieties of amnesic syndrome
What is impaired and...
Explicit and implicit memory
Do temporal lobe, diencephalic...
ow independently semantic is...
What determines the pattern...
!an retro"rade amnesia e#er...
Does psycho"enic amnesia in#ol#e...
ow and when do...
!onclusions
$eferences
%bbre#iations1 99 B antero)rade a&nesiaA C#* 2- B D
1:
4Efl'oro-deoxy )l'cose 2-A
*9B9 B )a&&a-a&ino b'tyric acidA $ B ?5no;+A -9 B %ost-tra'&atic a&nesiaA -,#
B %ost-tra'&atic stress disorderA 7 B ?re&e&ber+A 79 B retro)rade a&nesiaA ,2(- B
sin)le %"oton e&ission to&o)ra%"yA -29 B transient e%ile%tic a&nesiaA -*9 B transient
)lobal a&nesiaA - B t"ia&ine %yro%"os%"ateA =9F,-7 B =ec"sler 9d'lt Fntelli)ence
,caleG7evisedA =!,-7 B =ec"sler !e&ory ,caleG7evised
Introduction

?,o ;e beat on, boats a)ainst t"e c'rrent, borne bac5

ceaselessly into t"e %ast.+

.C. ,cott Cit@)erald, The Great Gatsby, 1925.0

?-"e %laces t"at ;e "ave 5no;n belon) no; only to t"e

little ;orld of s%ace on ;"ic" ;e
&a% t"e& for o'r o;n convenience.

None of t"e& ;as ever &ore t"an a t"in slice, "eld
bet;een t"e

conti)'o's i&%ressions t"at co&%osed o'r life at t"at ti&eA

re&e&brance of a
%artic'lar for& is b't re)ret for a %artic'lar

&o&entA and "o'ses, roads, aven'es are as
f')itive, alas, as

t"e years.+

.!. ro'st, Remembrance of Things Past, 1918.0

-"e literat're on &e&ory and its disorders "as %roliferated,

%artic'larly since t"e advent
of recent ne'roi&a)in) tec"ni<'es.

4o;ever, controversy re&ains concernin) certain
broad iss'es,

selected exa&%les of ;"ic" ;ill be considered in t'rn in t"is

revie;,
follo;in) a brief s'rvey of t"e clinical disorders t"at

)ive rise to an a&nesic syndro&e.
-"e foc's of t"e revie; ;ill

be on ne'ro%syc"olo)ical st'dies of %atients ;it" disorders

of e%isodic or se&antic &e&ory, and t"ere ;ill be only brief

reference to f'nctional
ne'roi&a)in) st'dies of "ealt"y individ'als.

Varieties of amnesic syndrome

-"e amnesic syndrome can be defined as1 ?9n abnor&al &ental

state in ;"ic" &e&ory
and learnin) are affected o't of all %ro%ortion

to ot"er co)nitive f'nctions in an ot"er;ise
alert and res%onsive

%atient+ .Victor et al., 1931 0.

-"e Korsakoff syndrome can be defined as t"e sa&e b't ;it" t"e

follo;in) %"rase added1
?... res'ltin) fro& n'tritional

de%letion, notably t"ia&ine deficiency.+

Fn fact, Victor et al. .1931 0 'sed t"e first descri%tion as

a definition of t"e $orsa5off
syndro&e, b't t"e %resent a't"or

feels t"at it is i&%ortant to distin)'is" bet;een a&nesic
syndro&es

in )eneral .for ;"ic" t"e Victor et al. definition s'ffices0

and t"e %artic'lar
clinical condition described by $orsa5off

.1::9 0, ;"ose cases .;it" "indsi)"t0 can all
be vie;ed as "avin)

s'ffered n'tritional de%letion, ;"et"er of alco"olic or non-alco"olic

ca'sation. Vario's disorders can )ive rise to an a&nesic syndro&e,

incl'din) "er%es
ence%"alitis, severe "y%oxia, certain vasc'lar

lesions, "ead inH'ry, dee% &idline t'&o'rs,
basal forebrain

lesions and occasionally early de&entia.

The Korsa&off syndrome
9s H'st &entioned, t"e $orsa5off syndro&e is t"e res'lt of n'tritional

de%letion, i.e.
t"ia&ine deficiency. $orsa5off .1::9 0 described

t"is condition as res'ltin) fro& alco"ol
ab'se or fro& a n'&ber

of ot"er ca'ses, b't by far t"e &ost co&&on no;adays is alco"ol
ab'se.

-"ere are fre<'ent &is'nderstandin)s abo't t"e nat're of t"is

disorder. ?,"ort ter&
&e&ory+ is intact in t"e Ia&esian

sense of recall over a &atter of seconds .Jan);ill, 19KL
0, b't

learnin) over &ore %rolon)ed %eriods is severely i&%aired, and

t"ere is 's'ally a
retro)rade &e&ory loss ;"ic" c"aracteristically

extends bac5 &any years or decades
.$o%el&an, 19:9 A ar5in et

al., 1990 b0. $orsa5off .1::9 0 "i&self noted t"at "is
%atients

?reason abo't everyt"in) %erfectly ;ell, dra; correct

ded'ctions fro& )iven
%re&ises, &a5e ;itty re&ar5s, %lay c"ess

or a )a&e of cards, in a ;ord co&%ort
t"e&selves as &entally

so'nd %ersons.+ 4o;ever, "e also noted re%etitive <'estionin)s,

t"e extensive nat're of t"e retro)rade &e&ory loss, and a %artic'lar

%roble& in
re&e&berin) t"e te&%oral se<'ence of events, associated

;it" severe disorientation in
ti&e. 9s ;ill be disc'ssed belo;,

"e )ave exa&%les of confab'lation t"at reflected t"e
%roble&

;it" te&%oral se<'ence &e&ory, s'c" t"at real &e&ories ;ere

H'&bled '% and
retrieved ina%%ro%riately, o't of te&%oral context.

!any cases of t"e $orsa5off syndro&e are dia)nosed follo;in)

an ac'te =ernic5e
ence%"alo%at"y, involvin) conf'sion, ataxia,

nysta)&'s and o%"t"al&o%le)ia. Not all
t"ese feat'res are al;ays

%resent, and t"e o%"t"al&o%le)ia in %artic'lar res%onds ra%idly

to treat&ent ;it" "i)"-dose vita&ins. -"ese feat'res are often

associated ;it" a
%eri%"eral ne'ro%at"y. 4o;ever, t"e disorder

can also "ave an insidio's onset .('ttin),
193: a0, and s'c"

cases are &ore li5ely to co&e to t"e attention of %syc"iatrists1

in t"ese
cases, t"ere &ay be either no 5no;n "istory or only

a transient "istory of =ernic5e
feat'res. -"ere are also re%orts

t"at t"e c"aracteristic =ernic5eM$orsa5off
ne'ro%at"olo)y

.see belo;0 is dia)nosed &'c" &ore co&&only at a'to%sy in alco"olics

t"an in life .-orvi5 et al., 19:2 A 4ar%er, 19:8 0, i&%lyin) t"at

&any cases are bein)
&issed.

-"e $orsa5off syndro&e is 'n's'al a&on) &e&ory disorders in

t"at t"ere is a distinct
ne'roc"e&ical %at"olo)y ;it" i&%ortant

i&%lications for treat&ent. Collo;in) ani&al
researc" in t"e

1980s and 19K0s, and t"e i&%ortant observations of de =ardener

and
/ennox .19K3 0 and ot"ers in &alno'ris"ed %risoners-of-;ar,

t"ia&ine de%letion ;as
establis"ed as t"e &ec"anis& )ivin) rise

to t"e ac'te =ernic5e e%isode, and t"e
s'bse<'ent .$orsa5off0

&e&ory i&%air&ent. 4o;ever, t"e )enetic factor, %redis%osin)

certain "eavy drin5ers to develo% t"is syndro&e before "e%atic

or )astro-intestinal
co&%lications, re&ains 'nclear. Ft ;as

t"o')"t t"at a trans5etolase )ene &i)"t acco'nt for
t"is, beca'se

trans5etolase is t"e en@y&e t"at re<'ires t"ia&ine %yro%"os%"ate

.-0 as a
co-factor. ,'c" a )ene "as been identified .!c(ool

et al., 1998 0, b't it does not acco'nt
for all t"e %ro%erties

of trans5etolase and only very ;ea5ly for %redis%osition to

t"e
$orsa5off syndro&e. !oreover, it re&ains 'nclear "o; t"ia&ine

de%letion %rod'ces t"e
%artic'lar ne'ro%at"olo)y fo'nd in =ernic5eM$orsa5off

%atients, alt"o')" =itt .19:5 0
%ointed o't t"at six ne'rotrans&itter

syste&s .incl'din) acetylc"oline, *9B9, )l'ta&ate0
are affected

by t"ia&ine de%letion, eit"er by red'ction of --de%endent

en@y&e activity
or by direct str'ct'ral da&a)e .see also B'tter;ort",

19:9 0. ="atever t"e %recise
&ec"anis&, treat&ent as soon as

%ossible ;it" "i)" doses of %arenterally ad&inistered
&'lti-vita&ins

is essential. -"e =ernic5e feat'res res%ond ;ell to "i)" doses

of vita&ins,
and s'c" treat&ent can %revent t"e occ'rrence of

a c"ronic $orsa5off state .Victor et al.,
1931 A /is"&an, 199: 0.

-"e s&all ris5 of ana%"ylaxis is co&%letely o't;ei)"ed by t"e

lar)e ris5 of severe brain da&a)e if s'c" treat&ent is not ad&inistered.

-"e c"aracteristic ne'ro%at"olo)y in ;"at is often 5no;n as

t"e ?=ernic5eM$orsa5off
syndro&e+ consists

of ne'ronal loss, &icro-"ae&orr"a)es and )liosis in t"e
%araventric'lar

and %eri-a<'ed'ctal )rey &atter .Victor et al., 1931 0. 4o;ever,

t"ere "as
been debate as to ;"ic" %artic'lar lesions are critical

for t"e c"ronic &e&ory disorder to
arise. Victor et al. .1931 0

%ointed o't t"at all 2K of t"eir cases, in ;"o& t"e &edial-
dorsal

n'cle's of t"e t"ala&'s ;as affected, "ad a clinical "istory

of %ersistent &e&ory
i&%air&ent .$orsa5off syndro&e0, ;"ereas

five cases, in ;"o& t"is n'cle's ;as
'naffected, "ad a "istory

of =ernic5e feat'res ;it"o't any recorded clinical "istory of

s'bse<'ent &e&ory disorder. By contrast, t"e &a&illary bodies

;ere i&%licated in all t"e
=ernic5e cases, ;"et"er or not t"ere

;as s'bse<'ent &e&ory i&%air&ent. 4o;ever,
!air et al. .1939 0

%rovided a caref'l %at"olo)ical and ne'ro%syc"olo)ical descri%tion

of
t;o $orsa5off %atients, ;"ose a'to%sies s"o;ed lesions in

t"e &a&illary bodies and t"e
anterior and &idline t"ala&'s,

incl'din) t"e %aratenial b't not t"e &edial-dorsal n'clei.
!air

et al. s'))ested t"at t"e lesions t"ey described &i)"t ?disconnect+

a critical &e&ory
circ'it r'nnin) bet;een t"e te&%oral lobes

and t"e frontal cortex. !ayes et al. .19:: 0
obtained very si&ilar

findin)s in t;o f'rt"er $orsa5off %atients, ;"o ;ere also very

caref'lly described bot" ne'ro%syc"olo)ically and at a'to%sy.

!ore recently, 4ardin) et
al. .2000 0 fo'nd ne'ronal loss and

atro%"y in t"e &a&illary bodies and &edial-dorsal
t"ala&ic n'cle's

of 18 =ernic5e %atients, ;"et"er or not t"ey develo%ed .$orsa5off0

a&nesia. (o&%arison of ei)"t $orsa5off ;it" five ?=ernic5e

only+ %atients s"o;ed
differences confined to t"e anterior

%rinci%al t"ala&ic n'cle's, s'))estin) t"at atro%"y in
t"is

str'ct're is critical for t"e develo%&ent of a&nesia.

-"ere is also ne'ro%at"olo)ical evidence of )eneral cortical

atro%"y in $orsa5off %atients
%artic'larly involvin) t"e frontal

lobes .-orvi5 et al., 19:2 A 4ar%er et al., 19:3 0, and
t"is is

associated ;it" ne'ro%syc"olo)ical evidence of ?frontal+

or ?exec'tive+ test
dysf'nction in t"ese %atients

./en) and ar5in, 19:: A Iacobson et al., 1990 A ,"o<eirat
et

al., 1990 A $o%el&an, 1991 0.

-"ere "ave been a n'&ber of ne'roi&a)in) st'dies in t"e $orsa5off

syndro&e. (- scan
st'dies indicated a de)ree of )eneral cortical

atro%"y, %artic'larly involvin) t"e frontal
lobes .,"i&a&'ra

et al., 19:: A Iacobson and /is"&an, 1990 0. !7F st'dies "ave

indicated &ore s%ecific atro%"y in dience%"alic and frontal

str'ct'res .Ierni)an et al.,
1991 A (olc"ester et al., 2001 0.

-"e findin)s in sin)le %"oton e&ission to&o)ra%"y
.,2(-0 and

2- st'dies "ave been &ore variable, so&e st'dies s"o;in) ;ides%read

"y%o%erf'sion and "y%o&etabolis& .4'nter et al., 19:9 A aller

et al., 1993 0, ot"er
st'dies s"o;in) very little c"an)e relative

to "ealt"y controls .!artin et al., 1992 0.
="ite &atter and

dience%"alic c"an)es "ave also been i&%licated .7eed et al.,

2002 0.

Victor et al. .19310 re%orted t"at 25N of $orsa5off %atients

?recovered+, 50N s"o;ed
i&%rove&ent t"ro')" ti&e

and 25N re&ained 'nc"an)ed. ="ilst it is 'nli5ely t"at any
establis"ed

$orsa5off %atient s"o;s co&%lete recovery, t"e %resent a't"or+s

ex%erience is
t"at s'bstantial i&%rove&ent does occ'r over a

&atter of years, if %atients refrain fro&
alco"ol1 it is %robably

correct to say t"at 35N of $orsa5off %atients s"o; a variable

de)ree of i&%rove&ent, ;"ilst 25N s"o; no c"an)e.

erpes encephalitis
-"is can )ive rise to a %artic'larly severe for& of a&nesic

syndro&e .=ilson and
=earin), 1995 0. -"e &aHority of cases are

said to be %ri&ary infections, and a "istory of
a %recedin)

?cold sore+ on t"e li% is 'nco&&on. -"ere is c"aracteristically

a fairly abr'%t
onset of ac'te fever, "eadac"e and na'sea. ,ei@'res

can occ'r, and t"ere &ay be
be"avio'ral c"an)es. -"e f'lly develo%ed

clinical %ict're ;it" nec5 ri)idity, vo&itin),
and &otor and

sensory deficits &ay ta5e several days to e&er)e .eto and I'el-Iensen,

199L 0. #ia)nosis is by a raised titre of antibodies to t"e vir's

in t"e (,C, b't often t"is is
&issed and a %res'&%tive dia)nosis

is &ade on t"e basis of t"e clinical %ict're as ;ell as
severe

si)nal alteration in t"e te&%oral lobes on !7F brain i&a)in),

associated ;it" loss
of tiss'e vol'&e, "ae&orr"a)e and oede&a.

Ne'ro%at"olo)ical and ne'roi&a)in) st'dies s"o; t"at t"ere is

's'ally extensive bilateral
te&%oral lobe da&a)e .4ierons et

al., 193: A N. $a%'r et al., 199K A Ooneda et al., 199K
A (olc"ester

et al., 2001 0 alt"o')", occasionally, t"e c"an)es are s'r%risin)ly

'nilateral
.,tan"o%e and $o%el&an, 2000 0. -"ere are often frontal

c"an)es, &ost co&&only in t"e
orbito-frontal re)ions, and t"ere

is a variable de)ree of )eneral cortical atro%"y. -"e
&edial

te&%oral lobe str'ct'res are %artic'larly severely affected,

incl'din) t"e
"i%%oca&%i, a&y)dalae, and t"e entor"inal, %erir"inal

and %ara"i%%oca&%al cortices.
2vidence fro& st'dies of bilateral

te&%oral lobecto&y .,coville and !ilner, 1953 A
!ilner, 19LL 0,

as ;ell as ani&al lesion st'dies .Jola-!or)an et al., 19:9 A

!'rray et
al., 1998 A Jola et al., 2000 0, "as indicated t"at

t"ese str'ct'res are %artic'larly critical
in &e&ory for&ation.

-"e c"ronic &e&ory disorder in "er%es ence%"alitis s"o;s &any

rese&blances to t"at in
t"e $orsa5off syndro&e, consistent ;it"

t"e fact t"at t"ere are &any ne'ral connections
bet;een t"e

t"ala&i, &a&illary bodies and t"e "i%%oca&%i .9))leton and ,a'nders,

1993
0. 2nce%"alitis, li5e "ead inH'ry, can also i&%licate basal

forebrain str'ct'res, ;"ic" )ive
c"oliner)ic o't%'ts to t"e

"i%%oca&%iA since t"ese are t"o')"t to &od'late "i%%oca&%al

f'nction, t"is &ay f'rt"er exacerbate t"e da&a)e .#a&asio et

al., 19:5 A "illi%s et al.,
19:3 A von (ra&on and ,c"'ri, 1992 0.

(ontrary to ;"at ;as %ost'lated in t"e 19:0s, t"ere a%%ears

to be no difference bet;een
$orsa5off %atients and "er%es ence%"alitis

%atients in ter&s of for)ettin) rates .$o%el&an
and ,tan"o%e,

1993 0, b't "er%es %atients s"o; better ?insi)"t+

into t"e nat're of t"eir
disorder .$o%el&an et al., 199: a0 and

a ?flatter+ te&%oral )radient to t"eir retro)rade

&e&ory loss .i.e. less s%arin) of early &e&ories0. -"ey &ay

"ave a %artic'larly severe
deficit in s%atial &e&ory, es%ecially

;"en t"e ri)"t "i%%oca&%'s is i&%licated
.$o%el&an et al., 1993 0.

,e&antic &e&ory is &ore co&&only affected in "er%es
%atients,

and t"is res'lts fro& t"e involve&ent of t"e left infero-lateral

te&%oral lobe,
%rod'cin) i&%air&ents in na&in), readin) .?s'rface

dyslexia+0 and ot"er as%ects of
lexical-se&antic &e&ory.

7i)"t te&%oral lobe da&a)e &ay lead to a %artic'larly severe

i&%air&ent in face reco)nition &e&ory or 5no;led)e of %eo%le

.e.). 2slin)er et al., 199L
0.

Se#ere hypoxia
,evere "y%oxia can )ive rise to an a&nesic syndro&e follo;in)

carbon &onoxide
%oisonin), cardiac and res%iratory arrests,

or s'icide atte&%ts by "an)in) or %oisonin)
;it" t"e ex"a'st

%i%e fro& a car. #r') overdoses &ay %reci%itate %rolon)ed
'nconscio'sness

and cerebral "y%oxia, and t"is <'ite co&&only occ'rs in "eroin

ab'sers.
9 recent revie; "as %rovided a ti&ely re&inder t"at

t"e ne'ro%at"olo)ical and co)nitive
conse<'ences of "y%oxia

are variable and can be ;ides%read .(aine and =atson, 2000 0,

a %oint ;idely acce%ted in clinical %ractice b't so&eti&es ne)lected

in t"e
ne'ro%syc"olo)ical literat're.

Jola-!or)an et al. .19:L0 described a %atient ;"o, follo;in)

re%eated e%isodes of
"y%oxia and cardio-res%iratory arrest,

develo%ed &oderately severe antero)rade a&nesia.
9t a'to%sy

L years later, t"is %atient ;as s"o;n to "ave a severe loss

of %yra&idal cells in
t"e (91 re)ion of t"e "i%%oca&%i bilaterally,

;"ereas t"e rest of t"e brain a%%eared
relatively nor&al. ress

et al. .19:90 re%orted "i%%oca&%al atro%"y on !7F in t"ree
a&nesic

%atients, and $o%el&an et al. .2001 0 fo'nd &edial te&%oral lobe

atro%"y in
"y%oxic %atients alt"o')" t"ese %atients also s"o;ed

red'ced )l'cose &etabolis& in t"e
t"ala&i on D
1:
4Efl'oro-deoxy

)l'cose 2- .C#* 2-0 .7eed et al., 1999 A co&%are
!ar5o;itsc"

et al., 1993 b0. Fn brief, t"e &e&ory disorder is li5ely to res'lt

fro& a
co&bination of "i%%oca&%al and t"ala&ic c"an)es, related

to t"e &any co&&on ne'ral
%at";ays bet;een t"ese t;o str'ct'res

.Ca@io et al., 1992 A 9))leton and ,a'nders, 1993
0. 9 recent

findin) of fornix atro%"y and &e&ory deficits follo;in) carbon

&onoxide
%oisonin) .$esler et al., 2001 0 is consistent ;it"

t"is vie;.

Vascular disorders
Vasc'lar disorders can %artic'larly affect &e&ory, as o%%osed

to )eneral co)nitive
f'nctionin), in .i0 t"ala&ic, &edial te&%oral

or retros%lenial infarction, and .ii0 s'b
arac"noid "ae&orr"a)e.

Fn an ele)ant (- scan st'dy, von (ra&on et al. .19:5 0 s"o;ed

t"at it ;as da&a)e to t"e
anterior t"ala&'s t"at ;as critical

in %rod'cin) an a&nesic syndro&e .co&%are ;it"
4ardin) et al.,

2000 0. ="en t"e %at"olo)y ;as confined to t"e &ore %osterior

re)ions of
t"e t"ala&'s, &e&ory f'nction ;as relatively 'naffected.

-"e anterior re)ion of t"e
t"ala&'s is variably s'%%lied by

t"e %olar or %ara&edian arteries in different individ'als,
bot"

of ;"ic" are, 'lti&ately, branc"es of t"e %osterior cerebral

circ'lation. !ore
recently, Van der =erf et al. .2000 0, in a

revie; of t"e literat're, ar)'ed t"at it is da&a)e
to t"e &a&illo-t"ala&ic

tract, ;"ic" %roHects into t"e anterior n'clei of t"e t"ala&'s,

t"at
is critical in %rod'cin) an a&nesic syndro&e. 9 relatively

%'re lesion of t"e anterior
t"ala&'s %rod'ces antero)rade a&nesia

.990 ;it" &ini&al retro)rade a&nesia .790
.*raff-7adford et

al., 1990 A ar5in and 4'n5in, 1998 A $a%'r et al., 199La 0. (ases

of
&ore extensive 79 .4od)es and !c(art"y, 1998 0 or de&entia

"ave been described,
%robably reflectin) t"e extent to ;"ic"

fronto-cortical %roHections are i&%licated in t"e
infarction.

-"e "i%%oca&%i are s'%%lied by t"e anterior and %osterior c"oroidal

arteries, branc"es of
t"e internal carotid and %osterior cerebral

arteries, res%ectively .=als", 19:3 0. Unilateral
infarction

tends to )ive rise to &aterial-s%ecific &e&ory loss and bilateral

da&a)e to
)lobal a&nesia .O+(onnor and Verfaellie, 2002 0.

-"e retros%leni'& is also s'%%lied
fro& t"e %osterior cerebral

artery, and infarction or "ae&orr"a)e can %rod'ce a&nesia by

disr'%tin) connections to anterior t"ala&'s, entor"inal and

%ara"i%%oca&%al cortices
.Valenstein et al., 19:3 A !a)'ire,

2001 0.

,'barac"noid "ae&orr"a)e follo;in) r'%t're of an ane'rys& can

res'lt in &e&ory
i&%air&ent, ;"et"er t"e anterior cerebral or

%osterior cerebral circ'lation in t"e (ircle of
=illis is involved

.7ic"ardson, 19:9 , 19910. 7'%t'red ane'rys&s in t"e anterior

co&&'nicatin) artery can i&%licate t"e basal forebrain and ventro-&edial

frontal
str'ct'res. *ade and ot"ers "ave ar)'ed t"at it is ;"et"er

or not t"e se%tal n'cle's of t"e
basal forebrain is da&a)ed

t"at deter&ines ;"et"er a %ersistent a&nesic syndro&e occ'rs

in s'c" %atients .*ade, 19:2 A *ade and !ortensen, 1990 0. Ot"ers

"ave attrib'ted t"e
florid confab'lation t"at so&eti&es occ'rs

to associated ventro-&edial frontal da&a)e
.e.). #e /'ca and

(icerone, 1991 A !oscovitc" and !elo, 1993 A *ilboa and
!oscovitc",

2002 0.

ead in'ury
4ead inH'ry can )ive rise to eit"er transient or %ersistin)

a&nesia. Ft is i&%ortant to
distin)'is" bet;een 79, ;"ic" is

's'ally .b't not necessarily0 relatively brief, %ost-
tra'&atic

a&nesia .-90, and ?islands+ of %reserved &e&ory

;it"in t"e a&nesic )a%
.7'ssell and Nat"an, 19KL A /is"&an, 19L: ,

1938 , 199: 0. Occasionally, -9 &ay
exist ;it"o't any 79, alt"o')"

t"is is &ore co&&on in cases of %enetratin) lesions.
,o&eti&es

t"ere is a %artic'larly vivid &e&ory for i&a)es or so'nds occ'rrin)

i&&ediately before t"e inH'ry, on re)ainin) conscio'sness, or

d'rin) a l'cid interval
bet;een t"e inH'ry and t"e onset of

-9.

-9 is )enerally ass'&ed to reflect t"e de)ree of 'nderlyin)

diff'se brain %at"olo)y, in
%artic'lar rotational forces )ivin)

rise to diff'se axonal inH'ry. $in) .1993 0 "as re%orted
t"at

-9 can be assessed ;it" reasonable reliabilityA t"e len)t"

of -9 is %redictive of
event'al co)nitive .Broo5s, 19:K 0, %syc"iatric

./is"&an, 19L: 0, and social o'tco&e
.7'ssell and ,&it", 19L1 A

Broo5s, 1991 0. 4o;ever, t"e d'ration of -9 is often not
;ell

doc'&ented in &edical records, and t"ese relations"i%s are often

;ea5er t"an is
)enerally ass'&ed. -9 needs to be distin)'is"ed

fro& %ersistin) antero)rade &e&ory
i&%air&ent, ;"ic" &ay be

detected on clinical assess&ent or co)nitive testin) lon) after

t"e %eriod of -9 "as ended. /evin et al. .19:: a0 fo'nd t"at,

d'rin) -9, "ead inH'ry
%atients s"o;ed accelerated for)ettin)

of learned infor&ation, ;"ereas after -9
for)ettin) rates ;ere

nor&al .co&%are ;it" Baddeley et al., 19:3 0.

#a&a)e to t"e frontal and anterior te&%oral lobes by direct

tra'&a res'lts in cont'sion,
"ae&ato&a and "ae&orr"a)e. (ontre-co'%

da&a)e, intracranial "ae&orr"a)e and
"y%oxia can all res'lt

.-easdale and !endelo;, 19:K 0. 7otational and accelerationM
deceleration

forces &ay %rod'ce s"earin) or tensile stretc"in) of axons ;it"

s'bse<'ent
)liosis .,tric" et al., 195L A O%%en"ei&er, 19L: 0.

-"e res'ltin) intra-axonal c"an)es
lead to a fail're in axo%las&ic

trans%ort, axonal s;ellin), and, 'lti&ately, to
disconnection,

de-afferentation and loss of syna%tic bo'tons over a &atter

of "o'rs or
days .Bl'&ber)s et al., 1995 A ovlis"oc5 and ("rist&an,

1995 0.

!e&ory is co&&only t"e last co)nitive f'nction to s"o; i&%rove&ent

follo;in) an
ac'te tra'&a .(on5ey, 198: 0, and %atients can s"o;

t"e c"aracteristic feat'res of an
a&nesic syndro&e .Baddeley

et al., 19:3 A /evin et al., 19:: b0. #is%ro%ortionate 79
"as

been described .see belo;0, %artic'larly in association ;it"

da&a)e to t"e frontal or
anterior te&%oral re)ions. Cor)etf'lness

is also a co&&on co&%laint ;it"in t"e context of
&ild conc'ssion

./is"&an, 19:: A Cle&in)er, 2000 0. ="ilst researc" st'dies s"o;

t"at
recovery fro& &ild "ead inH'ry is ty%ically ex%ected 1M8

&ont"s %ost-inH'ry, so&e
%atients de&onstrate sy&%to&s far beyond

t"is ti&e .Bart" et al., 199L 01 lon) %eriods of
disability and

end'rin) %ost-conc'ssion sy&%to&s are related, in %art, to individ'al

v'lnerabilities. Fn s'c" cases, co&%laints c"aracteristically

%ersist lon) after t"e
settle&ent of any co&%ensation iss'es

.!ers5ey and =oodforde, 1932 A !iller, 1939 A
-ars" and 7oyston,

19:5 0.

7ecent controversy "as concerned ;"et"er severe "ead inH'ry

and a&nesia excl'de t"e
%ossibility of %ost-tra'&atic stress

disorder .-,#0 sy&%to&s. O+Brien and N'tt .199: 0
ar)'ed

t"at a "ead inH'ry ca'sin) co&a and severe a&nesia %revents

t"e ?re-ex%eriencin)+
sy&%to&s of -,#. 4o;ever,

!c!illan .199L0 described 10 %atients ;it" "ead inH'ry
of varyin)

severity, in ;"o& -,# arose. -"ese 10 %atients re%orted ?;indo;s+

of
ex%erience, in ;"ic" e&otional dist'rbance ;as s'fficient

to ca'se -,#, even t"o')"
-9 ;as of relatively lon) d'ration

.t"ree %atients "ad -9s of &ore t"an 2 ;ee5s0.
-"ese ?;indo;s+

involved recall of events close to i&%act ;"en 79 ;as brief

.e.). of a
lorry bearin) do;n0, or of distressin) events soon

after t"e accident .;"en -9 ;as
s"ort0 or of ?islands+

of &e&ory .e.). "earin) t"e screa&in) of ot"ers0. Ot"ers re%orted

distressin) recollections t"at a%%eared to be self-)enerated

or t"at "ad been re%orted to
t"e %atient by ot"ers. !cNeil and

*reen;ood .199L 0 re%orted -,# sy&%to&s in a
yo'n) traffic accident

victi&, ;"o "ad an 79 of 2 days and a -9 of K ;ee5s. #es%ite

t"ese cases, t"e to%ic re&ains controversial, and so&e a't"ors

"ave even dis%'ted
;"et"er victi&s of &ild "ead inH'ry can "ave

-,# .for revie;, see 4arvey et al., 2002
0.

Transient "lobal amnesia
-ransient )lobal a&nesia .-*90 &ost co&&only occ'rs in t"e &iddle-a)ed

or elderly,
&ore fre<'ently in &en, and res'lts in a %eriod

of a&nesia lastin) several "o'rs. 9s is
;ell 5no;n, it is c"aracteri@ed

by re%etitive <'estionin), and t"ere &ay be so&e
conf'sion,

b't %atients do not re%ort any loss of %ersonal identity. Ft

is so&eti&es
%receded by "eadac"e or na'sea, a stressf'l life

event, a &edical %roced're, intense
e&otion or vi)oro's exercise.

4od)es and =ard .19:9 0 fo'nd t"at t"e &ean d'ration of
a&nesia

;as K " and t"e &axi&'& 12 ". Fn 25N of t"eir sa&%le, t"ere

;as a %ast "istory
of &i)raine, ;"ic" ;as considered to "ave

a %ossible aetiolo)ical role. Fn a f'rt"er 3N,
t"e %atients

s'bse<'ently develo%ed 'ne<'ivocal feat'res of e%ile%sy in t"e

absence of
any %revio's "istory of sei@'res. -"ere ;as no association

;it" eit"er a %ast "istory of or
ris5 factors for vasc'lar disease,

nor ;it" clinical si)ns indicatin) a vasc'lar %at"olo)y. Fn
%artic'lar, t"ere ;as no association ;it" transient isc"ae&ic

attac5s. Fn L0M30N of t"e
sa&%le, t"e 'nderlyin) aetiolo)y

;as 'nclear.

,o&e;"at si&ilarly, !iller et al. .19:3 0, in a sa&%le of 150

&en and 123 ;o&en, fo'nd
t"at t"e &ean d'ration of t"e e%isode

;as L.2 ", ran)in) fro& 2 to 12 ". 9 sei@'re
disorder ;as noted

in ei)"t %atients and, as in t"e 4od)es and =ard .19:9 0 re%ort,

t"e
incidence of cerebro-vasc'lar events .stro5e0 ;as no "i)"er

t"an ;o'ld be ex%ected in
t"is a)e )ro'%.

="ere ne'ro%syc"olo)ical tests "ave been ad&inistered to %atients

d'rin) t"eir ac'te
e%isode of transient a&nesia .$ritc"evs5y

et al., 19:: A 4od)es and =ard, 19:9 0, t"e
%atients "ave s"o;n

a %rofo'nd 99 on tests of bot" verbal and non-verbal &e&ory,

b't
79 ;as variable, 's'ally bein) relatively brief b't occasionally

%rolon)ed, and ;orse for
recall of e%isodes t"an facts .2vans

et al., 1998 A *'illery et al., 2000 0. Collo;in) t"e
attac5,

$ritc"evs5y and ,<'ire .19:9 0 re%orted t"at t"ere ;as co&%lete

recovery after
several ;ee5s. !iller et al. .19:3 0 stated t"at

?a selective verbal &e&ory deficit+ ;as
fo'nd ?in

a fe; %atients. .. Dalt"o')"E %ersistin) &e&ory co&%laints or

fran5 de&entia
follo;in) -*9 ;ere seldo& noted.+ 4o;ever,

4od)es and Oxb'ry .1990 0, in a follo;-
'% of K1 cases at L &ont"s,

fo'nd statistically si)nificant i&%air&ents on tests &eas'rin)

t"e recall of stories, fa&o's faces, and a'tobio)ra%"ical &e&ories.

-"e )eneral consens's is t"at t"e a&nesic disorder res'lts fro&

transient dysf'nction in
li&bic-"i%%oca&%al circ'its, cr'cial

to &e&ory for&ation. Cor exa&%le, ,till"ard et al.
.1990 0 re%orted

severe bite&%oral "y%o%erf'sion d'rin) an e%isode of -*9 'sin)

,2(- and, after recovery fro& t"e e%isode, cerebral %erf'sion

ret'rned to nor&al.
2vans et al. .1998 0 obtained si&ilar findin)s,

also in a ,2(- st'dy, ;it" recovery to
nor&al 3 ;ee5s later.

C'Hii et al. .19:9 0 'sed C#* 2- in -*9 %atients 8 &ont"s after

t"e e%isode, obtainin) nor&al findin)s in all b't one of t"e

%atients. ,i&ons and 4od)es
.2000 0 and *oldenber) .2002 0 "ave

cited evidence t"at c"an)es on diff'sion-;ei)"ted
!7F &i)"t

indicate isc"ae&ic disf'nction in t"ese brain re)ions.

Transient epileptic amnesia
-"is refers to t"e &inority of -*9 cases in ;"o& e%ile%sy a%%ears

to be t"e 'nderlyin)
ca'se .4eat"field et al., 1938 A Cis"er,

19:2 A !iller et al., 19:3 A 4od)es and =ard,
19:9 0. -"e &ain

%redictive factors for an e%ile%tic aetiolo)y are brief e%isodes

of
&e&ory loss .1 " or less0 and &'lti%le attac5s .!iller et

al., 19:3 A 4od)es and =arlo;,
1990 A $a%'r, 1990 0. Ft is i&%ortant

to note t"at standard 22* and (- scan findin)s are
often nor&al.

4o;ever, an e%ile%tic basis to t"e disorder &ay be revealed

on slee% 22*
.$o%el&an et al., 199K a0. $a%'r .1990 0 coined t"e

%"rase ?transient e%ile%tic a&nesia+
.-290 to describe

s'c" attac5s1 t"is is a 'sef'l ter&, alt"o')" it does not distin)'is"

bet;een e%isodes t"at are ictal .al&ini et al., 1992 A V'ille'&ier

et al., 199L 0 and
t"ose t"at are post-ictal in nat're .-assinari

et al., 1991 0.

atients ;it" -29 &ay s"o; resid'al deficits in bet;een t"eir

attac5s, associated ;it"
t"eir 'nderlyin) ne'ro%at"olo)y. -"ese

&ay involve antero)rade &e&ory .$o%el&an et
al., 199K a0 or as%ects

of re&ote &e&ory .$a%'r et al., 19:L , 19:9 A Je&an et al.,
199: 0.

!ost co&&only, t"e %atients co&%lain of ?)a%s+ in

t"eir %ast &e&ory. Ft see&s
%la'sible t"at t"e %atients &ay

"ave "ad brief r'ns of sei@'re activity in t"e %ast1 t"ese
;o'ld

"ave been 'ndetected clinically b't res'lted in fa'lty .antero)rade0

encodin) of
very s%ecific ite&s in a'tobio)ra%"ical &e&ory .$o%el&an,

2000 a0. 9n alternative
inter%retation is t"at t"e c'rrent e%ile%sy

so&e"o; %revents t"e retrieval of old,
a'tobio)ra%"ical &e&ories

.$a%'r, 2000 A !anes et al., 2001 0.

2%ile%sy &ay, of co'rse, )ive rise to a'to&atis&s or %ost-ictal

conf'sional states
./o)sdail and -oone, 19:: A Cen;ic5, 1990 A

/is"&an 199: 0. ="ere t"ere is an
a'to&atis&, t"ere is al;ays

bilateral involve&ent of t"e li&bic str'ct'res involved in
&e&ory

for&ation, incl'din) t"e "i%%oca&%al and %ara"i%%oca&%al str'ct'res

bilaterally
as ;ell as t"e &esial dience%"alon .Centon, 1932 0.

(onse<'ently, a&nesia for t"e %eriod
of a'to&atic be"avio'r

is al;ays %resent and is 's'ally co&%lete .$nox 19L: A Cen;ic5,

1990 , 19980.

Summary
-"e above exa&%les do not, of co'rse, co&%rise a co&%re"ensive

list of t"e disorders
;"ic" )ive rise to transient or %ersistent

a&nesia. ="at t"ey "ave in co&&on is t"at
te&%orary or %er&anent

dysf'nction or da&a)e in &edial te&%oralPdience%"alic circ'itry

.or in t"e basal forebrain c"oliner)ic in%'ts to t"at circ'itry0

%rod'ces t"e a&nesia. -"e
re&ainder of t"is article concerns

t"eoretical iss'es t"at arise in o'r 'nderstandin) of t"e
f'nctionin)

of t"is circ'itry and its interaction ;it" frontal lobe and

te&%oral neocortical
&ec"anis&s.

What is impaired and what is spared in
antero"rade amnesia(

9s is ;ell 5no;n, a distinction is 's'ally dra;n bet;een so-called

?;or5in) &e&ory+,
;"ic" "olds infor&ation for brief

%eriods of ti&e .a &atter of seconds0 and allocates
reso'rces,

and secondary &e&ory in ;"ic" infor&ation is stored on a %er&anent

or se&i-
%er&anent basis .Ia&es, 1:90 A 4ebb, 19K9 A Baddeley and

=arrin)ton, 1930 A ,"allice
and =arrin)ton, 1930 0. ,econdary

&e&ory can, in t'rn, be s'bdivided into an e%isodic
.or ?ex%licit+0

co&%onent, se&antic &e&ory and i&%licit &e&ory. 2%isodic &e&ory

refers to incidents or events fro& a %erson+s %ast .allo;in)

t"at %erson ?to travel bac5
&entally in ti&e+0 and

it is c"aracteristically affected in t"e a&nesic syndro&e, ;"ereas

se&antic &e&ory refers to 5no;led)e of facts, conce%ts and lan)'a)e

.-'lvin), 1932 0.
F&%licit &e&ory incl'des classical conditionin),

t"e %roced'ral learnin) of %erce%t'o-
&otor s5ills and t"e facilitation

of res%onses in t"e absence of ex%licit &e&ory 5no;n as
?%ri&in)+.

!'c" ne'ro%syc"olo)ical researc" "as exa&ined ;"ic" of t"ese

co&%onents
are affectedPs%ared in a&nesia, and t"e 'nderlyin)

&ec"anis&s of deficit.

Encodin", stora"e and retrie#al
Over t"e years, t"ere "as been extensive debate concernin) ;"et"er

t"e %ri&ary deficit in
t"e a&nesic syndro&e lies in eit"er .i0

t"e initial encodin) of e%isodic &e&ories, or .ii0
so&e 5ind

of %"ysiolo)ical ?consolidation+ into secondary

&e&ory, or .iii0 fa'lty
encodin) and stora)e of context'al infor&ation,

or .iv0 accelerated for)ettin) of
infor&ation, or .v0 in retrieval

%rocesses .=arrin)ton and =eis5rant@, 1930 A B'tters and
(er&a5,

19:0 A !e'dell and !ayes, 19:2 0.

)i* +aulty encodin"
,o&e t"eories %ro%ose t"at t"ere is a deficit in t"e %syc"olo)ical

%rocesses involved in
t"e initial ?re)istration+

or re%resentation of infor&ation. Fn %artic'lar, it "as been

ar)'ed
t"at, ;"ilst $orsa5off %atients are able to ?encode+

t"e direct, sensory %ro%erties of
infor&ation, t"ey "ave diffic'lties

in ?%rocessin)+ its &ore &eanin)f'l .se&antic0
<'alities

.e.). B'tters and (er&a5, 19:0 0. Cor exa&%le, t"ey %erfor& %artic'larly

badly
at learnin) ;ord-%airs .e.). "'n)ryMt"in0, t"e recall

of ;"ic" is nor&ally facilitated by
t"in5in) of se&antic lin5s

bet;een t"e ;ords .('ttin), 193: bA B'tters and (er&a5,
19:0 0.

On t"e ot"er "and, )ivin) instr'ctions or orientin) tas5s t"at

enco'ra)e t"e
extraction of &eanin) fro& a sti&'l's %rod'ces,

at &ost, a relatively s&all en"ance&ent
in t"e a&nesic %atients+

s'bse<'ent recall of t"at sti&'l's, an effect closely si&ilar

to t"at
seen in "ealt"y controls .!e'dell et al., 1939 A !c#o;all,

19:1 A !e'dell and !ayes,
19:2 0. (onse<'ently, it see&s 'nli5ely

t"at a fail're to encode se&antic infor&ation is
t"e f'nda&ental

&e&ory deficit in a&nesia.

)ii* +aulty consolidation
9 second ty%e of "y%ot"esis %ro%oses t"at t"ere is i&%air&ent

in t"e %"ysiolo)ical
%rocesses t"at are ass'&ed to occ'r shortly

after an initial re%resentation is laid do;n in
order to establis"

.?consolidate+0 infor&ation into so&e relatively

%er&anent for& .e.).
!e'dell et al., 1939 A !oscovitc", 19:2 0.

-"ese %rocesses ;ere traditionally t"o')"t to
involve t"e ?transfer+

of infor&ation fro& %ri&ary to secondary &e&ory, o%eratin)
d'rin)

a ti&e-%eriod of so&et"in) less t"an a &in'te. -"e "y%ot"esis

;as s'))ested by
t"e findin) t"at s%an test scores and ot"er

&eas'res of %ri&ary &e&ory .re<'irin) t"e
recall of infor&ation

;it"in a fe; seconds0 ;ere relatively intact in $orsa5off, &edial

te&%oral lesion and "ead inH'ry %atients .e.). !ilner, 19LL A

Broo5s, 19:K 0. Fn one
%artic'larly intri)'in) st'dy, /ync" and

Oarnell .1938 0 intervie;ed six conc'ssed
9&erican footballers

;it"in 80 s of t"eir inH'ry and at intervals t"ereafter. 9lt"o')"

t"ey
;ere initially able to )ive a l'cid acco'nt of events occ'rrin)

H'st before t"e blo;, t"ey
s'bse<'ently develo%ed a ?relatively

co&%lete+ 79, s'))estin) t"at t"eir i&&ediate
&e&ories

"ad not been effectively consolidated. 9 %roble& ;it" t"is t"eory

is t"at it
cannot by itself ex%lain ;"y 79 of &ore t"an a fe;

seconds or &in'tes s"o'ld occ'r
'nless one %ost'lates a distinct

?slow+ consolidation %rocess lastin) years or decades.

-"is
iss'e ;ill be disc'ssed belo;.

)iii* +aulty encodin" or stora"e of contextual information
4'%%ert and iercy %ro%osed a very s%ecific deficit in a&nesic

%atients+ ac<'isition of
context'al .e.). te&%oralPs%atial0

infor&ation .4'%%ert and iercy, 193L , 193: 0, and
t"is idea

;as ado%ted by several ot"er )ro'%s .e.). !ayes et al., 19:5 0.

-"ere is certainly
s'bstantial evidence of dis%ro%ortionate

context'al &e&ory deficits in a&nesic %atients,
b't diffic'lties

;it" t"is t"eory incl'de .a0 t"e fact t"at t"e %attern of context'al

&e&ory
deficits &ay vary s'bstantially across %atient )ro'%s

.e.). ar5in et al., 1990 a0, and t"at
dis%ro%ortionate context'al

&e&ory deficits are not al;ays fo'nd .,<'ire, 19:2 A (ave
and

,<'ire 1991 0A and .b0 s'c" deficits "ave often been attrib'ted

.%er"a%s erroneo'sly0
to conco&itant frontal lobe %at"olo)y,

rat"er t"an to t"e &edial te&%oralPdience%"alic
lesions critical

to t"e develo%&ent of antero)rade a&nesia. 7ecently, t"is t"eory

"as
evolved into a &ore )enerali@ed notion of a deficit in bindin)

co&%lex associations
.!ayes and #o;nes, 1993 0 or &e&ory for

relations bet;een ite&s .(o"en et al., 1993
0. Fn t'rn, t"is

notion relates to vario's distinctions to be considered belo;1

bet;een recall
and reco)nition &e&ory, re&e&berin) and 5no;in),

and bet;een ex%licit and i&%licit
&e&ory.

)i#* %ccelerated for"ettin"
9 fo'rt" %ossibility foc'ses '%on ?stora)e+ .retention0

rat"er t"an learnin) %rocesses. On
t"e basis of findin)s in

a sin)le-case .4!0, 4'%%ert and iercy .1939 0 ar)'ed t"at
%atients

;it" "i%%oca&%al lesions s"o; accelerated for)ettin), even after

&aterial "as
been ade<'ately learnedA "o;ever, t"is ;as not

t"e case in %atients ;it" dience%"alic
%at"olo)y. ,o&e s'%%ort

for t"is findin) "as been obtained in ot"er %atients ;it"
"i%%oca&%al

%at"olo)y .ar5in and /en), 19:: A Cris5 and !ilner, 1990 0 and

also in
9l@"ei&er %atients .4art et al., 19:3 0. 4o;ever, t"ere

;ere ceilin) effects in t"e first
t;o of t"ese st'dies, and

a fail're to 'se e<'ivalent &eas'res at different ti&e-%eriods

in
t"e t"ird. !oreover, Creed et al. .19:3 0 failed to re%licate

4'%%ert and iercy+s ori)inal
observation in 4! .4'%%ert

and iercy, 1939 0 in 4! ;"en "e ;as st'died re%eatedly
across

t"e sa&e ti&e delaysA and $o%el&an .19:5 0 and !c$ee and ,<'ire

.1992 0
fo'nd no difference in for)ettin) bet;een dience%"alic

%atients and t"ose ;it" 9l@"ei&er
de&entia or focal te&%oral

lobe lesions. 7ecent st'dies "ave de&onstrated an a%%arent
difference

bet;een reco)nition &e&ory, in ;"ic" for)ettin) rates in a&nesic

%atients
;ere nor&al once initial learnin) "ad been acco&%lis"ed,

and recall &eas'res in ;"ic"
a&nesic %atients s"o;ed accelerated

for)ettin) over delays of a fe; &in'tes .$o%el&an
and ,tan"o%e

1993 A ("ristensen et al., 199: A Fsaac and !ayes 1999 a, bA *reen

and
$o%el&an, 2002 0. 4o;ever, t"ere ;ere no differences in t"ese

latter st'dies bet;een
%atients ;it" %ri&arily dience%"alic

or te&%oral lobe %at"olo)y .$o%el&an and
,tan"o%e, 1993 A *reen

and $o%el&an, 2002 0. One inter%retation of s'c" findin)s is

t"at t"e &aHor deficit in a&nesia involves ac<'isition or learnin)

%rocesses, since
i&%air&ents in ne; learnin) can be detected

on standard &eas'res of eit"er reco)nition
or recall &e&ory,

b't t"at t"ere is an additional, &ore s'btle i&%air&ent in retention

.or
?consolidation+0 over a %eriod of &in'tes, detectable

only on recall tests .$o%el&an and
,tan"o%e, 1993 , 2001A *reen

and $o%el&an, 2002 0.

)#* +aulty retrie#al
-;o ty%es of retrieval "y%ot"esis "ave been %ost'lated. ?'re+

retrieval "y%ot"eses
%ost'late a retrieval deficit arisin) inde%endently

of any fail're of ?ac<'isition+ %rocesses.
Cor exa&%le,

=arrin)ton and =eis5rant@ %ost'lated t"at a&nesic %atients are

'nable to
s'%%ress ina%%ro%riate res%onses d'rin) recall or

reco)nition tas5s .=arrin)ton and
=eis5rant@, 19L: , 19300. -"ey

noted t"at a&nesic %atients so&eti&es res%ond
erroneo'sly to

&e&ory tests ;it" ;"at "ad been t"e correct re%lies to %revio's

tas5s and,
secondly, t"at t"e %rovision of retrieval c'es can

i&%rove t"eir %erfor&ance. On t"e ot"er
"and, it "as been s"o;n

t"at "ealt"y s'bHects also ex"ibit t"ese %"eno&ena ;"en )iven

recall tests at relatively lon) retention intervals .e.). a

;ee50, s'))estin) t"at t"ey &ay be
a conse<'ence of %oor &e&ory

rat"er t"an its ca'se .=oods and iercy, 193K A !ayes
and !e'dell,

19:1 0. !oreover, restrictin) t"e n'&ber of c"oices in a reco)nition

or c'ed
recall test does not necessarily i&%rove a&nesic %atients+

%erfor&ance relative to controls
.4'%%ert and iercy, 193L A

=arrin)ton and =eis5rant@, 193: 0. Fn recent years, t"e
=arrin)ton

and =eis5rant@ findin) of a beneficial effect of c'es "as been

reinter%reted as
de&onstratin) %reserved %ri&in) in t"e %resence

of i&%aired ex%licit &e&ory .e.).
,"i&a&'ra, 19:L 0, and t"ere

is also evidence t"at intact %ri&in) can %reci%itate
interference

effects ;"en ex%licit &e&ory is i&%aired .!ayes et al., 19:3 0.

9 &odified
retrieval "y%ot"esis stresses evidence t"at retrieval

%rocesses are "eavily de%endent '%on
t"e nat're of initial encodin),

and t"at retrieval deficits arise as a conse<'ence of an
initial

encodin) i&%air&ent .-'lvin) and -"o&%son, 1938 0. !oreover,

a retrieval deficit
&ay be i&%ortant ;"ere t"ere is an extensive

79 .=eis5rant@, 19:5 A $o%el&an, 19:9 A
see belo;0.

In summary, several st'dies %oint to a %roble& in t"e initial

ac<'isition or ?consolidation+
of infor&ation ;it"

a &ore s'btle i&%air&ent in retention .for)ettin)0 detectable

only on
recall testin). -"ere &ay also be a secondary deficit

in retrieval %rocesses. 4o;ever, t"e
%recise nat're of t"is

ac<'isition deficit re&ains ill defined. One %ossibility is

t"at t"ere is
a deficit in ?bindin)+ different ty%es

of &aterial incl'din) context'al infor&ation, and t"at
t"is

relates to vario's distinctions dra;n ;it"in e%isodic &e&ory,

;"ic" ;ill no; be
considered.

$ecall and reco"nition memory
-"ere &ay be a differentiation bet;een recall and reco)nition

&e&ory. On t"e basis of a
&eta-analysis of sin)le case and s&all

)ro'% st'dies of a&nesic %atients, 9))leton and
,"a; .199L 0

clai&ed t"at %atients ;"ose %at"olo)y ;as confined to circ'itry

involvin)
t"e "i%%oca&%i, fornices, &a&illary bodies, &a&illo-t"ala&ic

tract and anterior t"ala&i,
s"o;ed i&%air&ents on recall b't

not reco)nition testin). -"ey ar)'ed t"at in s'c"
%atients,

&e&ory based on fa&iliarity H'd)e&ents .reco)nition0 ;as intact,

;"ereas recall
&e&ory, involvin) recollection of s'c" context'al

feat'res as ti&e and s%atial location,
;as i&%aired .see also

9))leton and Bro;n, 1999 0. -"ey %ost'lated t"at da&a)e to
ot"er

str'ct'res, s'c" as t"e %erir"inal cortex, ;as re<'ired to %rod'ce

an i&%air&ent in
reco)nition &e&ory. 9 %otential %roble& ;it"

t"is sin)le-dissociation is t"at it &i)"t
si&%ly reflect t"e

severity of a&nesia, &ilder a&nesic %atients .;it" less extensive

%at"olo)y0 s"o;in) relatively s%ared %erfor&ance at reco)nition

&e&ory. 9))leton and
,"a; .199L 0 tried to ar)'e a)ainst t"is

by co&%arin) t"e reco)nition &e&ory scores of
.i0 %atients ;it"

?li&bic+ lesions and .ii0 $orsa5offP?ot"er+

a&nesic %atients, ;"ere t"e
revised =ec"sler !e&ory ,cale .=!,-70

scores of t"e t;o )ro'%s did not differ
si)nificantly. 4o;ever,

t"ere ;ere trend differences in recall scores, several of ;"ic"

;ere close to ?floor+, &a5in) any inter%retation

e<'ivocal.

(onsistent ;it" t"e 9))leton "y%ot"esis, Var)"a-$"ade& et al.

.1993 0 described t"ree
%atients ;it" a develo%&ental a&nesia

for everyday events, res'ltin) fro& brain inH'ry in
infancy

or early c"ild"ood. -"ese %atients s"o;ed a %rono'nced loss

of "i%%oca&%al
vol'&e bilaterally, and t"eir ne'ro%syc"olo)ical

test %erfor&ance s"o;ed i&%air&ents on
recall b't not reco)nition

&e&ory, t"e latter bein) tested ;it" &aterial t"at incl'ded

lists
of ;ords, non-;ords, fa&iliar faces and 'nfa&iliar faces.

-"ese findin)s s'))ested t"at,
;"ilst recall of e%isodic &e&ories

.involvin) a te&%oral and s%atial context0 ;as
i&%aired as a

res'lt of t"ese %atients+ "i%%oca&%al %at"olo)y, reco)nition

&e&ory and
se&antic &e&ory ;ere s%ared, beca'se t"ey do not

necessarily involve recollection of
te&%oralPs%atial context1

t"is allo;ed t"ese individ'als to co%e ;it"in &ainstrea&
ed'cation

.see also Baddeley et al., 2001 0.

Fn contrast, ,<'ire and collea)'es "ave s"o;n i&%aired %erfor&ance

in a series of st'dies
on &eas'res of reco)nition &e&ory by

%atients ;"ose %at"olo)y ;as a%%arently li&ited
to t"e "i%%oca&%al

for&ation1 t"eir st'dies incl'ded 29 &eas'res of verbal and

non-
verbal reco)nition &e&ory .yesPno, forced-c"oiceA 7eed and

,<'ire, 1993 0 and t"e
#oors and eo%le test .!anns and ,<'ire,

1999 0. !oreover, Jola et al. .2000 0 s"o;ed
t"at &on5eys ;it"

lesions li&ited to t"e "i%%oca&%al re)ion ;ere i&%aired on t;o

tas5s
of reco)nition &e&ory .b't see Baxter and !'rray, 2001 a,

bA Jola and ,<'ire, 2001 ,
on ani&al findin)s0. Fn "'&an a&nesic

%atients, $o%el&an et al. .2001 0 fo'nd no
obvio's differences

bet;een recall and reco)nition &e&ory tests in t"e si@e of t"eir

correlations ;it" an !7F &eas're of "i%%oca&%al atro%"y.

-"e &aHor difference bet;een t"e Var)"a-$"ade& and ,<'ire findin)s

&ay be in t"e
severity of t"e &e&ory disorder in t"eir res%ective

%atients. -"is is so&e;"at diffic'lt to
deter&ine beca'se different

tests ;ere e&%loyed, b't t"e Var)"a-$"ade& et al. .1993 0
s'bHects

"ad a &ean =ec"sler 9d'lt Fntelli)ence ,caleG7evised .=9F,-70

verbal FQ
&in's =ec"sler .=ec"sler, 19K5 0 &e&ory <'otient discre%ancy

of 11.L3 %oints .,# R
11.150, ;"ereas t"e 7eed and ,<'ire

.1993 0 %atients ;it" "i%%oca&%al lesions "ad a
&ean =9F,-7 FQ

&in's =!,-7 )eneral &e&ory index difference of 88.L3 %oints

.,# R
13.920. Fn t"is connection, %atient O7, re%orted

by !ayes et al. is relevant .!ayes et al.,
2001 , 20020. O7 "ad

bilateral "i%%oca&%al atro%"y and a 8K-%oint =9F,-7 FQ &in's

=!,-7 )eneral &e&ory discre%ancy. 9cross K8 reco)nition &e&ory

tests, O7 did s"o;
si)nificant i&%air&ent relative to controls,

b't t"e i&%air&ent ;as very &inor .&ean z B
M0.50 and

clinically si)nificant .S2 ,#0 in only 10N of tests. Fn contrast,

"er i&%air&ent
on recall tests ;as dis%ro%ortionately severe

.&ean z B M8.L0 and clinically si)nificant in
95N of tests.

In summary, it "as been "y%ot"esi@ed t"at "i%%oca&%al da&a)e

%rod'ces a deficit only in
recall &e&ory, ;"ereas %erir"inal

%at"olo)y i&%licates reco)nition &e&ory as ;ell. -"e
%ossibility

t"at completely s%ared reco)nition &e&ory in t"e "i%%oca&%al

cases si&%ly
reflects a relatively &ild &e&ory i&%air&ent cannot

be excl'ded. 4o;ever, t"ere is no;
at least one case-re%ort

s'))estin) relatie s%arin) of reco)nition &e&ory in a %atient

;it" a fairly severe i&%air&ent of recall &e&ory.

$ememberin" and &nowin", recollection and familiarity
*ardiner and 7ic"ardson-$lave"n .2000 0 "ave distin)'is"ed bet;een

t"e s'bHective
states of ?re&e&berin)+ and ?5no;in)+.

-"ey defined remembering as ?intensely %ersonal
ex%eriences

of t"e %ast, t"ose in ;"ic" ;e see& to recreate %revio's events

and
ex%eriences ;it" t"e a;areness of relivin) t"ese events

and ex%eriences &entally.+
Knowing referred to ?ex%eriences

of t"e %ast, in ;"ic" ;e are a;are of 5no;led)e t"at
;e %ossess

b't in a &ore i&%ersonal ;ay . . . no a;areness of relivin)

. . . DaE )eneral
sense of fa&iliarity . . . DofE facts, ;it"o't

relivin) t"e& &entally.+

Vario's ex%eri&ental st'dies "ave identified a n'&ber of factors

differentiatin)
?re&e&ber+ .70 and ?5no;+

.$0 &e&ories .*ardiner and Iava, 1990 A *ardiner and
ar5in,

1990 A 7aHara&, 1998 0. One iss'e t"at arises is ;"et"er 7 &e&ories

are .i0 a
s%ecial for& of, and "ave t"e %otential to beco&e

$ &e&ories .?red'ndancy+0, .ii0 t"e
o'tco&e of t;o

overla%%in) .?inde%endent+0 %rocesses .one of ;"ic"

is ?5no;in)+0, or
.iii0 t"e &anifestation of an

entirely se%arate %rocess .?excl'sivity+0. 9 second

iss'e is
"o; t"ese s'bHective states .7, $0 relate to &ore obHective

&eas'res of ?recollection+ and
?fa&iliarity+.

Fn a&nesic %atients, $no;lton and ,<'ire .1995 0 fo'nd red'ced

7 and $ &e&ories
.res%onses0 on a reco)nition &e&ory test, relative

to "ealt"y controls. -"e controls+
%erfor&ance 1 ;ee5

later rese&bled t"at of t"e a&nesic %atients 10 &in after sti&'li

%resentation. !oreover, &any of t"e controls+ 7 res%onses

at 10 &in beca&e $ res%onses
at 1 ;ee5, i&%lyin) t"at 7 and

$ &e&ories ;ere eit"er red'ndant or inde%endent, b't not
&'t'ally

excl'sive. -"e a't"ors concl'ded t"at a&nesic %atients s"o;

i&%air&ents in
bot" t"e 7 and $ co&%onents of ex%licit &e&ory,

and t"at $ res%onses cannot be
identified as reflectin) ?i&%licit

&e&ory+.

Ot"ers "ave ar)'ed t"at reco)nition &e&ory H'd)e&ents can be

&ade either on t"e basis
of feelin)s of familiarity, ;"ic" &ay

be relatively %reserved in a&nesia .see above0, or on
t"e basis

of context'al &e&ories .e.). ;"en or ;"ere so&et"in) "a%%ened0

i.e. conscious
recollection .e.). 4'%%ert and iercy, 193: A

Oonelinas, 2001 0. -"ere "ave been vario's
atte&%ts to <'antify

t"ese &e&ory co&%onents1 for exa&%le, Iacoby and collea)'es+

%rocess dissociation %roced're relies on s'bHects+ abilityPinability

to incl'de or excl'de
ite&s ;"ose so'rce or context is recollected

.Iacoby et al., 1998 0. Oonelinas .2001 0 "as
ar)'ed t"at recollection

and fa&iliarity are independent %rocesses ;it" very different

res%onse c"aracteristics. Fn reco)nition &e&ory, recollection

%rod'ces all-or-none ?"i)"
confidence+ res%onses

;"en t"e context'al infor&ation retrieved exceeds a certain

t"res"old. Ca&iliarity %rod'ces a &'c" &ore variable %attern

of res%ondin) accordin) to
trace stren)t" .dT0 and res%onse

bias .!0 .si)nal detection t"eory0, and t"e %attern of t"ese

res%onses ."its 1 false alar&s0 at different levels

of confidence .res%onse bias0 can be
%lotted on a so-called

receiver o%eratin) c"aracteristics .7O(0 c'rve.

Oonelinas et al. .199: 0 e&%"asi@ed t"e i&%ortance of controllin)

for res%onse bias in
bot" 7P$ and %rocess dissociation ex%eri&ents.

-"ey 'sed a si)nal-detection %roced're to
recalc'late findin)s

fro& s'c" st'dies in a&nesia .e.). Verfaellie and -read;ell,

1998 A
,c"acter et al., 199L a0, and t"ey also re%orted t"eir

o;n findin)s %lottin) 7O( c'rves in
a s&all )ro'% of a&nesic

%atients. 9cross all t"ree tec"ni<'es, a&nesia %rod'ced a
%rono'nced

red'ction in recollection &e&ory and a s&aller b't consistent

red'ction in
fa&iliarity res%onses1 t"is s'))ested t"at 7 and

$ are s'bHective states &eas'rin) .or
reflectin)0 recollection

and fa&iliarity and, conversely, t"at obHective &eas'res of

t"e
latter can be 'sed to %redict t"e occ'rrence of t"ese conscio's

states .a %oint contested by
*ardiner, 2001 0.

In summary, t"e evidence s'))ests t"at &e&ories based on ?recollection+

.and its
s'bHective co'nter%art, ?re&e&berin)+0

are severely i&%aired in a&nesia, and t"at t"ose
based on fa&iliarity

H'd)e&ents .or ?5no;in)+0 are also si)nificantly

i&%aired, alt"o')"
less severely so.

Explicit and implicit memory

#es%ite t"eir severe i&%air&ent of ex%licit or e%isodic &e&ory,

&any as%ects of i&%licit
&e&ory are %reserved in a&nesic %atients.

Cor exa&%le, t"ey s"o; intact ac<'isition and
retention of t"e

classically conditioned eye-blin5 res%onse .=eis5rant@ and =arrin)ton,

1939 A #a'& et al., 19:9 0, %rovided no interval is inter%osed

bet;een t"e offset of t"e
conditioned sti&'l's and t"e onset

of t"e 'nconditioned sti&'l's .(lar5 and ,<'ire, 199:
0. 9&nesic

%atients can also s"o; %reserved ac<'isition and retention of

%erce%t'o-
&otor s5ills .%roced'ral &e&ory0 .(or5in, 19L5 A !oscovitc",

19:2 A (er&a5 and
O+(onnor, 19:8 A =ilson and =earin), 1995 0.

Fn contrast, %roced'ral &e&ory is
i&%aired in certain s'bcortical

de&entias s'c" as ar5inson+s disease and 4'ntin)ton+s

disease on, for exa&%le, %'rs'it rotor and t"e ?-o;er

of 4anoi+ tas5s .4eindel et al., 19::
A ,aint-(yr et al.,

19:: 0. !ore recently, 7eber and ,<'ire .1999 0 "ave de&onstrated

t"at s5ill learnin) is not a sin)le entity1 ar5inson %atients

;ere i&%aired at ?"abit
learnin)+, i&%licatin) t"e

neostriat'&, b't s"o;ed intact learnin) of artificial )ra&&ars

and dot %attern %rototy%es, ;"ic" ;ere %ost'lated to reflect

brain re)ions o'tside bot" t"e
neostriat'& and t"e &edial te&%oral

lobes.

ri&in) refers to t"e facilitation of a res%onse in t"e absence

of conscio's a;areness
.ex%licit &e&ory0 t"at t"e sti&'l's "as

occ'rred before. -"is facilitation can occ'r across
%erce%t'ally

or conce%t'ally si&ilarPidentical sti&'li. !any st'dies "ave

de&onstrated
intact %ri&in) in a&nesic %atients on tas5s s'c"

as ste&-co&%letion .e.). *raf et al., 19:K
A ,"i&a&'ra and ,<'ire,

19:K A *raf and ,c"acter, 19:5 0, and 4a&ann and ,<'ire
.1993 0

re%orted f'lly intact %ri&in) in a %rofo'ndly a&nesic %atient

.fro& "er%es
ence%"alitis0, des%ite "is %erfor&in) at c"ance

on reco)nition &e&ory tests. ,'c"
findin)s "ave been inter%reted

as de&onstratin) t"at %ri&in) is not de%endent on
dience%"alicP&edial

te&%oral brain str'ct'res, b't t"at cortical re)ions &'st be

critical
.e.). ,"i&a&'ra, 19:L A ,c"acter, 19:3 A 9lvare@ and

,<'ire, 1995 A 7eber and ,<'ire,
1999 0.

$eane et al. .1995 0 contrasted t"e %erfor&ance of t;o %atients

on &eas'res of ex%licit
and i&%licit &e&ory. -"e first ;as /4,

;"o "ad s'ffered a severe closed "ead inH'ry 28
years earlier1

"e "ad 'nder)one a ri)"t te&%oral lobecto&y and insertion of

a s"'nt for
"ydroce%"al's. Fn addition, "e "ad da&a)e to t"e

ri)"t %arietal and occi%ital lobes and a
left "e&is%"ere ;"ite

&atter lesion, ;"ic" extended fro& t"e inferior te&%oral )yr's

to
H'st belo; t"e occi%ital "orn. -"e &edial te&%oral str'ct'res

;ere s%ared )ross str'ct'ral
da&a)e. -"e second %atient ;as

4! .,coville and !ilner, 1953 A (or5in et al., 1993 0,
;"o beca&e

%rofo'ndly a&nesic follo;in) a bilateral &edial te&%oral lobecto&y.

4is
!7F s"o;ed bilateral %at"olo)y incl'din) t"e &edial te&%oral

%olar cortex, t"e
a&y)dalae and t"e entor"inal cortex. -"e resection

involved a%%roxi&ately t"e anterior 2
c& of t"e "i%%oca&%al

for&ation, and t"ere ;as atro%"y in t"e %osterior 2 c& of t"e

"i%%oca&%al fields. -"e %osterior %erir"inal and %ara"i%%oca&%al

cortices ;ere only
sli)"tly da&a)ed. $eane et al. .1995 0 s"o;ed

t"at /4 ;as intact at &eas'res of
.ex%licit0 reco)nition &e&ory,

b't i&%aired at %erce%t'al %ri&in) .e.). %erce%t'al
identification,

;ord-co&%letion %ri&in)0. Fn contrast, 4! s"o;ed severely i&%aired

reco)nition &e&ory, b't intact %erce%t'al %ri&in). On t"e basis

of t"ese findin)s, $eane
et al. .1995 0 ar)'ed t"at vis'o-%erce%t'al

%ri&in) de%ends on t"e inte)rity of occi%ital
circ'its t"at

;ere co&%ro&ised in /4 b't ;ere %reserved in 4!. Fn contrast,

ex%licit
reco)nition &e&ory is critically de%endent on intact

&edial te&%oral lobes.

9n alternative vie; "as been %ost'lated by Oster)aard, ;"o ar)'ed

t"at, ;"en a tas5 is
easy and s'bHects ac"ieve "i)" levels of

baseline %erfor&ance .%rod'cin) t"e ?correct+
;ords

to c'es in t"e absence of any %ri&es0, t"e %ossible &a)nit'de

of %ri&in) effects is
constrained and &ay not reflect t"e a&o'nt

of infor&ation available fro& t"e st'dy
.?%ri&in)+0

e%isode .Oster)aard, 199K , 199:, 19990. ="en a tas5 ;as &ade

&ore
diffic'lt, a si)nificant correlation bet;een %ri&in) and

ex%licit &e&ory .reco)nition0
scores e&er)ed in "ealt"y s'bHects

.Oster)aard, 199: 0 and a&nesic %atients s"o;ed a
%ri&in) i&%air&ent

not seen on an easier tas5 .Oster)aard, 1999 0. !oreover, t;o

st'dies
e&%loyin) <'antitative str'ct'ral !7F .Ierni)an and

Oster)aard, 1998 A Ierni)an et al.,
2001 0 fo'nd correlations

bet;een i&%aired %ri&in) and &edial te&%oralP"i%%oca&%al
vol'&e

loss in &ixed )ro'%s of a&nesic, de&entin), and "ealt"y control

s'bHects.
Oster)aard and collea)'es "ave concl'ded t"at a&nesic

%atients s"o; i&%aired %ri&in),
related to &edial te&%oral da&a)e,

after controllin) for baseline %erfor&ance
.Oster)aard, 199K,

1999 0. !ore recently, $inder and ,"an5s .2001 0 "ave %ost'lated

t"at a sin)le &e&ory syste& &ay 'nderlie a%%arent ?dissociations+

in %erfor&ance on
reco)nition &e&ory and %ri&in) tas5s, ar)'in)

t"at differential rates of learnin) bet;een
%atients and controls

and differin) tas5 de&ands across %roced'res can ex%lain t"e

%attern
of res'lts obtained.

*oodin) et al. .2000 0 "ave %'t for;ard a t"ird %osition. -"ey

cond'cted a &eta-analysis
of st'dies of i&%licit &e&ory for

fa&iliar or novel infor&ation in a&nesic %atients and
"ealt"y

controls. Fn 8L st'dies involvin) 59 se%arate &eas'res, t"e

%atients and controls
s"o;ed e<'ivalent %ri&in) on tests 'sin)

fa&iliar infor&ation .t"e control )ro'% doin)
better 11 ti&es

o't of 28 investi)ations0. 4o;ever, t"e controls %erfor&ed si)nificantly

better t"an t"e a&nesic %atients on %ri&in) tas5s involvin)

novel ite&s .1: o't of 28
st'dies0 or novel associative infor&ation

.e.). 'nrelated ;ord %airs s'c" as &o'ntainM
sta&%0 .nine

o't of 18 st'dies0. ,o&e;"at si&ilarly, ("'n and "el%s .1999 0

fo'nd t"at
a&nesic %atients s"o;ed nor&al %erce%t'o-&otor .%roced'ral0

s5ill learnin) on a vis'al
searc" tas5, b't 'nli5e "ealt"y s'bHects,

t"ey failed to s"o; additional benefits fro&
context'al c'ein)1

t"e benefits of t"is context'al c'ein) ;ere ?i&%licit+

in t"e "ealt"y
s'bHects beca'se t"ey co'ld not identify t"e

context on s'bse<'ent reco)nition testin).
Cro& s'c" findin)s,

*oodin) et al. .2000 0 concl'ded t"at t"e "i%%oca&%i are essential

for encodin) and storin) .or ?bindin)+0 novel infor&ation

;it" its associates incl'din)
context, or in &a5in) new associations

bet;een establis"ed ite&s, ;"et"er i&%licitly or
ex%licitly.

In summary, t"e conventional ;isdo& is t"at %ri&in) is s%ared

in a&nesia, and t"at
da&a)e to sites of %at"olo)y beyond t"e

&edial te&%oral lobes is re<'ired to %rod'ce
i&%aired %ri&in)

.e.). involvin) occi%ital circ'itry0. 4o;ever, a&nesic %atients

do s"o;
i&%aired %ri&in) in certain ex%eri&ental conditions,

e.). in ?diffic'lt+ tas5s ;"ere
baseline res%ondin)

"as been controlled or in associative learnin). -"is i&%licates

a
contrib'tion of &edial te&%oralPdience%"alic str'ct'res in

%ri&in) in t"ese
circ'&stances.

Do temporal lobe, diencephalic and frontal lobe
amnesias differ(

-esion studies
-"ere "as been considerable interest in ;"et"er or not t"e %attern

of &e&ory deficits
differs across %atients ;it" eit"er &edial

te&%oral, dience%"alic, or frontal %at"olo)y. 9s
already &entioned,

4'%%ert and iercy .1939 0 ar)'ed t"at te&%oral lobe %at"olo)y

)ives
rise to accelerated for)ettin), ;"ereas dience%"alic a&nesia

does not, b't t"is observation
"as )enerally not been s'%%orted.

9n alternative "y%ot"esis is t"at t"e dience%"alon and
&edial

te&%oral lobes differ in t"eir contrib'tion to context &e&ory.

ar5in and ot"ers
ar)'ed t"at dience%"alic lesions %rod'ce lar)er

deficits in te&%oral order &e&ory t"an
does &edial te&%oral

lobe %at"olo)y, ;"ereas s%atial context deficits are lar)er

for t"e
latter )ro'% .ar5in et al., 1990 aA 4'n5in et al., 199K 0.

Ot"er st'dies "ave %rovided
so&e s'%%ort for t"ese findin)s,

b't t"e findin)s are )enerally &'c" less clear-c't
.("alfonte

et al., 199L A $o%el&an et al., 1993 0.

(ontext &e&ory "as also been invo5ed as a %artic'lar role of

t"e frontal lobes. !ilner
.19:2 0 and B'tters et al. .199K 0 fo'nd

i&%air&ents in te&%oral context &e&ory in
%atients ;it" lar)e

frontal lesions, alt"o')" !ilner et al. .1991 0 and $o%el&an

et al.
.1993 0 fo'nd t"at t"is deficit ;as confined to %atients

;it" lesions %enetratin) t"e dorso-
lateral frontal cortical

&ar)ins. O;en et al. .1990 0 described s%atial ;or5in) &e&ory

i&%air&ents follo;in) frontal lobe lesions, and ,c"acter et

al. .19:K 0 re%orted t"at
deficits in &e&ory for t"e so'rce of

infor&ation ;ere correlated ;it" i&%aired
%erfor&ance on tests

of frontalPexec'tive f'nction. !ore )enerally, %at"olo)y to

t"e
frontal lobes "as been t"o')"t to be da&a)in) to e"ecutie

f'nctions s'c" as %lannin), t"e
or)ani@ation of &aterial, &onitorin)

of res%onses, t"e in"ibition of ina%%ro%riate
res%onses, and

&e&ory for context .e.). ,"i&a&'ra et al., 1991 A ,"i&a&'ra,

199K A
Baldo and ,"i&a&'ra, 2002 0. ,t'dies "ave dra;n attention,

for exa&%le, to i&%air&ent
on s'bHect-ordered tas5s .etrides

and !ilner, 19:2 0, release fro& %roactive interference
.!oscovitc",

19:2 0, or as%ects of t"e ?exec'tive+ co&%onent of

;or5in) &e&ory
.Baddeley, 19:L A etrides, 2000 0. -"is co&bination

of %lannin), or)ani@ational and
context &e&ory deficits ;as

considered to ex%lain ;"y, at least in so&e st'dies, %atients

;it" frontal lobe lesions s"o;ed dis%ro%ortionate i&%air&ent

on &eas'res of recall
&e&ory relative to reco)nition .Iano;s5y

et al., 19:9 A ,"i&a&'ra et al., 1990 0.
C'rt"er&ore, t"e abnor&ality

in te&%oral context &e&ory "as been %ost'lated as t"e
basis

for s%ontaneo's confab'lation so&eti&es fo'nd in %atients ;it"

frontal lobe lesions
.see belo;0.

-"ere are vario's %roble&s ;it" a si&%le notion t"at t"e frontal

lobes %lay only a
s%eciali@ed ?exec'tive+ role in

&e&ory. Cirst, it "as beco&e clear t"at t"e li&bic-
dience%"alic

&e&ory syste& also contrib'tes to vario's as%ects of context

&e&ory, as
already indicated. ,t'dies in a&nesic %atients "ave

s"o;n t"at deficits in &e&ory for
te&%oral context .$o%el&an,

19:9 A ar5in et al., 1990 a0, s%atial context .,"o<eirat
and

!ayes, 1991 0 or t"e &odality of infor&ation .ic5erin) et al.,

19:9 0 are often
attrib'table to da&a)e in t"e dience%"alon or

&edial te&%oral lobes, rat"er t"an t"e
frontal cortex. ,econdly,

so&e ani&al st'dies "ave indicated a critical role for t"e frontal

lobes in ?tar)et+ &e&ory .Bac"evalier and !is"5in,

19:L A ar5er and *affan, 199: 0.
-"irdly, ="eeler et al. .1995 0

re-exa&ined t"e effects of frontal lesions on &eas'res of
"'&an

recall and reco)nition &e&ory. Fn a &eta-analysis of t"e literat're,

t"ey fo'nd t"at,
in virt'ally all st'dies, frontal lesion %atients

;ere i&%aired on tests of recall &e&ory,
relative to a)e-&atc"ed

"ealt"y controls, even if t"e decre&ent did not act'ally reac"

statistical si)nificance in all individ'al st'dies. !oreover,

t"ere ;as a .lesser0 de)ree of
i&%air&ent in frontal lesion

%atients on tests of reco)nition &e&ory in t"e vast &aHority

of st'dies. Co'rt"ly, individ'al %atients "ave been described,

in ;"o& frontal lesions
"ave %rod'ced a dis%ro%ortionate i&%air&ent

in reco)nition &e&ory, res'ltin) fro& a
"i)" rate of false-%ositive

errors, rat"er t"an in recall &e&ory, ;"ic" is nor&ally &ore

de&andin) of ?exec'tive+ reso'rces .#elbec<-#ero'esnU

et al., 1990 A ar5in et al., 199L
A ,c"acter et al., 199L b0.

Collo;in) s'c" observations, $o%el&an and ,tan"o%e cond'cted

a series of
investi)ations co&%arin) )ro'%s of &e&ory-disordered

%atients, ;"ose %ri&ary
%at"olo)y ;as eit"er in t"e frontal

lobes, t"e dience%"alon or t"e te&%oral lobes.
9lt"o')" definite

differences ;ere fo'nd bet;een t"e frontal and ot"er )ro'%s,

t"ese
;ere relatively s'btle, and t"ere ;as considerable overla%

in t"e )ro'%s+ %atterns of
&e&ory i&%air&ent. !e&ory-disordered

%atients ;it" frontal lobe lesions %erfor&ed
si&ilarly to ot"er

a&nesic %atients in &a5in) fre<'ency H'd)e&ents .,tan"o%e et

al.,
199: 0 and in t"eir overall %erfor&ance at recall and reco)nition

&e&ory &eas'res
.$o%el&an and ,tan"o%e, 199: 0, alt"o')" t"eir

scores ;ere en"anced by t"e %rovision
of ?bloc5ed+

cate)ory c'es in a ;ay t"at ;as not seen in t"e ot"er a&nesic

%atients
.$o%el&an and ,tan"o%e, 199: 0. ,econdly, %atients ;"ose

frontal lesions %enetrated t"e
dorso-lateral cortex de&onstrated

severe i&%air&ent on a &eas're of te&%oral context
&e&ory, b't

not on a &eas're of s%atial context &e&ory, ;"ereas ot"er %atients

;it"
lar)e frontal lesions s"o;ed intact %erfor&ance on bot"

&eas'res .$o%el&an et al., 1993
0. -"irdly, dience%"alic and

te&%oral lobe %atients s"o;ed accelerated for)ettin),
relative

to controls, on &eas'res of recall &e&ory, ;"ereas frontal lesion

%atients s"o;ed
only a non-si)nificant trend in t"e sa&e direction

.$o%el&an and ,tan"o%e, 1993 A b't
also see Ietter et al., 19:L 0.

Co'rt"ly, t"e frontal %atients s"o;ed i&%air&ent on t;o
&eas'res

of re&ote &e&ory .recall of a'tobio)ra%"ical incidents and of

fa&o's ne;s
events0, co&%arable in severity to ot"er a&nesic

%atients, b't .'nli5e t"e latter )ro'%0
t"ey ;ere not i&%aired

in t"e retrieval of ;ell-re"earsed %ersonal facts. Fn t"e latter

case,
acc'rate retrieval %res'&ably de&ands less active or ?effortf'l+

reconstr'ction %rocesses.

.euroima"in" in#esti"ations
C'nctional ne'roi&a)in) st'dies "ave been revie;ed else;"ere

.e.). 7')), 2002 0, and it
is beyond t"e sco%e of t"is %a%er

to %rovide a detailed acco'nt. One of t"e earliest
investi)ations

.*rasby et al., 1998 0 %ointed to t"e stren)t" of frontal activations

in
&e&ory encodin) and retrieval. -"e 4279 ."e&is%"eric encodin)

and retrieval
asy&&etry0 "y%ot"esis .-'lvin) et al., 199K a0

;as based on findin)s t"at t"e left frontal
re)ion ;as %artic'larly

involved in t"e encodin) of e%isodic &e&ories, ;"ereas t"e ri)"t

frontal re)ion, to)et"er ;it" t"e %rec'ne's, ;as of %artic'lar

i&%ortance in e%isodic
&e&ory retrieval .,. $a%'r et al., 199K A

,"allice et al., 199K A -'lvin) et al., 199K bA
Cletc"er et al.,

1995 0. Fn contrast, a &ore recent f'nctional !7F .f!7F0 st'dy

"as
%rod'ced evidence of traditional leftMri)"t &aterial-s%ecific

asy&&etries d'rin) encodin)
in bot" t"e &edial te&%oral lobes

and t"e %re-frontal cortex .*olby et al., 2001 0. 9s in
lesion

st'dies, it "as beco&e i&%ortant to differentiate t"e role of

frontal and &edial
te&%oral str'ct'res in e%isodic &e&ory encodin)

and retrieval.

Cletc"er and 4enson .2001 0 "ave revie;ed evidence for a s'bclassification

of frontal
f'nctionin). -"ey ar)'ed t"at ventro-lateral frontal

activation is associated ;it" t"e
'%datin)P&aintenance of infor&ation,

dorso-lateral activation ;it" t"e
selectionP&ani%'lationP&onitorin)

of t"at infor&ation, and anterior activation ;it" t"e
selection

of %rocessesPs'b)oals. On t"e ot"er "and, 7')) and =ildin) .2000 0

%ost'lated
fo'r different classes of retrieval %rocess b't ar)'ed

t"at, to date, t"ere is little evidence to
fractionate t"eir

ne'ral correlates.

Ot"er st'dies "ave exa&ined &edial te&%oral lobe activation.

Cor exa&%le, ,c"acter et
al. .199Lc 0 fo'nd t"at "i%%oca&%al

activation ;as associated ;it" t"e s'ccessf'l
retrieval of e%isodic

&e&ories, ;"ereas left %refrontal activation ;as associated

;it" less
s'ccessf'l b't ?effortf'l+ retrieval.

Bre;er et al. .199: 0 fo'nd t"at %ara"i%%oca&%al
activation and

a re)ion of ri)"t dorsolateral frontal activation ;ere %redictive

of
s'bse<'ent re&e&berin) of %ict'res. *abrieli et al. .1993 0

fo'nd t"at encodin) of novel
infor&ation activated %osterior

&edial te&%oral re)ions, ;"ereas s'ccessf'l retrieval ;as
associated

;it" &ore anterior &edial te&%oral activations. 4o;ever, on

t"e basis of &eta-
analyses, /e%a)e et al. .199: 0 ar)'ed t"at

anterior &edial te&%oral activations ;ere
associated ;it" &e&ory

encodin) and posterior activations ;it" retrieval, ;"ereas
,c"acter

and =a)ner .1999 0 fo'nd t"at encodin) activations ;ere associated

;it" both
anterior and %osterior activations. =a)ner et al.

.199: 0 "ave ar)'ed t"at t"e
%ara"i%%oca&%al )yri en)a)e )eneral

encodin) &ec"anis&s, and t"at t"e left %refrontal
re)ions &ay

contrib'te to t"e or)ani@ation of event attrib'tes in ;or5in)

&e&ory, servin)
as in%'t to t"e &edial te&%oral &e&ory syste&.

$o%el&an et al. .199: b0 fo'nd t"at
verbal encodin) of ne; sti&'li

;as associated ;it" bot" a left "i%%oca&%al and a left
%refrontal

activation, b't t"at, as incre&ental learnin) to re%eated sti&'li

occ'rred, t"e
left &edial te&%oral activation re&ained b't t"e

left frontal activation disa%%eared.
7e%eated %resentation of

sti&'li ;as associated ;it" activation in a ri)"t %refrontal

and
%rec'ne's ?retrieval+ circ'it, s'))estin) t"at

?consolidation+ of learnin) en)a)es bot" t"e
ri)"t

"e&is%"ere circ'it .to retrieve %revio'sly learned &aterial0

and t"e left &edial
te&%oral re)ion .d'rin) incre&ental learnin)0.

Ot"er st'dies "ave indicated t"at &edial
te&%oral activation

&ay be %artic'larly related to associative encodin). -"is "as

been
fo'nd 'sin) f!7F and verbal sti&'li .!ayes et al., 199: 0,

,2(- and %ict'res of
co&%lex scenes .!ontaldi et al., 199: 0,

and t"e encodin) of inter-ite& associations
bet;een %ict'res

or ;ords 'sin) 2- .4en5e et al., 1993 0.

Summary
7ecent lesion and ne'roi&a)in) st'dies indicate a closer interaction

and overla% in
f'nction t"an ;as %revio'sly ass'&ed bet;een

t"e frontal lobe and &edial
te&%oralPdience%"alic str'ct'res

en)a)ed in &e&ory for&ation. ="ilst a broad
distinction bet;een

exec'tive %rocesses and encodin)Pretrieval &ec"anis&s re&ains

valid, t"ere is considerable overla% in t"e effect of focal

lesions involvin) t"ese brain
re)ions, and in t"e brain activations

associated ;it" s%ecific &e&ory tas5s. 4o;ever,
lesion st'dies

indicate t"at dien ce%"alicP&edial te&%oral %atients s"o; faster

for)ettin)
on recall tas5s t"an frontal %atients, and are less

li5ely to res%ond to t"e %rovision of
s%ecific cate)ory c'es,

s'))estin) a &ore f'nda&ental a&nesic deficit. ,i&ilarly, so&e

f'nctional i&a)in) st'dies s'))est t"at &edial te&%oral activations

are &ore closely
related to s'ccessf'l or incre&ental re&e&berin)

t"an are frontal activations. /esion
st'dies indicate t"at t"ere

&ay be differences bet;een s%ecific sites of frontal %at"olo)y

on te&%oral context &e&ory and in effects '%on recall and reco)nition

&e&oryA in
f'nctional i&a)in), dissociations in frontal activation

%atterns are c'rrently bein)
ex%lored.

ow independently semantic is semantic memory(

-"e distinction bet;een se&antic and e%isodic &e&ory ;as &ade

by -'lvin) .1932 0,
alt"o')" it "ad "istorical %recedents .e.).

*illes%ie, 1983 0. ,e&antic &e&ory is t"e
syste& ;"ic" %rocesses,

stores, and retrieves infor&ation abo't t"e &eanin) of ;ords,

conce%ts and facts. Ft is affected in se&antic de&entia, a variant

of fronto-te&%oral
de&entia, and also in "er%es ence%"alitis,

9l@"ei&er de&entia, and occasionally in "ead
inH'ry. Ft see&s

to be rarely affected in cerebro-vasc'lar disorders .?stro5e+0

beca'se of
d'al blood s'%%ly fro& t"e &iddle and %osterior cerebral

arteries to t"e critical re)ions in
t"e left infero-lateral

te&%oral lobe .atterson and 4od)es, 1995 , 20000.

!ate"ory/specific deficits
(ate)ory-s%ecific i&%air&ents are fre<'ently re%orted in se&antic

&e&ory disorders.
=arrin)ton and ,"allice .19:K 0 de&onstrated

t"at, in "er%es ence%"alitis %atients,
identification of livin)

t"in)s and foods ;as severely i&%aired, relative to t"e %atients+

ability to identify inani&ate obHects, and t"at t"is ;as inde%endent

of t"e &odality of
%resentation. Ot"er dissociations "ave also

been %ost'lated1 for exa&%le, bet;een
%"ysical obHects and t"eir

;ord for&s, abstract and concrete ;ords, "i)" and lo;
fre<'ency

;ords, no'ns and verbs, and content and f'nction ;ords .=arrin)ton

and
,"allice, 19:K A (ara&a@@a, 199: 0.

,everal %roble&s "ave arisen. On t"e one "and, not all st'dies

"ave controlled ade<'ately
for %ro%erties s'c" as ;ord fre<'ency

and fa&iliarity, as %ointed o't by C'nnell and
,"eridan .1992 0

and ,te;art et al. .1992 0, alt"o')" t"e &ore recent st'dies

"ave done
so .e.). /a&bon 7al%" et al., 199: A !oss et al., 199: 0.

,econdly, 'n's'al dissociations
.e.). %atients ;"o are severely

i&%aired in %rocessin) ani&als b't not ot"er livin) t"in)s
s'c"

as fr'its and ve)etables0, "i)"ly s%ecific i&%air&ents .e.).

confined to fr'its and
ve)etables0, or reversed dissociations

.ani&als intact, inani&ate obHects i&%aired0 "ave
all been re%orted

.=arrin)ton and !c(art"y, 19:3 A (ara&a@@a, 199: 0. !ost
%artic'larly,

t"ere is little a)ree&ent abo't t"e interpretation of t"ese

cate)ory-s%ecific
i&%air&ents. Ft "as been s'))ested t"at livin)

t"in)s &ay be %ri&arily %rocessed in ter&s
of t"eir sensory

%ro%erties, ;"ereas inani&ate obHects are %rocessed &ainly in

ter&s of
t"eir f'nction .e.). =arrin)ton and ,"allice, 19:K A

Basso et al., 19:: A Bor)o and
,"allice, 2001 0. 4o;ever, !oss

et al. .199: 0 re%orted a %atient in ;"o& t"e vis'al and
f'nctional

attrib'tes of livin) t"in)s ;ere i&%licated e<'ally in a na&in)

tas5, ;"ereas
bot" ty%es of attrib'te ;ere relatively s%ared

;it" res%ect to obHects. !any alternative
t"eories "ave been

%ro%osed. Not only is t"ere disa)ree&ent abo't t"e i&%aired

%rocesses
'nderlyin) cate)ory-s%ecific deficits b't, &ore )enerally,

abo't ;"et"er t"ey %rovide
'na&bi)'o's %roof of 'nderlyin) brain

&ec"anis&s dedicated to t"ose cate)ories
.(ara&a@@a, 199: 0.

9n alternative is t"at cate)ory-s%ecific deficits are si&%ly

an
?e&er)ent %ro%erty+ of differences in t"e str'ct're

and content of conce%ts t"e&selves
.-yler and !oss, 2001 0.

,o&e f'nctional i&a)in) st'dies "ave %rovided so&e s'%%ort for

a ne'robiolo)ical basis
to cate)ory s%ecificity. !artin and

collea)'es fo'nd differin) %atterns of activation for
)eneratin)

colo'r ;ords, action ;ords and for& ;ords .!artin et al., 1995 ,

199L A ("ao
et al., 1999 0. #ifferent classes of ite& ;ere associated

;it" activation of differin)
net;or5s in discrete cortical re)ions1

identifyin) andPor na&in) ani&als ;as associated
;it" activations

in t"e lateral f'sifor& )yr's, &edial occi%ital cortex and s'%erior

te&%oral s'lc's, ;"ereas identifyin) andPor na&in) tools ;as

associated ;it" activations
in t"e &edial f'sifor& )yr's, left

&iddle te&%oral )yr's and left %re&otor cortex. ,o&e
of t"ese

brain re)ions overla%%ed ;it" t"ose t"at are atro%"ied in %atients

;it" cate)ory-
s%ecific deficits .!oss et al., 199: A !oss and

-yler, 2000 0. On t"e ot"er "and, #evlin
et al. .2002 0 identified

a ne'ral syste& co&&on to livin) t"in)s and obHects, b't failed

to
find rob'st evidence of f'nctional se)re)ation by do&ain

or cate)ory.

Semantic dementia
=arrin)ton .1935 0 described t"ree cases of a de)enerative disorder,

selectively affectin)
se&antic &e&ory. ,no;den et al. .19:9 0

coined t"e ter& ?se&antic de&entia+ to
describe

so&e si&ilar cases, and t"is ter& ;as s'bse<'ently ado%ted by

4od)es et al.
.1992 0 and ot"ers. ,e&antic de&entia is really

a ?te&%oral lobe+ variant of fronto-
te&%oral de&entia

or de)eneration .,no;den et al., 199L bA 4od)es et al., 199: 0.

Ft is
c"aracteri@ed by a %ro)ressive disorder of se&antic 5no;led)e1

t"is involves a %rofo'nd
loss of &eanin), enco&%assin) verbal

and non-verbal &aterial and res'ltin) in severe
i&%air&ents

in na&in) and ;ord co&%re"ension. ,%eec" beco&es increasin)ly

e&%ty
and lac5in) in s'bstantives, b't o't%'t is fl'ent, effortless,

)ra&&atically correct and free
fro& %"one&ic %ara%"asias. erce%t'al

and reasonin) abilities are intact, and e%isodic
&e&ory is relatively

%reserved .,no;den et al., 19:9 , 199LaA 4od)es et al., 1992A

!'rre et al., 2001 0. ri&ary %ro)ressive non-fl'ent a%"asia

is anot"er disorder res'ltin)
fro& focal te&%oral lobe atro%"y

b't in t"is, 'nli5e se&antic de&entia, s%eec" o't%'t is
non-fl'ent,

co&%re"ension is relatively %reserved .at least initially0,

and t"ere are fre<'ent
%"one&ic errors .,no;den et al., 199L b0.

On !7F, se&antic de&entia %atients c"aracteristically s"o; severe

atro%"y of t"e inferior
and lateral te&%oral )yri ;it" relative

%reservation of &edial te&%oral str'ct'res
.,no;den et al.,

19:9 A 4od)es et al., 19920. 9 recent st'dy 'sin) voxel-based

&or%"o&etry fo'nd %rono'nced atro%"y of t"e left te&%oral %ole

and left antero-lateral
te&%oral lobe re)ion .!'&&ery et al.,

1999 0. 9 2- activation st'dy, exa&inin) a
se&antic decision

tas5 relative to a vis'al decision tas5, s"o;ed red'ced activity

in t"e
left %osterior inferior te&%oral )yr's1 t"is ;as inter%reted

as reflectin) disr'%ted te&%oral
lobe connections .!'&&ery et

al., 1999 0. -"e ri)"t te&%oral lobe is 's'ally affected to
a

lesser .b't variable0 de)ree, and it is %redo&inantly affected

in cases in ;"o& t"e &ost
%ro&inent %roble&s are in vis'al se&antics

.2vans et al., 1995 0. 9lt"o')" t"e %ri&ary
site.s0 of cortical

%at"olo)y are consistent across st'dies, t"ere is variability

in t"e
"isto%at"olo)ical findin)s, ;"ic" &ay involve s%on)ifor&

c"an)es and ne'ronal loss
.,no;den et al., 199L b0, ic5 cells

and bodies .4od)es et al., 199: 0, or 'bi<'itin-
%ositive ta'-ne)ative

incl'sion bodies identical to &otor ne'rone disease b't ;it"o't

involve&ent of brainste& or s%inal cord &otor ne'rones .7ossor

et al., 2000 0.

Fn se&antic de&entia, =arrin)ton .1935 0 fo'nd t"at 5no;led)e

of s'bordinate
cate)ories .e.). t"e na&e of a s%ecific ani&al0

;as &ore i&%aired t"an 5no;led)e of
s'%erordinate cate)ories

.e.). ani&als or birds0. ,i&ilar findin)s ;ere obtained by
,no;den

et al. .19:9 0 and 4od)es et al. .19920. 4od)es et al. .1995 0

&onitored t"e
deterioration of a sin)le %atient over t"e co'rse

of 2 years, findin) t"at t"e %attern of
relative %reservation

of s'%erordinate 5no;led)e ;as de&onstrable bot" d'rin) a
%artic'lar

test session and across test sessions lon)it'dinally. 4o;ever,

over t"e co'rse of
ti&e, t"ere ;as a %ro)ression fro& s'bordinate

to s'%erordinate errors. =arrin)ton .1935
0 "ad inter%reted t"is

%attern of deterioration as reflectin) t"e "ierarc"ical &anner

in
;"ic" 5no;led)e is or)ani@ed .co&%are (ollins and Q'illian,

19L9 0, b't 4od)es et al.
.1995 0 ar)'ed t"at s'%erordinate ;ords

"ave &'lti%le connections ;it"in t"e se&antic
net;or5, and conse<'ently

are less v'lnerable to de)radation, a vie; t"at does not
necessarily

i&%ly an "ierarc"ical str'ct're to t"e or)ani@ation of 5no;led)e.

9 t"ird vie;,
advocated by C'nnell .1995 0, is t"at bot" s'%erordinate

and s'bordinate 5no;led)e are
affected, after controllin) for

;ord fre<'ency and fa&iliarity effects, s'))estin) t"at t"e

a%%arent s%arin) of s'%erordinate 5no;led)e is an artefact of

factors s'c" as ;ord
fre<'ency and fa&iliarity. Ot"er classes

of 5no;led)e, e.). n'&ber and t"e ability to
%erfor& &ental

calc'lations, &ay be s'r%risin)ly ;ell %reserved des%ite t"e

severe
disr'%tion of verbal and vis'al se&antics in t"is disorder

.B'tter;ort" et al., 2001 A
(a%%elletti et al., 2001 0.

Interaction of semantic and episodic memory in semantic dementia
Fn a series of %'blications, ,no;den and collea)'es "ave %ointed

to a %'tative interaction
bet;een a'tobio)ra%"ical ex%erience

and se&antic 5no;led)e .,no;den et al., 199K ,
1995, 199La0.

-"ey fo'nd t"at %atients ;it" se&antic de&entia reco)ni@ed t"e

na&es of
%ersonal ac<'aintances better t"an t"ose of celebrities,

%ersonally fa&iliar %lace na&es
better t"an "i)" fre<'ency,

i&%ersonal %lace na&es, and %ersonal obHects better t"an
alternative

exa&%les of t"e sa&e obHect .,no;den et al., 199K 0. (o&%re"ension

of
;ords in conversation tended to be li&ited to t"ose as%ects

of &eanin) relevant to t"e
%atient+s o;n life, and definitions

)iven by a se&antic de&entia %atient "ad a &ar5edly
a'tobio)ra%"ical

<'ality .,no;den et al., 1995 0. C'rt"er&ore, ,no;den et al.

.199L a0
fo'nd t"at s'c" %atients+ 5no;led)e ;as better

for conte&%orary t"an for %ast or
"istorical celebrities, for

t"e conte&%orary deci&al &onetary syste& co&%ared ;it" t"e
old

.i&%erial0 Britis" &onetary syste&, and for recent %ersonal

facts and incidents
co&%ared ;it" earlier ones .see belo;0.

,no;den and collea)'es concl'ded t"at c'rrent
a'tobio)ra%"ical

ex%erience interacts ;it" and facilitates t"e &aintenance andPor

retrieval
of resid'al se&antic 5no;led)e in se&antic de&entia

%atients .,no;den et al., 199K ,
1995, 199LaA ,no;den, 2002 0.

#escri%tions of t;o &ore %atients "ave %rovided f'rt"er
evidence

of a'tobio)ra%"ical c'es facilitatin) se&antic 5no;led)e in

se&antic de&entia
%atients .C'nnell, 2001 A =est&acott et al.,

2001 0.

*ra"a& and collea)'es "ave contested t"is vie; .*ra"a& et al.,

1993 , 1999, 20000.
-"ey exa&ined 5no;led)e of )olf and bo;ls

in t;o %atients ;it" se&antic de&entia,
;"o ;ere re)'lar %artici%ants

in t"ese s%orts. *ra"a& et al. .1993 0 failed to find a direct

effect of a'tobio)ra%"ical ex%erience on %revio'sly learned

5no;led)e of t"ese s%orts,
alt"o')" one of t"eir %atients s"o;ed

so&e %reserved 5no;led)e of %eo%le s"e sa;
fre<'ently. *ra"a&

and collea)'es ar)'ed t"at t"is 5no;led)e ;as &ediated by t"e

"i%%oca&%i, and t"ey c"aracteri@ed it as ?se&antic-li5e+

and ?"i)"ly a'tobio)ra%"ically
constrained+ .*ra"a&

et al., 1993 , 19990. Fn re%ly, ,no;den et al. .1999 0 ar)'ed

t"at
t"is %reserved 5no;led)e is indeed ?se&antic+,

albeit i&%overis"ed, and t"at ;"at
deter&ines ;"et"er %revio'sly

establis"ed 5no;led)e is available is not t"e ti&e at ;"ic"
it ;as ac<'ired .as *ra"a& et al., 1993 , "ad s'))ested0 b't

its relevance to
conte&%orary a'tobio)ra%"ical ex%erience.

Dissociation of semantic and episodic memory(
(onsistent ;it" t"e ar)'&ent of *ra"a& et al. .1993 0, $itc"ener

et al. .199: 0 %rovided
f'rt"er evidence for t"e a%%arent se%aration

of t"e ne'robiolo)ical syste&s 'nderlyin)
se&antic and e%isodic

&e&ory .co&%are Var)"a-$"ade& et al., 1993 A Verfaellie et al.,

2000 0. -"ey described a severely a&nesic K9-year-old &an, ;"ose

!7F s"o;ed
co&%lete destr'ction of t"e left "i%%oca&%'s, %ara"i%%oca&%al

)yr's, entor"inal and
%erir"inal cortex. -"ere ;as isc"ae&ia

in t"e left t"ala&'s, and infarction involvin) t"e
left &edial

frontal lobe and basal forebrain. -"ere ;as also a circ'&scribed

area of
infarction in t"e ri)"t %osterior "i%%oca&%'s and %artial

atro%"y in t"e re&ainder of t"e
ri)"t "i%%oca&%'s. -"ere ;as

relative %reservation of infero-lateral te&%oral neocortex.

-"e %atient s"o;ed severe i&%air&ent across a ran)e of standard

e%isodic &e&ory tests.
4o;ever, t"e %atient did s"o; so&e resid'al

5no;led)e on tests involvin) reco)nition
&e&ory for %ersonal

events or %ersonally 5no;n individ'als, and on &ore se&antic

tas5s
involvin) fa&o's faces, events, na&es, or %re&orbidly

ac<'ired vocab'lary. Cro& t"eir
findin)s, t"e a't"ors ar)'ed

t"at ne; se&antic infor&ation can be ac<'ired in t"e absence
of ?any+ si)nificant antero)rade e%isodic &e&ory.

-"ey %ost'lated t"at t"is occ'rs by
&eans of a ?slo;-learnin)+

syste& in t"e .non-&edial0 te&%oral neocortex as a res'lt of

re%eated %resentation of t"e relevant infor&ation. 9lt"o')"

so&e of t"e a't"ors+ findin)s
;ere stri5in) .for exa&%le,

t"e %atient did not 5no; t"at "is da')"ter "ad )ro;n '% in t"e
last 18 years, des%ite 5no;in) ;"o 7onald 7ea)an ;as0, t"eir

inter%retation "as to be
ta5en ;it" ca'tion. -"e a't"ors co&%ared

a very severely i&%aired e%isodic &e&ory
;it" a better b't still

very %oor se&antic &e&ory, and t"ere ;as so&e %reservation of

t"e
ri)"t &edial te&%oral str'ct'res. ,o&e of t"e tests in ;"ic"

t"e %atient %erfor&ed
relatively ;ell .fa&o's faces, fa&o's

ne;s events0 are t"ose in ;"ic" ot"er st'dies "ave
s"o;n an

i&%ortant ri)"t te&%oral lobe contrib'tion .$itc"ener et al.,

1999 A $o%el&an
et al., 1999 0.

Summary
,e&antic &e&ory is severely affected in se&antic de&entia, and

is also affected in
disorders s'c" as "er%es ence%"alitis and

9l@"ei&er de&entia. (ate)ory-s%ecific effects
and relative %reservation

of s'%erordinate 5no;led)e "ave co&&only been re%orted in
t"ese

disorders. 4o;ever, t"e 'nderlyin) nat're of t"ese %"eno&ena

re&ains
controversial, as do t"e de)ree of se&anticMe%isodic

inde%endence in focal lesions and
se&anticMe%isodic interaction

in se&antic de&entia.

What determines the pattern and extent of
retro"rade memory loss(

Temporal "radients
7ibot .1::2 0 ar)'ed t"at1 ?-"e %ro)ressive destr'ction

of &e&ory follo;s a lo)ical
order M a la; . . . it be)ins

at t"e &ost recent recollections ;"ic", bein) . . . rarely
re%eated

and . . . "avin) no %er&anent associations, re%resent or)ani@ation

in its feeblest
for&.+ ,ince 1931, 7ibot+s ?la;+

"as been tested across a n'&ber of different tas5s,
&eas'rin)

5no;led)e of fa&o's %eo%lePfaces, ne;s events and ot"er %'blic

infor&ation,
facts abo't oneself .%ersonal se&antic &e&ory0,

and ?a'tobio)ra%"ical+ incidents .events0
fro& a

%erson+s %ast.

,anders and =arrin)ton .1931 0 ad&inistered tests of fa&o's faces

and %'blic ne;s
events to five a&nesic %atients, incl'din) t"ree

alco"olic $orsa5off %atients. On recall
and reco)nition versions

of t"eir fa&o's faces test, t"e a&nesic )ro'% ;as severely
i&%aired

co&%ared ;it" "ealt"y s'bHects, b't on neit"er test ;as t"ere

any clear evidence
of relative s%arin) of early &e&ories, 5no;n

as a ?te&%oral )radient+. 4o;ever,
s'bse<'ent st'dies

"ave fo'nd s'c" a )radient. !arslen-=ilson and -e'ber .1935 0

obtained only &ild i&%air&ent in $orsa5off %atients on a fa&o's

faces test coverin) t"e
1920s to t"e 1950s, b't severe i&%air&ent

on t"e faces fro& t"e 19L0s. 9lbert and
collea)'es .9lbert et

al., 1939 , 19:1 0 obtained evidence of a te&%oral )radient
extendin)

bac5 80 years .fro& t"e 1930s to t"e 19K0s0 and (o"en and ,<'ire

.19:1 0 for
t"e &ost recent t;o decades. On a &eas're of t"e

recall of a'tobio)ra%"ical incidents,
re<'irin) s'bHects to

retrieve &e&ories related to a c'e-;ord and to date t"e& .t"e

?(rovit@ test+A (rovit@ and ,c"iff&an, 193K 0, Jola-!or)an

et al. .19:8 0 fo'nd t"at
$orsa5off %atients "ad to delve &'c"

f'rt"er bac5 into t"eir re&ote %ast in order to
retrieve a'tobio)ra%"ical

&e&ories .&ean B 80.K years0 t"an did co&%arison )ro'%s of
non-$orsa5off

alco"olics .&ean B 20.1 years0 or "ealt"y controls .&ean B 12.3

years0.

Fn )eneral, t"e earlier st'dies in $orsa5off %atients did not

doc'&ent t"e d'ration fro&
t"e =ernic5e e%isode 'ntil t"e ti&e

of testin), so t"at it ;as diffic'lt to se%arate o't t"e
?retro)rade+

and ?antero)rade+ contrib'tions to ?recent+

&e&ory i&%air&ent. 4o;ever, in
t"e $o%el&an .19:9 0 st'dy, t"e

&ean d'ration of illness ;as doc'&ented as 5.35 years,
and t"ere

;as a retro)rade co&%onent to t"e $orsa5off %atients+

a&nesia of 15M25 years.
Fn addition, st'dies in $orsa5off

%atients "ave s"o;n t"e follo;in)1 .i0 t"e severity of 79
is

not ;ell correlated ;it" t"e severity of 99 .,"i&a&'ra et al.,

19:L A $o%el&an, 19:9
A ar5in, 1991 A !ayes et al., 199K 0A .ii0

t"e i&%air&ent in recallin) recent &e&ories
cannot be attrib'ted

to a s%ecific deficit in encodin) or retrievin) context'al infor&ation

.ar5in et al., 1990 b0A and .iii0 t"ere is also 79 ;it" a te&%oral

)radient for &ore %'rely
se&antic infor&ation, e.). 5no;led)e

of ;ords ;"ic" entered t"e lan)'a)e at different
ti&es .Verfaellie

et al., 1995 b0.

Ot"er )ro'%s of a&nesic %atients s"o; eit"er a te&%oral )radient

co&%arable ;it" t"at
seen in $orsa5off %atients .,<'ire et al.,

19:9 0 or a so&e;"at flatter or ?)entler+
)radient

.$o%el&an, 19:9 A $o%el&an et al., 1999 0. 2ven in 9l@"ei&er %atients,

t"ere is
)enerally a ?)entle+ te&%oral )radient

across a n'&ber of different tas5s ;it" relative
s%arin) of

early &e&ories .Beatty et al., 19:: A ,a)ar et al., 19:: A $o%el&an,

19:9 A
*reene et al., 1995 A *reene and 4od)es, 199L 0. !ore controversial

is ;"et"er t"ere is
a te&%oral )radient in %atients ;it" 4'ntin)ton+s

de&entia or frontal lobe %at"olo)y.
9lbert et al. .19:1 0 and

Beatty et al. .19:: 0 failed to obtain a te&%oral )radient in

4'ntin)ton %atients, as did !an)els et al. .199L 0 on recall

tests in %atients ;it" frontal
lobe lesions. On t"e ot"er "and,

$o%el&an et al. .1999 0 fo'nd a statistically si)nificant
te&%oral

)radient on a ne;s events test in frontal %atients, as did #+2s%osito

et al. .199L
0 on a fa&o's faces test, and *ade and !ortensen

.1990 0 obtained near-si)nificance .P
B 0.0580 on a %'blic events

test.

In summary, &any st'dies "ave %rod'ced evidence of a te&%oral

)radient, consistent ;it"
7ibot+s ?la;+. 4o;ever,

t"e stee%ness of t"at )radient a%%ears to differ across %atient

)ro'%s, )enerally bein) )entler in %atients ;it" de&entia or

frontal lobe atro%"y t"an in
%atients ;it" %'rer a&nesic syndro&es

fro&, for exa&%le, "y%oxia or t"e $orsa5off
syndro&e. !oreover,

differin) tas5 de&ands &ay %rod'ce so&e;"at varyin) %atterns

of
%erfor&ance1 retrievin) a'tobio)ra%"ical &e&ories in an ?'nconstrained+

&anner to a
c'e-;ord in t"e (rovit@ test .,a)ar et al., 19:: 0,

%rovidin) a ?free narrative+ of
i&%ortant events

fro& a %erson+s life .Cro&"olt and /arsen, 1991 , 19920,

or retrievin)
a'tobio)ra%"ical &e&ories ;"en constrained to

%artic'lar ti&e-%eriods .$o%el&an, 19:9
A !ac$innon and ,<'ire,

19:9 0 &ay eac" %rod'ce so&e;"at different %atterns of
res'lts.

Why do temporal "radients occur(
One vie; of t"e te&%oral )radient in $orsa5off %atients ;as

t"at it reflected bot" .i0 t"e
conse<'ences of da&a)e at t"e

ti&e of t"e =ernic5e e%isode and .ii0 a progressie
deterioration

in t"e antero)rade ac<'isition of &e&ories d'rin) t"e years

of "eavy
drin5in) .9lbert et al., 1939 , 19:10. -"ere are t"ree

%ieces of evidence a)ainst t"e latter
idea1 .a0 B'tters and

(er&a5 .19:L 0 obtained a te&%oral )radient in an e&inent
scientist,

J, ;"o develo%ed t"e alco"olic $orsa5off syndro&e s"ortly after

;ritin) "is
a'tobio)ra%"y. 4e ;as tested on infor&ation derived

fro& t"is boo5 t"at "e "ad
a%%arently been a;are of at t"e ti&e

of ;ritin). Nevert"eless, "e s"o;ed a stri5in)
te&%oral )radient

in recallin) t"is infor&ation after "is =ernic5e e%isode. .b0

-"ere is no
si)nificant correlation bet;een t"e severityPextensiveness

of t"e 79 loss in $orsa5off
%atients and t"e esti&ated d'ration

of t"eir "eavy drin5in) .$o%el&an et al., 1999 0. .c0
-e&%oral

)radients "ave also been described in non-$orsa5off a&nesic

%atients of ac'te
onset .$o%el&an et al., 19:9 A ,<'ire et al.,

19:9 0. -o t"e extent t"at $orsa5off %atients
"ave a stee%er

)radient t"an ot"er a&nesic %atients .$o%el&an et al., 1999 0,

t"is &ay
res'lt fro& an additional, %ro)ressive antero)rade

i&%air&ent arisin) fro& t"eir %eriod of
"eavy drin5in)A "o;ever,

t"is is li5ely to be a relatively &inor factor, addin) to a

te&%oral )radient ;"ic" a%%ears ac'tely for ot"er reasons .$o%el&an,

19:9 A ar5in et
al., 1990 b0.

9 second vie; is t"at t"e &edial te&%oral and dience%"alic str'ct'res

are critical not only
in initial &e&ory for&ation, b't also

in t"e %"ysiolo)ical ?consolidation+ of &e&ories

over 2 or 8 years or lon)er, after ;"ic" &e&ories are stored

in neocortex and are no
lon)er de%endent '%on t"e &edial te&%oralPdience%"alic

syste& .?str'ct'ral reallocation+A
e.). ,<'ire et

al., 19:K A ,<'ire and 9lvare@, 1995 A -en) and ,<'ire, 1999 0.

-"is is
consistent ;it" t"e findin) t"at %atients ;it" &edial

te&%oral lobe or dience%"alic lesions
often s"o; only a brief

or even absent 79 .Jola-!or)an et al., 19:L A #'soir et al.,

1990
A *raff-7adford et al., 1990 A 7eed and ,<'ire, 199: A $o%el&an

et al., 1999 0. Fn
contrast, %atients ;it" an 79 extendin) bac5

&any years tend to "ave eit"er extensive
%at"olo)y beyond t"e

&edial te&%oral lobesPdience%"alon or an alco"olic "istory.

="ere
t"at %at"olo)y is %artic'larly ;ides%read, as in 9l@"ei&er

de&entia or "er%es
ence%"alitis, t"e 79 tends to extend a lon)

;ay bac5 ;it" a relatively ?flat+ te&%oral
)radient.

(onnectionist net;or5s "ave been 'sed to &odel t"e te&%oral

)radients
associated ;it" %rolon)ed consolidation .9lvare@ and

,<'ire, 1995 A ,<'ire and 9lvare@,
1995 A !c(lelland et al., 1995 A

!'rre, 1993 0. 9 f'nda&ental %roble& for t"is t"eory
is t"at

an extensive te&%oral )radient, )oin) bac5 20 to 80 years, ;o'ld

i&%ly t"at
%"ysiolo)ical consolidation &'st contin'e for a very

lon) ti&e indeed .Nadel and
!oscovitc", 1993 0.

9 t"ird vie; is t"at, as e%isodic &e&ories are re"earsed, t"ey

ado%t a &ore semantic
for&, losin) t"eir context'al i&&ediacy

or vividness, b't protecting t"e& fro& t"e effects
of brain

da&a)e .(er&a5, 19:K 0. 2arly &e&ories are %artic'larly salient

and ;ell
re"earsed .(er&a5, 19:K A =eis5rant@, 19:5 A ,a)ar et

al., 19:: A $o%el&an, 19:9 0,
and t"erefore %rotected fro& any

retrieval deficit. -"ere are reasons for s'%%osin) t"at t"e
79 res'lts, at least in %art, fro& a deficit in retrieval %rocesses,

rat"er t"an fro& a
destr'ction of &e&ory stora)e per se. .a0

9&nesic and de&entin) %atients can &anifest a
dis%ro%ortionate

benefit fro& reco)nition or context'al c'es .co&%ared ;it" "ealt"y

controls0 des%ite very %oor recall %erfor&ance .$o%el&an, 19:9 A

ar5in et al., 1990 aA
Verfaellie et al., 1995 A $o%el&an et al.,

1999 0. .b0 9 deficit in retrievin) re&ote
&e&ories s'%eri&%osed

'%on an antero)rade learnin) deficit is consistent ;it" t"e

observation t"at t"ere are )enerally lo; and non-si)nificant

correlations bet;een t"e
severity of 99 and 79 .,"i&a&'ra and

,<'ire, 19:L A $o%el&an, 19:9 A ar5in, 1991
0. .c0 ,i)nificant

correlations bet;een re&ote &e&ory scores and frontalPexec'tive

%erfor&ance .$o%el&an, 1991 A Verfaellie et al., 1995 A #+2s%osito

et al., 199L 0
s'))est t"e %'tative role of frontalPneocortical

&ec"anis&s in or)ani@in) t"e retrieval of
re&ote &e&ories. -o

t"e extent t"at early &e&ories are ?se&antici@ed+

or "i)"ly
re"earsed, t"ey are %rotected fro& s'c" a retrieval

deficit, and a te&%oral )radient res'lts
.(er&a5, 19:K 0. 4o;ever,

a &aHor %roble& for t"is t"eory is t"e findin) t"at &e&ories

;"ic" are se&antic virt'ally fro& t"e o'tset, s'c" as 5no;led)e

of t"e &eanin) of ne;
;ords, can also s"o; a te&%oral )radient

.Verfaellie et al., 1995 a, b0.

9 fo'rt" vie; %ost'lated by Nadel and !oscovitc" is t"at t"e

"i%%oca&%i are
contin'o'sly involved in t"e stora)e and retrieval

.reactivation0 of &e&ory traces .Nadel
and !oscovitc", 1993 A

!oscovitc" and Nadel, 199: , 19990. Fn t"eir &odel, a
"i%%oca&%alMneocortical

ense&ble constit'tes t"e &e&ory trace for an e%isode, ;it"in

;"ic" a distrib'ted )ro'% of "i%%oca&%al ne'rones acts as an

index or %ointer to
neocortical .or ot"er0 ne'rones t"at re%resent

t"e infor&ation, bindin) t"e activations of
t"ese ne'rones into

a co"erent &e&ory trace .co&%are !orton et al., 19:5 A !'rre,

1993
0. Beca'se t"e "i%%oca&%i are al;ays involved in encodin)

%rocesses ;"en attention is
actively en)a)ed, Nadel and !oscovitc"

.1993 0 %ro%osed t"at t"e reactivation of a
&e&ory trace res'lts

in a ne;ly encoded trace1 re%eated reactivations ;ill %rod'ce

?multiple traces+ and t"e ?extraction+

of fact'al infor&ation fro& an e%isode and its
inte)ration into

%re-existin) .neocortical0 se&antic &e&ory stores. 7etrieval

beco&es
easier as t"e n'&ber of traces %roliferates ;it" re"earsal,

res'ltin) in a te&%oral )radient.
-"e a't"ors %redicted t"at

t"e extent of 79 and slo%e of t"e te&%oral )radient for
e%isodic

&e&ories ;o'ld de%end '%on t"e si@e of a "i%%oca&%al lesion,

and t"at
co&%lete "i%%oca&%al transection ;o'ld res'lt in a

flat )radient. 4o;ever, fact'al
infor&ation .e.). for %'blic

events and %ersonalities0 is event'ally stored in neocortex

inde%endently of t"e ."i%%oca&%al-&ediated0 e%isode in ;"ic"

it ;as ac<'ired1 t"is
as%ect of t"e t"eory rese&bles ?str'ct'ral

reallocation+ a%%roac"es and %redicts a stee%er
)radient

for ?se&antic+ t"an for a'tobio)ra%"ical re&ote

&e&ories. .Fn t"e &ost recent
acco'nts of t"is t"eory, t"e role

%revio'sly attrib'ted to t"e "i%%oca&%i is no; accorded
to t"e

&edial te&%oral lobesA see e.). C'Hii et al., 2000 .0

Nadel and !oscovitc" .1993 0 cited a lar)e n'&ber of st'dies,

in ;"ic" &edial te&%oral
lobe %at"olo)y ;as acco&%anied by a

te&%orally extensive 79 .see also C'Hii et al., 2000
0, and Vis5ontas

et al. .2000 0 fo'nd a flat )radient in &e&ory for a'tobio)ra%"ical

incidents in te&%oral lobe e%ile%sy %atients, ;"ereas t"ere

;as a stee%er )radient in t"e
recall of %ersonal se&antic facts.

,i&ilarly, (i%olotti et al. .2001 0 re%orted an extensive
79

in a %atient ;it" severe "i%%oca&%al atro%"y .alt"o')" t"ere

;as so&e evidence of a
)entle )radient in t"eir data0. On t"e

ot"er "and, virt'ally all of t"e %atients cited by
Nadel and

!oscovitc" .1993 0 "ad extensive %at"olo)y else;"ere. Ot"er re%orts

s'))est
only a brief 79 in %atients ;it" &edial te&%oral .or

dience%"alic0 lesions .Jola-!or)an
et al., 19:L A 7eed and ,<'ire,

199: 0A and $o%el&an and collea)'es did not find any
correlation

bet;een !7F &eas'res of &edial te&%oral vol'&e and t"e extent

or )radient
of 79 in "er%es and "y%oxic %atients .!. #. $o%el&an,

#. /asserson, #. $in)sley, C.
Bello, (. 7's", N. ,tan"o%e et

al., &an'scri%t s'b&itted0, contrary to Nadel and
!oscovitc"+s

%rediction.

$e#ersed temporal "radients in semantic dementia
9s &entioned briefly above, ,no;den et al. .199L 0 fo'nd recency

effects .reversed
te&%oral )radients0 in se&antic de&entia %atients+

&e&ory for %ersonal facts and
incidents. -"ey contrasted t"is

)radient ;it" t"at obtained &ore ty%ically in 9l@"ei&er or
a&nesic

%atients. *ra"a& and 4od)es .1993 0 re%licated t"is findin) on

a'tobio)ra%"ical
&e&ory tests and 4od)es and *ra"a& .199: 0 on

a fa&o's na&es test, alt"o')" t"e
latter findin) ;as so&e;"at

e<'ivocal beca'se of ?ceilin)+ and ?floor+

effects and t"e
fail're to re%ort )ro'% by ti&e-%eriod interactions.

One inter%retation of t"ese findin)s is
in ter&s of str'ct'ral

reallocation1 in se&antic de&entia, t"e te&%oral neocortex is

severely da&a)ed and early &e&ories are lost, ;"ereas &ore recent

&e&ories are
de%endent '%on &edial te&%oral str'ct'res, ;"ic"

are relatively %reserved .*ra"a& and
4od)es, 1993 0. 9n alternative

ex%lanation is t"at t"e ?a'tobio)ra%"ical+ i&%air&ent

in
t"ese %atients &ainly reflects t"eir lin)'istic and se&antic

deficit, associated ;it" left
te&%oral da&a)e, and t"at c'rrent

everyday ex%erience %rovides c'es for t"e retrieval of
recent

a'tobio)ra%"ical &e&ories stored else;"ere .!oscovitc" and Nadel,

1999 A
$o%el&an, 2000 0, H'st as it "el%s t"e& to &a5e conversational

'se of ;ords t"ey cannot
define .,no;den et al., 199L , 19990.

(onsistent ;it" t"is, !oss and collea)'es .!oss et
al., 2002 0

"ave bot", fo'nd a %rono'nced beneficial effect of c'ein) in

a'tobio)ra%"ical
&e&ory retrieval in se&antic de&entia %atients,

and t"is c'ein) effect ;as only very
;ea5ly related to ti&e

%eriod.

Dissociable forms of retro"rade amnesia
On t"e basis of an extensive revie; of t"e literat're, $a%'r

.1999 0 %ro%osed a
?%rovisional+ fra&e;or5, delineatin)

&aHor dissociable for&s of ne'rolo)ical retro)rade
a&nesia.

4e advocated a distinction bet;een e%isodic 79 and se&antic

79, and
s'bdivisions ;it"in t"ese &aHor )ro'%in)s. Fn e%isodic

79, "e distin)'is"ed bet;een
?%re-ictal+ and ?extended+

79, and in se&antic 79 bet;een 5no;led)e of %eo%le and
events.

=it"in ?%eo%le+, $a%'r ar)'ed t"at 5no;led)e of

na&es and faces can be
differentially affected and, ;it"in ?faces+,

"e differentiated 5no;led)e of fa&o's vers's
%ersonally fa&iliar

faces.

9 %roble& ;it" s'c" dissociations, as ;ill be disc'ssed belo;,

is t"at t"ey are lar)ely
based '%on sin)le cases and, ;"en lar)er

)ro'%s of %atients are investi)ated, considerable
overla% in

%erfor&ance can be fo'nd, and correlates of %erfor&ance &ay

e&er)e ;"ic"
&i)"t acco'nt for a%%arent dissociations .$o%el&an,

2000 a0. Cactors s'c" as levels of
ed'cation, &edia ex%os're

and intelli)ence can stron)ly affect %erfor&ance on se&antic

tests, and exec'tive dysf'nction &ay adversely affect %erfor&ance

on e%isodic tests1
sin)le case st'dies do not al;ays exa&ine

t"e infl'ence of s'c" factors. !oreover,
co)nitiveMbrain

relations"i%s are so&eti&es %ost'lated on t"e basis of %atients

;it" eit"er
very s'btle &ar5ers of brain dysf'nction .e.). on

22* or ,2(-0 or %atients ;it"
&'lti%le sites of %at"olo)y,

;"ic" &eans t"at t"e attrib'tion of ca'sality to a %artic'lar

site of ?%at"olo)y+ &ay be e<'ivocal even in t"e

%resence of an a%%arent ?do'ble
dissociation+ .$o%el&an,

2000 a0. -"e sa&e iss'e arises as in (ara&a@@a+s disc'ssion

of
cate)ory-s%ecific deficits .(ara&a@@a, 199: 01 does a differential

%attern of %erfor&ance
on co)nitive tas5s, even if s'%%orted

by a do'ble dissociation, necessarily i&%ly t"at
s%ecific brain

&ec"anis&s s"o'ld be %ost'lated as 'nderlyin) t"e cate)ories

affected or
s%ared>

#es%ite t"ese <'alifications, $a%'r+s .$a%'r, 1999 0 cate)ori@ation

does %rovide a 'sef'l
fra&e;or5 for vie;in) t"e %attern of %erfor&ance

of different %atients on %artic'lar types
of test. !oreover,

so&e )eneral observations abo't locali@ed effects can be &ade.

Q'antitative !7F evidence s'))ests t"at ;ides%read net;or5s

;it"in t"e te&%oral lobe,
frontal neocortex and t"ala&'s are

involved in t"e stora)e and retrieval of re&ote
&e&ories .!.

#. $o%el&an, #. /asserson, #. $in)sley, C. Bello, (. 7's", N.

,tan"o%e et
al., s'b&itted0, b't t"ere are al&ost certainly

focal or locali@ed contrib'tions ;it"in t"ese
re)ions. Ft see&s

li5ely t"at t"e frontal lobes contrib'te %artic'larly to t"e

%lannin),
&onitorin) and or)ani@ation of retrieval %rocesses

in re&ote &e&ory, %artic'larly ;"ere
&ore ?effortf'l+

or active reconstr'ction is re<'ired .!an)els et al., 199L A

$o%el&an et
al., 1999 0. -"e ri)"t te&%oral lobe a%%ears to be

%artic'larly i&%ortant for t"e
reconstr'ctive %rocesses en)a)in)

vis'al or &'lti-&odal i&a)ery necessary for retrievin)
a'tobio)ra%"ical

or ?event+ &e&ories .O+(onnor et al., 1992 A

O)den, 1998 A 7'bin and
*reenber), 199: A $o%el&an et al., 1999 A

b't contrast $itc"ener et al., 1999 0. On t"e
ot"er "and, t"e

left te&%oral lobe &ay be %artic'larly cr'cial for t"e stora)e

and access of
re&ote se&antic and lexical-se&antic infor&ation

.#e 7en@i et al., 19:3 A $o%el&an et
al., 1999 0.

.euroima"in" of remote memory
-o date, t"ere "ave been relatively fe; activation st'dies of

re&ote &e&ory retrieval.
Cin5 et al. .199L 0 s"o;ed ;ides%read

activation of ri)"t frontal and te&%oral lobe .and
left &edial

te&%oral0 re)ions ;"en "ealt"y s'bHects i&a)ined incidents fro&

t"eir re&ote
%ast, relative to t"eir brain state ;"en i&a)inin)

events 'nrelated to t"eir %ast. !a)'ire
and !'&&ery .1999 0 re<'ired

s'bHects to listen to state&ents t"at differed accordin) to

;"et"er or not t"ey constit'ted a'tobio)ra%"ical or )eneral

5no;led)e, and ;"et"er or
not t"ey "ad a s%ecific foc's in ti&e.

-"ere ;as en"anced activity for ti&e-s%ecific
a'tobio)ra%"ical

events in t"e left "i%%oca&%'s, &edial %refrontal cortex and

left
te&%oral %ole. Bilateral te&%oro-%arietal re)ions ;ere

activated %referentially for
%ersonal &e&ories, re)ardless of

ti&e-s%ecificity. (on;ay et al. .1999 0 obtained left
frontal,

inferior te&%oral and occi%ito-%arietal activations in verbal

c'ein) of
a'tobio)ra%"ical &e&ories and, li5e 7yan et al. .2001 0

and !a)'ire et al. .2001 0, did
not obtain any &edial te&%oral

differences in retrievin) re&ote or recent &e&ories. Fn
contrast,

4aist et al. .2001 0 re%orted te&%orally )raded &edial te&%oral

activations in
retrievin) t"e na&es of fa&o's faces fro& different

decades.

Summary
-e&%oral )radients of varyin) stee%ness are ;idely re%orted

in 79, b't t"eir 'nderlyin)
basis is dis%'ted. 9 %ro)ressive

deterioration in ac<'irin) ne; &e&ories cannot acco'nt
for te&%oral

)radients in %atients ;it" an ac'te onset of a&nesia. (onsolidation

t"eory
re<'ires %"ysiolo)ical c"an)es lastin) years or decades.

-"e se&antic %rotection
"y%ot"esis is 'nder&ined by te&%oral

)radients in ?%'rely+ se&antic &e&ories. -"e
&'lti%le

trace "y%ot"esis "as %roble&s ;it" cases of brief 79 in %atients

;it" isolated
&edial te&%oral or dience%"alic %at"olo)y. -"e

%recise role of s%ecific brain str'ct'res in
storin)Pretrievin)

re&ote &e&ories is c'rrently an active to%ic of researc"A and

t"e
ne'robiolo)ical stat's of t"e &any dissociations %ost'lated

in 79 re&ains 'nclear.

!an retro"rade amnesia e#er
be 0isolated1(

!ase reports in the literature
Fn t"e early 19:0s, a series of %atients ;it"
a%%arently isolated

79 ;ere described .7o&an-
(a&%os et al., 19:0 A *oldber) et al.,

19:1 , 19:2 A
9ndre;s et al., 19:2 A 7o'ssea'x et al., 19:K 0. Of

Top
Summary
Introduction
Varieties of amnesic syndrome
What is impaired and...
Explicit and implicit memory
Do temporal lobe, diencephalic...
ow independently semantic is...
What determines the pattern...
!an retro"rade amnesia e#er...
Does psycho"enic amnesia in#ol#e...
ow and when do...
!onclusions
$eferences
t"ese, t"e &ost convincin) st'dy ;as by *oldber) et al. .19:1 0,

;"o described a 8L-
year-old %atient ;it" an o%en s5'll fract're

associated ;it" extensive ri)"t te&%oral lobe
c"an)es on (-

scan and a lesser de)ree of left te&%oral abnor&ality. -"e %atient

"ad an
initially ?%rofo'nd+ 99, b't i&%roved to

a state in ;"ic" a severe i&%air&ent in re&ote
&e&ory ;as acco&%anied

by only &ini&al antero)rade deficits.

$a%'r et al. .1992 0 described a 2L-year-old ;o&an ;"o "ad fallen

fro& a "orse. ,"e
"ad s'stained left and ri)"t frontal cont'sions,

evident on (- scan ;it" associated re)ions
of lo; atten'ation.

9n !7F scan 1: &ont"s later s"o;ed lesions in t"e left and ri)"t

te&%oral %oles, ;it" t"e "i%%oca&%i a%%arently intact. On co)nitive

testin), t"e %atient
"ad an esti&ated %re-&orbid FQ .N97-0 of

105 ;it" a c'rrent .=9F,-70 FQ of 9:.
CrontalPexec'tive test

%erfor&ance ;as intact, b't t"e %atient ;as i&%aired across

all t"e
re&ote &e&ory tests e&%loyed. !ost antero)rade &e&ory

tas5s ;ere %erfor&ed
nor&ally, b't it s"o'ld be noted t"at t"e

%atient %erfor&ed %oorly at i&&ediate lo)ical
&e&ory .18t" %ercentile0

and on t"e faces co&%onent of t"e reco)nition &e&ory test .Kt"
%ercentileA =arrin)ton, 19:K 0.

,o&e;"at si&ilarly, /evine et al. .199: 0 described a %atient,

!/, ;"o "ad s'ffered a
road traffic accident res'ltin) in a

ri)"t frontal lesion, involvin) t"e 'ncinate fascic'l's,
as

;ell as left %refrontal "ae&orr"a)es. -"is %atient "ad an initially

severe 99, consistent
;it" a re%orted %ost-tra'&atic a&nesia

of at least 8K days .and co&a for L days0, b't t"is
resolved

leavin) a verbal &e&ory index .=!,-70 of 12: and a severe 79.

4o;ever,
findin)s on a tas5 re<'irin) t"e s'bHect to &a5e re&e&berP5no;

H'd)e&ents in
antero)rade &e&ory over varyin) delays, as ;ell

as in a 2- activation st'dy, indicated
t"at t"ere ;ere, in

fact, s'btle i&%air&ents of antero)rade &e&ory.

9 f'rt"er st'dy, co&&only cited in t"is literat're, is t"at

by O+(onnor et al. .1992 0.
-"ese a't"ors described a %atient

;"o, follo;in) extensive ri)"t te&%oral lobe da&a)e
fro& "er%es

ence%"alitis, &anifested a severe loss of a'tobio)ra%"ical &e&ories,

relative
to 5no;led)e of %'blic infor&ation. -"e %atient also

s'ffered a vis'al a)nosia, and t"e
a't"ors s'))ested t"at t"e

loss of a'tobio)ra%"ical &e&ories &i)"t be related to a
%roble&

in conH'rin) '% vis'al i&a)ery. -"e a't"ors ;rote1 ?;"ilst

s"e "as only a &ild to
&oderate erbal D&y e&%"asisE learnin)

deficits, s"e contin'es to de&onstrate very
severe retro)rade

a&nesia.+ Fndeed, t"is %atient "ad a severe deficit in

vis'o-s%atial
antero)rade &e&ory as ;ell as "er 79.

-"ere are &any ot"er cases cited in t"is literat're, ;"ic" "as

been revie;ed extensively
else;"ere .$o%el&an, 2000 a0. ,o&e

of t"ese %atients "ad definite str'ct'ral brain
da&a)e .e.).

!aravita et al., 1995 A 2vans et al., 199L A !attioli et al.,

199L A (arlesi&o
et al., 199: 0. Fn ot"er cases, t"e evidence

for brain %at"olo)y ;as e<'ivocal .7o&an-
(a&%os et al., 19:0 A

,t'ss and *'@&an, 19:: A ,tar5stein et al., 1993 A Venneri and

(affarra, 199: 0, or ne)li)ible .e.). ,tracciari et al., 199K A

#e 7en@i et al., 1995 A
/'cc"elli et al., 1995 A a%a)no, 199: 0.

#e 7en@i et al. .1993 0 ar)'ed t"at s'c" cases
s"o'ld be called

?f'nctional a&nesia+ on t"e )ro'nds t"at or)anic

c"an)e &ay event'ally
be discovered ?at t"e cell'lar level+

.see also /'cc"elli et al., 1995 , 199:A $a%'r, 199L A
,%innler

et al., 199L 0.

!riti2ue of the literature
Fn a recent revie; of t"is to%ic .$o%el&an, 2000 aA see also

$a%'r, 2000 A $o%el&an,
2000 b0, t"e %resent a't"or concl'ded

s'c" cases can be classified 'nder a n'&ber of
different cate)ories,

as follo;s.

.i0 ,o&e cases "ad severe a'tobio)ra%"ical a&nesia and "ave

been ;idely cited in t"is
literat're, des%ite t"e fact t"at

t"ere ;as evidence of very %oor antero)rade &e&ory,
so&eti&es

%artic'larly for vis'o-s%atial &aterial .e.). O+(onnor

et al., 1992 A
!ar5o;itsc" et al., 1998 A O)den, 1998 A Bro;n and

("obor, 1995 0. ,'c" cases cannot
be described as instances of

?isolated+, ?focal+ or ?dis%ro%ortionate+

79, des%ite t"e fact
t"at t"e relations"i% bet;een &e&ories

for vis'o-s%atial &aterial and a'tobio)ra%"ical
a&nesia %oses

interestin) t"eoretical <'estions.

.ii0 -"ere are ot"er cases in t"e literat're ;"o s"o;ed %oor

antero)rade &e&ory in
&oderate or &ore s'btle for& across a

n'&ber of antero)rade tests, %artic'larly lo)ical
&e&ory, face

reco)nition &e&ory and tests of delayed recall .$a%'r et al.,

19:L , 19:9,
1992, 199LbA ,t'ss and *'@&an, 19:: A 4'n5in et al.,

1995 A !aravita et al., 1995 A
(alabrese et al., 199L A !attioli

et al., 199L A (arlesi&o et al., 199: A *ainotti et al.,
199: A

/'cc"elli and ,%innler, 199: A Je&an et al., 199: 0. Fn t"ese

instances, t"e %"rase
?dis%ro%ortionate+, rat"er

t"an ?focal+ or ?isolated+, 79 see&s

&ore a%%ro%riate.
!oreover, t"ese cases %ose t"e <'estions of

;"y t"ere are s'c" c"aracteristic %atterns,
and ;"et"er ?li5e+

"as really been co&%ared ;it" ?li5e+ across retro)rade

and antero)rade
&e&ory> ossibly t"e 79 tas5s re<'ire )reater

'se of vis'al or &'lti-&odal i&a)ery,
;"ic" &ay be critical

for a'tobio)ra%"ical incident recall .O+(onnor et al.,

1992 A
O)den, 1998 A Bro;n and ("obor, 1995 A Van der /inden et

al., 199L A *ainotti et al.,
199: A 7'bin and *reenber), 199: 0.

9lternatively, re&ote &e&ory tas5s &ay be &ore
?effortf'l+,

re<'irin) &ore s%ontaneo's strate)ies and ?exec'tive+

%lannin) and
or)ani@ation in retrieval .Baddeley and =ilson,

19:L A #ella ,ala et al., 1998 0 t"an do
antero)rade tas5s. 9

t"ird %ossibility is t"at t"e re&ote &e&ory deficit reflects

i&%aired
consolidation, &anifest on recall tests at %rolon)ed

delays .#e 7en@i and /'cc"elli, 1998
A !aravita et al., 1995 A

2vans et al., 199L A $a%'r et al., 199Lb A !attioli et al., 199L
0.

-"e latter is an attractive "y%ot"esis, alt"o')" ins'fficiently

tested as yet, b't it i&%lies
t"at t"e %roble& is not s%ecific

to retro)rade &e&ory. Only one recent st'dy .!annin),
2002 0

"as serio'sly addressed t"e iss'e of co&%arin) ?li5e+

;it" ?li5e+, b't t"e %atient
%erfor&ed nor&ally

or near nor&ally at standardi@ed re&ote &e&ory tas5s .9!F, #ead-
or-9live0.

.iii0 ,o&e of t"e cases in t"e literat're s"o;ed a s%ecific

i&%air&ent in a'tobio)ra%"ical
&e&ory .2vans et al., 199L 0,

ot"ers s"o;ed %oor recall for %'blic or se&antic
5no;led)e .e.).

$a%'r et al., 19:L , 19:9A $a%'r, 199K 0, ;"ilst yet ot"ers s"o;ed

79
across a ;ide ran)e of tas5s .e.). $a%'r et al., 1992 0. 9ll

t"ese cases tend to )et cited as
instances of ?focal 79+,

b't it is i&%ortant to e&%"asi@e t"at t"ey are not t"e sa&e.

Ff
dissociations are to be %ost'lated, it o')"t to be s"o;n

t"at t"e %atient fails on &ore t"an
one tas5 reflectin) a %'r%orted

deficit, ot"er;ise t"e findin)s &ay si&%ly reflect c"ance
variation

in test %erfor&ance. ,econdly, so&e %atterns of findin)s can

be ex%lained in
ot"er ter&s1 for exa&%le, %oor recall for %'blic

or se&antic 5no;led)e &ay reflect lo;
intelli)ence or %oor ed'cation

and &edia ex%os're .$o%el&an, 19:9 A $a%'r et al., 1999
0, ;"ereas

%rofo'ndly i&%aired a'tobio)ra%"ical &e&ory retrieval in t"e

%resence of
s'r%risin)ly )ood se&antic 5no;led)e &i)"t reflect

severe frontalPexec'tive i&%air&ent
.$o%el&an, 2000 a0.

.iv0 -"e &ost convincin) cases in t"is literat're "ave been

t"ose in ;"o& t"ere ;as an
initially severe antero)rade a&nesia,

as ;ell as an extensive 79, %artic'larly follo;in)
"ead inH'ry.

-"e 79 re&ained %rofo'nd, ;"ereas t"e 99 beca&e only &oderate,

&ild or
&ini&al .7o'ssea'x et al., 19:K A *oldber) et al., 19:1 A

4'n5in et al., 1995 A
(alabrese et al., 199L A 2vans et al., 199L A

$a%'r et al., 199Lb A !attioli et al., 199L A
(arlesi&o et al.,

199: A /evine et al., 199: A !ar5o;itsc" et al., 1999 0. Fn s'c"

cases,
t"e iss'e is not so &'c" ;"et"er 79 can occ'r ;it"o't

99, b't what determines
differential patterns of recoery. -"is

is a so&e;"at different <'estion, b't one on ;"ic"
;e "ave s'r%risin)ly

little infor&ation. Ft is li5ely t"at both %"ysiolo)ical .s'c"

as ne'ral
reor)ani@ation0 and %syc"olo)ical factors infl'ence

recovery %rocesses, %artic'larly
follo;in) "ead inH'ry ./is"&an,

1938 , 19::A Ne;co&be, 19:2 0. -"e fact t"at recovery
fro& 99

and 79 can occ'r at differential rates does not necessarily

&ean t"at t"ey are
co&%letely dissociated.

.v0 9 second )ro'% of %atients ;ere t"ose ;it" -29, ;"o co&&only

re%ort ?)a%s+ in t"eir
a'tobio)ra%"ical &e&ory and

"ave often been cited in t"is literat're. ,o&e of t"ese cases

"ad eit"er &oderately severe .$o%el&an et al., 199K a0 or &ore

s'btle .$a%'r et al.,
19:9 0 antero)rade &e&ory i&%air&ent, and

t"ey often %erfor&ed ;ell on standardi@ed
tests of a'tobio)ra%"ical

and re&ote &e&ory .$a%'r et al., 19:L A $a%'r, 1998 A
$o%el&an

et al., 199K aA Je&an et al., 199: A !anes et al., 2001 0. Fn s'c"

cases, t"e
&ost %arsi&onio's ex%lanation is t"at t"e %atients

"ad brief r'ns of sei@'re activity in t"e
%ast, ;"ic" ;ere not

detected clinically, and t"at t"ese res'lted in fa'lty .antero)rade0

encodin) of very s%ecific ite&s in a'tobio)ra%"ical &e&ory .$o%el&an,

2000 aA b't see
$a%'r, 2000 0.

.vi0 9 final )ro'% of cases ;ere t"ose in ;"o& %syc"o)enic factors

&i)"t ;ell "ave been
%redo&inant. Brain disease and %syc"iatric

%roble&s co&&only co-occ'r .$o%el&an and
(ra;ford, 199L A !ar5o;itsc",

199L 0, and %syc"olo)ical factors &ay exacerbate
co)nitive deficits

andPor infl'ence %atterns of recovery. -"is &ay "ave been %artic'larly

tr'e of %atients in ;"o& &ild conc'ssion ;as acco&%anied by

dis%ro%ortionate
i&%air&ents in retro)rade &e&ory. Fn so&e case

descri%tions of focal 79, evidence of
%ossible %syc"iatric factors

;as only briefly or scantily disc'ssed .7o&an-(a&%os et al.,

19:0 A 9ndre;s et al., 19:2 A ,t'ss and *'@&an, 19:: A ,tracciari

et al., 199K A #e
7en@i et al., 1995 A !attioli et al., 199L 0,

alt"o')" a f'ller disc'ssion "as been incl'ded
in &ore recent

st'dies .Barbarotto et al., 199L A /'cc"elli et al., 199: A a%a)no,

199: A
!ac5en@ie 7oss, 2000 0. (ases in ;"o& 79 "as been reversed

d'rin) an intervie; 'nder
sodi'& a&ytal .,t'ss and *'@&an, 19:: A

$o%el&an, 2000 a0 add ;ei)"t to t"e
%ossibility t"at %syc"olo)ical

factors can be cr'cial, even in t"e %resence of so&e
evidence

of or)anic brain da&a)e.

Summary
#is%ro%ortionate 79 does occ'r, alt"o')" ?focal+

or ?isolated+ 79 is &'c" &ore
e<'ivocal. Ft is an

exa&%le of a %ost'lated dissociation ;"ose ne'robiolo)ical stat's

re&ains 'nclear. -"e evidence t"at it is related to a s%ecific,

locali@ed brain %at"olo)y is
t"in, ;"en acco'nt is ta5en of

t"e above factors. C't're researc" needs to concentrate not

so &'c" on t"e existence of ?dissociation+, b't

on t"e deter&inants of differential
recovery %rocesses in "ead

tra'&a, t"e nat're of &e&ory ?)a%s+ in -29, and

on t"e
interaction of %syc"o)enic factors and brain %at"olo)y

in %rod'cin) 79.

Does psycho"enic amnesia in#ol#e the same
mechanisms as or"anic amnesia(

syc"olo)ical for&s of &e&ory loss can eit"er be ?)lobal+

or ?sit'ation s%ecific+. -"e
for&er refers to a

%rofo'nd loss of %ast &e&ories, often enco&%assin) t"e sense

of
%ersonal identity, and t"e latter to &e&ory loss for a %artic'lar

incident or %art of an
incident.

3lobal amnesia
-"is is exe&%lified by t"e so-called ?f')'e state+

.4'nter, 19LK 0, so&eti&es 5no;n as
?f'nctional retro)rade

a&nesia+ .,c"acter et al., 19:2 0. -"e f')'e state refers,

in essence,
to a syndro&e consistin) of a s'dden loss of all

a'tobio)ra%"ical &e&ories and
5no;led)e of self or %ersonal

identity, 's'ally associated ;it" a %eriod of ;anderin), for

;"ic" t"ere is a s'bse<'ent a&nesic )a% '%on recovery. C')'e

states 's'ally last a fe;
"o'rs or days only1 if %rolon)ed,

t"e s's%icion of si&'lation &'st al;ays arise.

-"ere a%%ear to be t"ree &ain %redis%osin) factors for f')'e

e%isodes. Cirst, f')'e states
are al;ays %receded by a severe,

%reci%itatin) stress, s'c" as &arital or e&otional discord
.$an@er,

1989 0, bereave&ent .,c"acter et al., 19:2 0, financial %roble&s

.$an@er, 1989
0, a c"ar)e of offendin) .=ilson et al., 1950 0

or stress d'rin) ;arti&e .,ar)ant and
,later, 19K1 A arfitt

and *all, 19KK A 4'nter, 19LK 0. ,econdly, de%ressed &ood is an

extre&ely co&&on antecedent for %syc"o)enic f')'e states. Berrin)ton

et al. .195L0
;rote1 ?Fn nearly all f')'es, t"ere a%%ears

to be one co&&on factor, na&ely a de%ressive
&ood. ="et"er t"e

individ'al in t"e f')'e is %syc"otic, ne'rotic, or %syc"o%at"ic,

de%ression see&s to start off t"e f')'e.+ Cor exa&%le,

9beles and ,c"ilder .19850
described a ;o&an ;"o deserted "er

"'sband for anot"er &an1 after a ;ee5, s"e
deter&ined to ret'rn

to "er fa&ily b't, as s"e descended into t"e 'nder)ro'nd rail;ay

station, s"e ;as conte&%latin) s'icide. -"e a't"ors tersely

re%orted t"at ?instead a&nesia
develo%ed+. -"e t"ird

factor is a "istory of a transient, or)anic a&nesia fro& s'c"

ca'ses
as e%ile%sy .,ten)el, 19K1 0, "ead inH'ry .Berrin)ton

et al., 195L 0 or alco"olic blac5o'ts
.*ood;in et al., 19L9 0.

Cor exa&%le, Berrin)ton et al. .195L 0 re%orted t"at 19 of t"eir

83 f')'e cases "ad %revio'sly ex%erienced a "ead inH'ry of so&e

de)ree of severity. Ft
a%%ears t"at %atients ;"o "ave %revio'sly

ex%erienced a transient or)anic a&nesia, and
t"en beco&e de%ressed

andPor s'icidal, are %artic'larly li5ely to )o into a f')'e

in t"e
face of a severe %reci%itatin) stress.

-"e clinical and ne'ro%syc"olo)ical %"eno&ena in s'c" cases

bear interestin)
rese&blances to or)anic a&nesia. Cor exa&%le,

t"ere &ay be islets or fra)&ents of
%reserved &e&ory ;it"in

t"e a&nesic )a%. 9 ;o&an ;"o ;as d'e to &eet "er "'sband to
disc'ss divorce recalled t"at s"e ;as ?s'%%osed to &eet

so&eone+ .$an@er, 1989 0. 9
yo'n) &an, ;"o sli%%ed into

a f')'e follo;in) "is )randfat"er+s f'neral, recalled

a
cl'ster of details fro& t"e year ;"ic" "e described .after

recovery0 as "avin) been t"e
"a%%iest of "is life .,c"acter

et al., 19:2 0. -"e s'bHect &ay ado%t a detac"ed attit'de to

t"ese &e&ory fra)&ents, describin) t"e& as ?stran)e and

'nfa&iliar+ .(oriat, 1903 A
!ar5o;itsc", 199L 0. Fn &any

cases, se&antic 5no;led)e re&ains 'ni&%aired .e.).
$an@er, 1989 A

,c"acter et al., 19:2 0, ;"ereas in ot"ers it is also i&%licated

.(oriat,
1903 A 9beles and ,c"ilder, 1985 A $an@er, 1989 0. ,i&ilarly,

%erfor&ance of verbal
learnin) tests can be 'naffected .9beles

and ,c"ilder, 1985 A $o%el&an et al., 199K b0,
&ildly i&%aired

.,c"acter et al., 19:2 0 or &ore severely i&%aired .*'dHonsson

and
-aylor, 19:5 0. !e&ory for %roced'ral s5ills is often %reserved

.e.). (oriat, 1903 A
Bradford and ,&it", 1939 0 and t"is &ay also

be tr'e of ot"er as%ects of i&%licit &e&ory
.(a&%odonico and

7ediess, 199L A $i"lstro& and ,c"acter, 2000 0. #eliberate c'ein)

of
&e&ories is seldo& s'ccessf'l .(oriat, 1903 A $an@er, 1989 A

$o%el&an et al., 199K b0,
b't &e&ory retrieval is often facilitated

by c"ance c'es in t"e environ&ent .e.). 9beles
and ,c"ilder,

1985 A ,c"acter et al., 19:2 0. $o%el&an .1995 b0 )ave t"e exa&%le

of a
%atient ;"o, on seein) an a't"or+s na&e on t"e s%ine

of a boo5, recalled t"at "e "ad a
friend of t"at na&e ;"o ;as

dyin) of cancer. On transfer to a %syc"iatric ;ard, "e
recollected

t"e details of anot"er %syc"iatric "os%ital ad&ission fro& years

earlier.

Situation/specific amnesia
,it'ation-s%ecific a&nesia can arise in a variety of circ'&stances,

incl'din) co&&ittin)
an offence, bein) t"e victi& of an offence

or of c"ild sex'al ab'se, and in a variety of
ot"er circ'&stances

res'ltin) in %ost-tra'&atic stress disorder .-,#0.

4ffences
Offenders as ;ell as victi&s of cri&es co&&only clai& a&nesia,

%artic'larly in violent
offences, and %syc"iatrists or ne'rolo)ists

are often called '%on to co&&ent '%on t"is.
9&nesia is clai&ed

by 25MK5N of offenders in cases of "o&icide, a%%roxi&ately

:N of
%er%etrators of ot"er violent cri&es and a s&all %ercenta)e

of non-violent offenders
.$o%el&an, 19:3 b, 1995b0. ,'c" findin)s

"ave been &ade in different settin)s,
incl'din) a re&and %rison

.-aylor and $o%el&an, 19:K 0, a forensic %syc"iatric
o't%atient

clinic .Bradford and ,&it", 1939 0 and a 4o&e Office 7e)ister

of /ife
Offenders .y@sora et al., 2002 0.

Ft is necessary to excl'de 'nderlyin) ne'rolo)ical factors s'c"

as an e%ile%tic a'to&atis&,
%ost-ictal conf'sional state, "ead

inH'ry, "y%o)lycae&ia, slee%;al5in) .4o;ard and
d+Orban,

19:3 A Cen;ic5, 1990 , 19980, or 72! .ra%id eye &ove&ent0 slee%

disorder
.,c"enc5 et al., 19:L A (lar5e et al., 2000 0. ,'c" %at"olo)y

can be )ro'nds for a so-
called ?insane+ a'to&atis&

in 2n)lis" la; .if t"e res'lt of a brain disease0 or a ?sane+

a'to&atis& .if t"e conse<'ence of an external a)ent0. Cor exa&%le,

t"e %resent a't"or
sa; a yo'n) diabetic %atient, nor&ally obsessive

in &aintainin) "is control, ;"o too5 "is
ins'lin before dinner,

delayed %re%arin) "is &eal ;"en "e ;as intri)'ed by a television

%ro)ra&&e and ;"o s'bse<'ently attac5ed "is flat&ate ;it" a

bread 5nife. On t"e arrival
of t"e %olice, t"e &an s"o;ed c"aracteristic

feat'res of "y%o)lycae&ia. 4e ;as ac<'itted
on t"e )ro'nds of

a ?sane+ a'to&atis&. Fn ot"er circ'&stances, a&nesia

per se does not
constit'te )ro'nds for alleviation of res%onsibility.

9&nesia for an offence is &ost co&&only associated ;it"1 .i0

states of extre&e
e&otional aro'sal, in ;"ic" t"e offence is

'n%re&editated, and t"e victi& 's'ally a lover,
;ife, or fa&ily

&e&ber, &ost co&&only in "o&icide .?cri&es of %assion+0.

Fn s'c" cases,
t"ere is often a %recedin) "istory of de%ression

and, on ?co&in) ro'nd+, t"e %erson
reali@es ;"at

"as been done and &ay ;ell call t"e %olice "i&selfP"erself .*'dHonsson

and
!ac$eit", 19:: A $o%el&an, 1995 b0. .ii0 9lco"olic intoxication

.so&eti&es in
association ;it" ot"er s'bstances0, 's'ally involvin)

very "i)" %ea5 levels as ;ell as a
lon) "istory of alco"ol ab'se.

9lco"ol can %rod'ce a&nesia fro& a so-called ?blac5o't+

or
as a state-de%endent %"eno&enon .*ood;in et al., 19L9 0. -"e

offence &ay vary fro&
cri&inal da&a)e, t"ro')" assa'lt, to "o&icide,

and t"e victi& &ay be 'n5no;n to t"e
offender. .iii0 Clorid

%syc"otic states1 occasionally, offenders describe a del'sional

acco'nt of ;"at "as "a%%ened .-aylor and $o%el&an, 19:K 0, <'ite

at odds ;it" ;"at
;as observed by ot"ers, so&eti&es res'ltin)

in confessions to ?cri&es+ t"at t"e %erson
co'ld

not act'ally "ave co&&itted .a ?%ara&nesia+ or ?del'sional

&e&ory+0.

5TSD
-,# can occ'r in association ;it" "ead inH'ry, road traffic

accidents, bein) t"e victi& of
violent cri&e, or in &aHor disasters

.e.). t"e sin5in) of t"e 4erald of Cree 2nter%rise at
Jeebr'))e,

t"e $in)+s (ross fire0. 9s is ;ell 5no;n, it is c"aracteri@ed

by intr'sive
t"o')"ts and &e&ories .?flas"bac5s+0

abo't t"e tra'&atic ex%erience. Bre;in and
collea)'es "ave %ost'lated

a d'al syste& of &e&ory re%resentation to try to acco'nt for

t"e s%ontaneity and invasiveness of t"ese flas"bac5s .Bre;in

et al., 199L A Bre;in, 2001
0. 4o;ever, t"ere &ay be instances

of %artial &e&ory loss .?fra)&entary+ &e&ories0,

distortions, or even fran5 confab'lations. Cor exa&%le, t"e

a't"or sa; a victi& of t"e
4erald of Cree 2nter%rise at Jeebr'))e,

;"o described tryin) to resc'e a close friend still
on board

t"e s"i%, ;"en ot"er ;itnesses re%orted t"at t"is close friend

"ad not been seen
fro& t"e &o&ent t"e s"i% t'rned over. 9lt"o')"

factors s'c" as "ead inH'ry or
"y%ot"er&ia &ay confo'nd t"e

inter%retation of s'c" cases, it is of interest t"at -,#
sy&%to&s

"ave been re%orted to occ'r even ;"en t"e s'bHect a%%ears to

"ave been
co&%letely a&nesic for t"e e%isode .!cNeil and *reen;ood,

199L A 4arvey et al., 2002
0. !oreover, -,# victi&s &ay s"o;

deficits in antero)rade &e&ory on for&al tas5s
&any years after

t"e ori)inal tra'&a .Bre&ner et al., 1998 0, and t"ere are clai&s

t"at
t"ey s"o; a loss of "i%%oca&%al vol'&e on !7F brain scan

.Bre&ner et al., 1995 0,
;"ic" "as been attrib'ted to effects

of )l'cocorticoid &etabolis& .!ar5o;itsc", 199L 0.
-"ese !7F

findin)s "ave to be inter%reted ;it" ca'tion in vie; of t"e

rat"er cr'de
&eas're&ents e&%loyed1 in t"e Bre&ner et al. st'dy,

even t"e control val'es ;ere a lon)
;ay o't of line ;it" t"ose

in &ost ot"er investi)ations .see (olc"ester et al., 2001 0.

9 recent revie; doc'&ented evidence of a&nesia in t"e victi&s

of li)"tnin) flas"es,
flood disasters, %i%eline ex%losions,

eart"<'a5es, concentration ca&% and "oloca'st
s'rvivors, ref')ees

and tra'&ati@ed soldiers fro& t"e t;o ;orld ;ars and Vietna&

.Bro;n et al., 1999 0. Ot"er revie;s "ave cited instances of

5idna% and tort're .van der
$ol5 and Cisler, 1995 0. -"e %syc"olo)ical

i&%act of a road traffic accident &ay also
)ive rise to a&nesia

.4arvey, 2000 A $o%el&an, 2000 c0, alt"o')" it &ay be diffic'lt

to
se%arate t"is fro& t"e effects of conc'ssion. Bro;n et al.

.19990 also fo'nd evidence of
for)ettin) in all L: st'dies t"at

t"ey revie;ed on t"e fra')"t iss'e of &e&ory for c"ild
sex'al

ab'se. ="ilst t"ere are indeed %roble&s in eval'atin) self-re%orts

of a&nesia for
c"ild ab'se, so&e s&aller scale st'dies "ave

exa&ined t"e corroborative evidence for t"e
tra'&a and t"e s'bse<'ent

for)ettin) in so&e detail .,c"ooler et al., 1993 0, and ot"ers

"ave %ro%osed &ec"anis&s ;"ereby s'c" for)ettin) and s'bse<'ent

.%ossibly erroneo's0
re-retrieval &i)"t occ'r .,c"acter, 199L A

,"i&a&'ra, 1993 0, %artic'larly in relation to
certain c'es or

tri))ers .9ndre;s et al., 2000 0.

6echanisms in#ol#ed in psycho"enic and or"anic amnesia
Ft "as been clai&ed t"at sit'ation-s%ecific a&nesia involves

a narro;in) of conscio'sness
;it" attention foc'sed on central

%erce%t'al details, so&eti&es evolvin) into a&nesia
.("ristianson,

19:K 0, andPor t"at e&otional or tra'&atic events are %rocessed

differently
fro& ?ordinary &e&ories+ .van der $ol5

and Cisler, 1995 0. Fn %artic'lar, e&otional
&e&ories, es%ecially

t"ose involvin) fear, &ay i&%licate a&y)daloid circ'its, distinct

fro& t"ose involved in ?nor&al+ learnin) .9dol%"s

et al., 199K A (a"ill et al., 1995 A
Oo'n) et al., 1995 A Cine

and Blair, 2000 0. 4o;ever, it &ay also be t"e case t"at, ;"en

so&et"in) extraordinary "a%%ens, ;e as5 o'rselves to recall

far &ore detail t"an ;e
;o'ld nor&ally ex%ect, and ex%erience

t"e s"ortfall as ?)a%s+ in &e&ory .$o%el&an,
2000 c0.

!ar5o;itsc" .199L 0 %ointed to vario's co&&onalities bet;een

%syc"o)enic and or)anic
a&nesia, and $o%el&an .2000 a0 %ro%osed

a &odel to try to ta5e acco'nt of t"e inter%lay
of %syc"osocial

factors and brain syste&s in %rod'cin) %syc"o)enic a&nesia.

Fn Ci). 1,
social and %syc"olo)ical factors are indicated in

t"e ovals, ;"ilst t"e relevant brain
syste&s are in t"e boxes.

="ilst stress so&eti&es "as a direct effect '%on t"e &edial

te&%oralPdience%"alic syste&, t"ereby %rod'cin) an i&%air&ent

in ne; learnin), t"e
&odel %ost'lates t"at, in ot"er cases,

stress %redo&inantly affects frontal
controlPexec'tive syste&s,

s'c" t"at t"e retrieval of a'tobio)ra%"ical &e&ories is
in"ibited.

-"is in"ibition ;ill be exacerbated, or &ade &ore li5ely, ;"en

a s'bHect is
extre&ely aro'sed, very de%ressed, or ;"en t"ere

"as been a ?learnin) ex%erience+ of a
%ast transient

or)anic a&nesia. Ff t"e stress is severe, t"e in"ibition &ay

even affect a
%ost'lated ?%ersonal se&antic belief syste&+,

res'ltin) in a transient loss of %ersonal
identity .das"ed arro;0.

Ff t"at occ'rs, t"ere is ?ne)ative feedbac5+ in

t"e for& of a severe
da&%enin) of t"e c'rrent e&otional state,

s'c" t"at a %atient in f')'e c"aracteristically
a%%ears e&otionally

?flat+ or %er%lexed, instead of aro'sed or de%ressed.

Fn s'c"
circ'&stances, ?ne;+ antero)rade learnin)

in res%onse to ?nor&al+ environ&ental sti&'li
can

still occ'r via t"e intact &edial te&%oralPdience%"alic syste&.

Fn brief, t"is &odel
adds to, b't is broadly consistent ;it",

5no;led)e of t"e brain syste&s i&%licated in
nor&al &e&ory and

or)anic a&nesia. !oreover, 9nderson and *reen .2001 0 "ave
recently

re%orted evidence t"at exec'tive &ec"anis&s can indeed be recr'ited

to %revent
'n;anted &e&ories fro& enterin) a;areness, and t"at

re%eated 'se of t"is strate)y
in"ibits t"e s'bse<'ent recall

of t"e s'%%ressed &e&ories.

View lar"er #ersion .81$01
Din t"is ;indo;E
Din a ne; ;indo;E

+i". 7 ,ocial factors and brain syste&s infl'encin)
a'tobio)ra%"ical &e&ory retrieval and %ersonal
identity. -"e fi)'re s"o;s .i0 frontal lobe %rocesses
involved in confab'lation .trace
s%ecificationPverificationA context &e&oryPso'rce
&onitorin)0, and .ii0 social factors .in ovals0 and
brain syste&s .in rectan)les0 involved in
%syc"o)enic a&nesia1 in"ibition leads to i&%aired
retrieval of incidents and facts, and severe in"ibition
affects orientation in %erson. .9da%ted fro& (o)n
Ne'ro%syc"ol 2000A 131 5:5ML21.0

Summary
syc"o)enic a&nesia can be eit"er local or sit'ation s%ecific.

-"e brain syste&s
&od'latin) e&otional &e&ories are be)innin)

to be 'nderstood, b't it is essential t"at
&odels of %syc"o)enic

a&nesia s"o'ld incor%orate t"e %syc"osocial context in ;"ic"

it
arises. -"at said, t"e dysf'nction can be 'nderstood in ter&s

of an interaction bet;een t"e
%redis%osin) %syc"osocial factors

and frontal in"ibitory &ec"anis&s o%eratin) ;it"in and
'%on

t"ose brain syste&s %revio'sly %ost'lated in st'dies of nor&al

&e&ory and or)anic
a&nesia.

ow and when do false memories arise(

Spontaneous confabulation in brain disease
Fn ?s%ontaneo's+ confab'lation, t"ere is a %ersistent,

'n%rovo5ed o't%o'rin) of erroneo's
&e&ories, often "eld ;it"

fir& conviction, so&eti&es bi@arre, and very often
%reocc'%yin).

Cor exa&%le, %atient 9B .$o%el&an et al., 1993 0 believed t"at

"er %arents
;ere fre<'ently visitin) "er in "os%ital .t"ey "ad

been dead K and 20 years0, and t"at "er
brot"er ;as a doctor

livin) on t"e 2Kt" floor of t"e b'ildin) s"e occ'%ied ."e ;as

not a
doctor, and s"e ;as on t"e to% floor, t"e 12t"0. ,%ontaneo's

confab'lation "as &ost
co&&only been described in association

;it" frontal lobe %at"olo)y, %artic'larly
involvin) t"e ventro-&edial

re)ions ./'ria, 193L A ,t'ss et al., 193: A $a%'r and
(o')"lan,

19:0 A ,"a%iro et al., 19:1 A Baddeley and =ilson, 19:L A !oscovitc"

and
!elo, 1993 0. 4o;ever, s%ontaneo's confab'lation can also

occ'r in ot"er, &ore
)enerali@ed disorders, s'c" as conf'sional

states .#e/'ca and (icerone, 1991 0 or t"e
)enerali@ed .&etabolic0

effects of carcino&a .$o%el&an et al., 1993 0. On t"e ot"er
"and,

frontal dysf'nction see&s to be a necessary, b't %er"a%s not

a s'fficient, condition
for s%ontaneo's confab'lation .$o%el&an

et al., 1993 A #alla Barba et al., 1999 01 it is a
tr'is& t"at

&any %atients ;it" frontal %at"olo)y do not confab'late, and

t"is does not
see& to be related %'rely to t"e %recise site

of %at"olo)y.

9 n'&ber of ex%lanatory t"eories for s%ontaneo's confab'lation

"ave been %ro%osed. -"e
first )ro'% of t"eories e&%"asi@es fa'lty

s%ecification and verification in &e&ory
retrieval. B'r)ess

and ,"allice .199L 0 %ost'lated deficits in a descri%tor %rocess,

an
editor %rocess, and a &ediator %rocess, ;"ic" all contrib'te

differently to t"e clinical
%"eno&ena of confab'lation. -"e

?descri%tor+ s%ecifies t"e ty%e of trace t"at ;o'ld

satisfy
t"e de&ands of a retrieval tas5, and ?noisy+

s%ecification increases t"e c"ance of an
ina%%ro%riate re%resentation

bein) %rod'ced as a candidate &e&ory. -"e ?editor+

c"ec5s
t"at t"e o't%'t fro& lon)-ter& stora)e fits ;it" %revio'sly

retrieved &e&ory ele&ents
and ;it" overall tas5 re<'ire&ents.

="en t"is ?editor+ is i&%aired, confab'lators res%ond

to a <'estion ;it"o't )ivin) it ade<'ate consideration, c"ec5in),

or self-correction. -"e
?&ediator+ controls strate)icP%roble&-solvin)

o%erations concernin) t"e ade<'acy or
%la'sibility of retrieved

&e&ory ele&entsA i&%air&ent res'lts in reasonin) errors and

in
bi@arre or ?fantastic+ res%onses .see also B'r)ess

and !cNeil, 1999 0. !oscovitc" and
!elo .1993 0 %ro%osed a so&e;"at

si&ilar t"eory, also identifyin) a n'&ber of %'tative
deficits

in c'e-retrieval, strate)ic searc" or fa'lty &onitorin), t"e

last res'ltin) in
erroneo's &e&ories not bein) edited o't or

s'%%ressed. /i5e;ise, ,c"acter et al. .199: 0
ar)'ed t"at an

ins'fficiently ?foc'sed+ retrieval descri%tion,

or an i&%air&ent in %ost-
retrieval &onitorin) and verification,

&ay )ive rise to confab'lation. Fn addition, t"ese
latter a't"ors

ar)'ed t"at encodin) i&%air&ents &ay &a5e s'bHects &ore liable

to
confab'latory errors at retrieval.

-"e second )ro'% of t"eories e&%"asi@es context'al &e&ory deficits,

%artic'larly ;it"
reference to te&%oral se<'ence andPor so'rce

&onitorin) deficits. $orsa5off .1::9 0
"i&self )ave several exa&%les

of conf'sions in te&%oral se<'ence, res'ltin) in
confab'lation,

s'c" as a ;o&an ;"ose acco'nt of %ast vacations conf'sed Ho'rneys

to
Cinland and t"e (ri&ea so t"at ;"olly ina%%ro%riate descri%tions

;ere )iven. Ot"er
a't"orities "ave also e&%"asi@ed te&%oral

context conf'sions as t"e basis of
confab'lation .e.). !oll,

1915 A Van der 4orst, 1982 A -alland, 19L5 A Victor et al., 1931
0.

!ost recently, ,c"nider et al. .199L 0 fo'nd t"at a s&all )ro'%

of s%ontaneo's
confab'lators ;ere differentiated fro& ot"er

a&nesic %atients and "ealt"y controls on t"e
basis of errors

at an ?i&%licit+ te&%oral context &e&ory tas5, b't

not on ot"er &e&ory or
exec'tive tas5s .see also ,c"nider and

ea5 , 19990. Fn a variant of t"is "y%ot"esis, #alla
Barba and

collea)'es "ave %ro%osed t"at ?te&%oral conscio'sness+

is intact b't
&alf'nctionin) in confab'latin) %atients .#alla

Barba, 1998a , b, 1999 A #alla Barba et
al., 1993 , 19990. ,'c"

%atients are a;are of a %ast, %resent and f't're .'nli5e severely

a&nesic %atients0 b't, in &a5in) te&%oral H'd)e&ents, t"ey e&%loy

only t"e &ost stable
ele&ents fro& t"eir lon)-ter& &e&ory stores.

9s5ed ;"at t"ey did yesterday or ;ill do
to&orro;, t"e %atients

re%ly ;it" ;ell-establis"ed ro'tines or t"e "abits of a lifeti&e,

"o;ever irrelevant to t"eir %resent sit'ation. !ore )enerally,

Io"nson and collea)'es
"ave ar)'ed t"at confab'lation &ay res'lt

fro& a ;ider ran)e of context, so'rce, or
reality &onitorin)

deficits .Io"nson, 1991 A Io"nson et al., 1998 , 20000. ,o'rce

&e&ory
or &onitorin) refers to t"e ability to identify t"e ?so'rce+

of infor&ation .,c"acter et al.,
19:K 0, and reality &onitorin)

to t"e ability to discri&inate ?real+ fro& i&a)ined

ex%erience.

9 t"ird )ro'% of t"eories e&%"asi@es t"at &'lti%le deficits

&ay contrib'te to
confab'lation. ,"a%iro et al. .19:1 0 e&%"asi@ed

.i0 i&%aired self-&onitorin), .ii0 a
res'ltin) fail're to in"ibit

&e&ory errors and .iii0 fre<'ent %erseverations as t"e basis

of
confab'lation1 eac" of t"ese deficits co'ld be related to

a different as%ect of
frontalPexec'tive f'nction. Fn a co&%arison

of a confab'latin) %atient and ot"er %atients
;it" frontal lesions,

Io"nson et al. .1993 0 concl'ded t"at confab'lation &ay reflect

an
interaction bet;een .i0 a vivid i&a)ination, .ii0 an inability

to retrieve a'tobio)ra%"ical
&e&ories syste&atically and .iii0

so'rce &onitorin) deficits. $o%el&an et al. .1993 0
fo'nd t"at

.i0 &any confab'lations &i)"t %la'sibly res'lt fro& t"e confo'ndin)

and
ina%%ro%riate retrieval of ?real+ &e&ory fra)&ents

o't of te&%oral se<'ence, b't t"at .ii0
ot"er confab'lations

res'lt fro& %erseverations, %artic'larly in se&antic &e&ory,

and .iii0
yet ot"ers a%%ear to be instantaneo's, ill-considered

and 'nc"ec5ed res%onses to
i&&ediate environ&ental and social

c'es.

In summary, it see&s li5ely t"at te&%oral context and so'rce

&onitorin) deficits
contrib'te to, b't are not t"e sole basis

of, t"e i&%air&ents of &e&ory s%ecification
.descri%tionA foc'sed

retrieval0 and verification .editin)A &onitorin)0 %ost'lated

in ot"er
t"eories. Ft see&s %la'sible t"at t"ese vario's deficits

reflect dysf'nction in differin)
co&%onents of frontal control

&ec"anis&s and t"at t"ey interact to)et"er .see Ci). 1 and
$o%el&an,

1999 A *ilboa and !oscovitc", 2002 0.

6omentary )0pro#o&ed1* confabulation
!'c" &ore co&&only, a&nesic %atients s"o; ?&o&entary+

or ?%rovo5ed+
confab'lations. -"ese are fleetin)

intr'sion errors or distortions &ade in res%onse to a
c"allen)e

to &e&ory, s'c" as a &e&ory test .Berlyne, 1932 A $o%el&an, 19:3 0.

Fn
a&nesic %atients, t"ese intr'sion errors are secondary to

t"e &e&ory deficit itself, rat"er
t"an necessarily reflectin)

frontalPexec'tive dysf'nction, H'st as, in "ealt"y s'bHects,

t"ey
arise ;"en &e&ory is ?;ea5+ for any reason.

,'c" errors reflect t"e essentially
?reconstr'ctive+

nat're of &e&ory retrieval1 ;"ere t"e trace is ;ea5 .e.). at

lon) delays0,
t"is reconstr'ction beco&es distorted or fran5ly

erroneo's, b't t"is does not necessarily
i&%ly a conscio'sPdeliberate

atte&%t to ?fill in+ &e&ory )a%s. -"e errors t"e&selves,

alt"o')" fleetin), can be <'ite stri5in) and s'r%risin). !o&entary

confab'lation in
"ealt"y s'bHects "as been de&onstrated in &any

ex%eri&ental investi)ations .Bartlett,
1982 A 4a&&ersley and

7ead, 19:L A $o%el&an, 19:3 A /indsay and 7ead, 199K A
7ead and

/indsay, 1993 A ,c"acter et al., 199: 0.

Fn a ne'roi&a)in) st'dy, ,c"acter et al. .199Ld 0 fo'nd t"at

t"e left &edial te&%oral
.%ara"i%%oca&%al0 &e&ory syste& ;as

activated d'rin) both acc'rate .?veridicial+0 and

fa'lty .?ill'sory+0 reco)nition of ;ord sti&'li.

9cc'rate reco)nition ;as associated ;it"
si)nificantly increased

blood flo; in t"e left te&%oro-%arietal cortex, ;"ereas fa'lty

reco)nition ;as associated ;it" activation in t"e %refrontal

and orbito-frontal cortices and
t"e cerebell'&, ;"ic" ;ere %ost'lated

as involved in error detection. -"e overla%
bet;een t"e brain

syste&s activated by tr'e and fa'lty &e&ories &ay "el% to ex%lain

t"e
%artic'lar salience of false &e&ories in &any circ'&stances.

+alse memories in other contexts
Calse &e&ories can also arise in &any ot"er circ'&stances .$o%el&an,

1999 0.
#el'sional &e&ories are a relatively rare %"eno&enon,

dis%layin) &any of t"e feat'res
of s%ontaneo's confab'lation,

exce%t t"at t"ey tend to revolve aro'nd a sin)le t"e&e and
are

not necessarily related to frontalPexec'tive dysf'nction .#avid

and 4o;ard, 199K A
$o%el&an et al., 1995 A Baddeley et al., 199L 0.

#el'sional &isidentification syndro&es
are often associated

;it" t"e conc'rrence of bilateral frontal lobe %at"olo)y and

ri)"t
%osterior lesions .9lexander et al., 1939 A 2llis and Oo'n),

1990 A Oo'n) et al., 1995 A
Ceinber) and 7oane, 1993 A Box et al.,

1999 A Ceinber) et al., 1999 0. (onfab'lations
"ave also been

described in sc"i@o%"renic %atients d'rin) story recall .Nat"aniel-Ia&es

and Crit", 199L A Nat"aniel-Ia&es et al., 199L 0, alt"o')" it

is %ossible t"at t"ese are
si&%ly ?&o&entary+ confab'lations,

colo'red by t"e %reocc'%yin) del'sional beliefs of
t"ese %atients

.$o%el&an, 1999 0. -"ere are also rare %syc"iatric syndro&es

in ;"ic"
%atients ado%t a false identity or identities, associated

;it" ?&e&ories+ in ;"ic" t"ey "ave
co&e to believe

.$o%el&an, 1999 0. -"ese incl'de ?%se'dolo)ia fantastica+,

so&eti&es
5no;n as ?%at"olo)ical lyin)+ .Cis", 19L3 0,

rare instances of %eo%le ;"o ado%t a ne;
role and identity follo;in)

a ty%ical f')'e e%isode .$o%el&an et al., 199K bA
!ar5o;itsc"

et al., 1993 a0 and so-called &'lti%le %ersonality disorder .dissociative

identity disorder0, in ;"ic" t"e ;idely varyin) )eo)ra%"ical

%revalence al&ost certainly
reflects differences in t"e reinforcin)

be"avio'r of doctors, %syc"olo)ists, and t"e o'tside
;orld .!ers5ey,

1992 , 19950.

!ost controversial are cases of alle)edly false &e&ory for c"ild

sex'al ab'se. -"e
%olari@ed %ositions of official re%orts .Brandon

et al., 199: A !orton et al., 1995 0
ill'strate t"e 'nresolved

and %olitical as%ects of t"is debate. 4o;ever, certain co&&on

t"e&es "ave e&er)ed in t"e scientific literat're on t"is to%ic.

2ven t"e &ost fort"ri)"t
%rota)onists of false &e&ory "ave fo'nd

t"at 19N of victi&s of ab'se re%orted for)ettin)
at so&e ti&e

in t"e %ast ./oft's et al., 199K 0. Ot"er researc"ers .,c"acter,

199L A
,c"acter et al., 1993 A ,"i&a&'ra, 1993 0 "ave s'))ested

t"at &e&ories for ab'se are
never act'ally co&%letely for)otten,

b't t"ey are retained in a va)'e 'nelaborated for&,
%oorly located

in te&%oral and s%atial context. ,'c" &e&ories ;o'ld be v'lnerable

to t"e
nor&al %rocesses of decay and interference, as ;ell as

to conscio's avoidance and
s'%%ression. 9%%ro%riate c'ein) &i)"t

res'lt in t"e re-retrieval of t"ese &e&ories, b't t"e
?;ea5

trace+ ;o'ld &a5e t"e& very v'lnerable to distortion and

a')&entation. Ft also
see&s very %la'sible t"at false &e&ories

for sex'al ab'se are &ost li5ely to arise in
%artic'lar social

contexts .$o%el&an, 1999 0.

9 related %"eno&enon is false confession to an offence, in ;"ic"

t"e victi& is oneself
rat"er t"an anot"er. *'dHonsson et al.

.1999 0 described a &an ;"ose conviction of
"o&icide ;as <'as"ed

after "e "ad served 25 years in %rison. 9t 13 years of a)e,

"e "ad
%ers'aded "i&self of "is ?)'ilt+ d'rin) K:

" of %olice intervie;s, in ;"ic" "e ;as in an
extre&ely distra')"t

and a)itated state ;it"o't a doctor or la;yer in attendance.

,"ortly
before t"e trial, and H'st after it "ad be)'n, "e ;as

)iven t"ree intervie;s 'nder sodi'&
a&ylobarbitone, ;"ic" ;ere

s'bse<'ently %art of t"e )ro'nds for a%%eal. 7ecently,
so&e;"at

si&ilar cases fro& 1952 and 19:8 "ave "ad t"eir convictions

overt'rned .7.
vers's 4ay *ordon, 2000A 7. vers's Cell, 20010.

*'dHonsson and collea)'es "ave
described t"e circ'&stances in

;"ic" %eo%le &a5e s'c" ?confessions+, ;"ic" t"ey

often
co&e to believe or ?internali@e+ .*'dHonsson

and !ac$eit", 19:2 , 19::, 1990A
*'dHonsson, 1992 0. Fn &any of

t"ese cases, so'rce &e&ory errors .abo't ;"ere
%artic'lar infor&ation

"ad been learned abo't t"e offence or victi&0 see& to "ave arisen

in t"e context of %ersonal self-do'bt .lo; self-estee& andPor

de%ression0 andPor external
coercion by t"e interro)ators. -"is

res'lted in a fa'lty attrib'tion, "eld s'bse<'ently ;it"
varyin)

de)rees of conviction.

Summary
Ft see&s t"at vario's deficits in frontal control &ec"anis&s,

involvin) trace s%ecification
or verification %rocesses and

context or so'rce &e&ory, &ay contrib'te to ne'rolo)ical
confab'lation.

B't false &e&ories can also arise in t"e absence of brain disease,

innoc'o'sly in t"e case of &o&entary confab'lation, or &ore

sinisterly in t"e case of
false &e&ory for sex'al ab'se or offences

.false confessions0. Fn t"ese latter instances
also, i&%air&ents

in context'al or so'rce &e&ory "ave been %ost'lated.

!onclusions

-"is revie; "as atte&%ted an overvie; of clinical &e&ory disorders

and selected
t"eoretical controversies t"at arise fro& t"e&.

!any of t"e %revio'sly %ost'lated
distinctions bet;een te&%oral

lobe and dience%"alic a&nesia .e.). in for)ettin) rates0
"ave

bro5en do;n. -"ere is even s'bstantial overla% bet;een t"ese

for&s of a&nesia and
t"at arisin) fro& lar)e frontal lobe lesions,

alt"o')" i&%ortant distinctions re&ain.
7ecent researc" in or)anic

a&nesia "as e&%"asi@ed i&%air&ents in ?bindin)+

different
ty%es of &aterial in antero)rade &e&ory, incl'din)

context'al infor&ation and
associations, and vario's distinctions

"ave been dra;n to e&%"asi@e t"e ty%es of &e&ory
c"aracteristically

affected or relatively s%ared in 991 recallPreco)nition &e&ory,

re&e&berin)P5no;in), recollectionPfa&iliarity and ex%licitPi&%licit

&e&ory. 4o; t"ese
vario's distinctions &a% %recisely on to one

anot"er re&ains controversial. One vie; of
79 is t"at &edial

te&%oralPdience%"alic str'ct'res are critical to &e&ory for&ation,

b't
t"at da&a)e to t"ese str'ct'res )ives rise to a relatively

brief .8 years or less0 79.
9ccordin) to t"is vie;, a s'%eri&%osed

retrieval deficit, related to ;ides%read frontal
andPor te&%oral

neocortical %at"olo)y, is re<'ired to %rod'ce a te&%orally extensive

79
.)reater t"an 8 years0 .e.). $o%el&an, 19:9 , 19910. syc"o)enic

factors co'ld
%reci%itate or exacerbate a retrieval deficit.

4o;ever, t"is ti&e-li&ited vie; of &edial
te&%oral lobe f'nction

is no; contested by t"ose ;"o %ost'late a &ore 'nifor& effect

of
&edial te&%oral lesions '%on 79 and 99 .Nadel and !oscovitc"

1993 A C'Hii et al.,
2000 A (i%olotti et al., 2001 0.

Fn t"is revie;, F "ave tried to e&%"asi@e co&&onalities .as

;ell as differences0 bet;een
lar)ely se%arate literat'res ;"ere

t"ey exist1 for exa&%le, bet;een t"e &ec"anis&s
'nderlyin) t"e

ne'rolo)ical and %syc"o)enic for&s of a&nesia and of false &e&ory

.confab'lation0. 9n i&%ortant %oint is t"at ne'ro%syc"olo)ical

a%%roac"es "ave been
"eavily infl'enced by t"e s'ccess of &od'lar

&odels in ex%lainin) ot"er ty%es of
co)nitive disorder, s'c"

as ac<'ired dyslexia and %roso%a)nosia. B't t"is a%%roac" "as

so&eti&es led to a so&e;"at li&itin) and static %ers%ective.

-"ere are %robably only a
finite n'&ber of f'nctional dissociations

to be discoveredA t"ey &ay "ave a n'&ber of
alternative ex%lanationsA

and t"eir ne'robiolo)ical stat's often re&ains 'nclear
.(ara&a@@a,

199: A $o%el&an, 2000 aA -yler and !oss, 2001 0. *reater e&%"asis

needs
to be %laced on a &ore dyna&ic a%%roac" ;"ic" exa&ines1

.i0 t"e interaction bet;een
different co)nitive syste&s .and

t"e ne'ral net;or5s or re)ions ;"ic" 'nderlie t"e&0, as
in ,no;den+s

investi)ations of t"e inter%lay bet;een a'tobio)ra%"ical and

se&antic
&e&ory in se&antic de&entia .,no;den, 2002 0A .ii0 t"e

deter&inants of differential
%atterns of co)nitive c"an)e t"ro')"

ti&e s'c" as t"e relative rates of recovery of 79 and
99 follo;in)

"ead inH'ryA and .iii0 t"e interaction of %syc"o)enic and &otivational

factors ;it" s%ecific brain syste&s or %at"olo)y, e.). follo;in)

"ead inH'ry, in
%syc"o)enic a&nesia, or in t"e ne'rolo)ical

and %syc"olo)ical for&s of false &e&ory
.confab'lation0. Fn

t"is to%ic, t"ere is still )reat sco%e for clinicians and researc"ers

to
infor& and learn fro& one anot"er.