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SPECIAL ARTICLES

Delusions and the Right Hemisphere: A Review of


the Case for the Right Hemisphere as a Mediator of
Reality-Based Belief
Lindsey Gurin, M.D., Sonja Blum, M.D., Ph.D.

Delusions are beliefs that remain fixed despite evidence that they are incorrect. Although the precise neural mechanism of
delusional belief remains to be elucidated, there is a predominance of right-hemisphere lesions among patients with de-
lusional syndromes accompanied by structural pathology, suggesting that right-hemisphere lesions, or networks with key
nodes in the right hemisphere, may be playing a role. The authors discuss the potential theoretical basis and empiric support
for a specific right-hemisphere role in delusion production, drawing on its roles in pragmatic communication; perceptual
integration; attentional surveillance and anomaly/novelty detection; and belief updating.
JNCN in Advance (doi: 10.1176/appi.neuropsych.16060118)

Delusions have fascinated clinicians, researchers and philoso- familiarity which determines…immediate recognition…
phers for centuries. While the DSM-IV-TR required that de- The patient, whilst picking up on a very narrow resemblance
between two images, ceases to identify them because of the
lusional beliefs be “false,” “based on incorrect inference” and
different emotions they elicit. (Capgras & Carrette 1924)5
“firmly sustained,” the updated DSM-5 defines delusions on
the basis of their fixedness alone, providing no commentary on While Capgras went on to propose a psychodynamic for-
their veracity or the reasoning used to reach them.1,2 This shift mulation of the syndrome based on oedipal conflict,5 modern
in definitional emphasis, from the content of beliefs to the te- studies in patients with what came to be known as the
nacity with which their proponents cling to them, parallels a Capgras delusion have borne out his original hypothesis.
historic shift in the general approach to delusions and the Using skin conductance response (SCR) as a marker of au-
mechanisms driving them. While psychoanalytic “motiva- tonomic activity, Ellis and Young6 found a SCR deficit in
tional” models of delusions once dominated, framing delusions Capgras patients presented with familiar faces, a finding that
as ego defenses protecting against distressing unconscious con- others have replicated.7,8 This model of the Capgras de-
flict with their unique content essential to understanding the lusion has at its core a fundamental abnormality of “covert,”
psychodynamic processes at play, more recent cognitive neu- or affective, facial recognition in the presence of preserved
roscience approaches suggest instead that a single unifying “overt,” or visual, recognition. In some patients with this
mechanism may be responsible for the broad range of de- abnormality—although notably not in all of them9—this dis-
lusional beliefs.3 Within these latter models, it is the way in crepancy is explained by way of delusion about a lookalike
which the belief is developed and maintained, rather than impostor.
content of the belief itself, which is most of interest. Capgras, a so-called monothematic delusion,10,11 is the
William James observed that delusions, in many cases, best studied of all the specific delusions in large part due to
were “certainly theories which patients invent to account its relative simplicity and the frequency with which it is
for their abnormal bodily sensations.”4 In their original encountered. Monothematic delusions in general have lent
1923 paper reporting the “illusion of doubles,” Capgras and themselves more easily to scientific study due to their highly
Reboul-Lachaux proposed an “agnosia of identification” trig- circumscribed nature. In addition to Capgras, other com-
gered by disconnect between cognitive and emotional recog- monly encountered types of monothematic delusions in-
nition of faces: clude Frégoli, in which strangers are believed to be close
Some faces that [the patient] sees with their normal features, friends or relatives in disguise12; Cotard, in which the indi-
the memory of which is not altered in any way, are never- vidual feels that he or she is dead13,14; mirror agnosia, in
theless no longer accompanied by this feeling of exclusive which the individual’s own reflection is believed to be

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DELUSIONS AND THE RIGHT HEMISPHERE

someone else15; reduplicative paramnesia, the belief that a internal and external events. Delusions arise as a cognitive
person or place has been duplicated16,17; anosognosia, de- attempt to explain this powerful, fundamental feeling
lusional denial of illness (often left hemiplegia after right about an event’s importance.32 Anosognosia, pathologic
hemisphere stroke)18,19 and the related phenomenon of unawareness of a neurologic deficit, is a delusional denial in
asomatagnosia, in which a body part is denied as one’s which a failure of normal sensory feedback (e.g., from a
own20; delusional jealousy (Othello Syndrome)21; delusional paralyzed limb, or from the visual system in the case of
fidelity (Reverse Othello Syndrome)22; and erotomania (de Anton syndrome) allows for the development of a belief that
Clérambault syndrome).23 should be, but is not, rejected on the basis of overwhelming
Polythematic delusions, in contrast, involve multiple de- evidence that it is incorrect.33 In all of these cases, as in
lusional beliefs, sometimes interrelated, covering a wide Capgras, the perceptual anomaly can exist without the de-
range of topics. The complex, multiform delusions professed lusion and thus must not be sufficient to produce it. A second
by famed mathematician John Nash during the height of process interfering with the rejection of implausible beliefs
his psychosis exemplify this phenomenon: Nash believed, at must be invoked.
various times, that he was at the center of a secret effort to Coltheart, Davies and colleagues propose that while a
build a world government and that he would be the Emperor delusion can be triggered by any of a variety of abnormal
of Antarctica within this government; that his picture was on perceptual experiences, the second factor common to all
the cover of Life magazine disguised as Pope John XXIII; delusions is likely a defective “belief evaluation system”
that he was the left foot of God and that God was walking the housed in the right frontal lobe.34 Devinsky similarly sug-
earth through him; and that he was a Go board upon which a gests that delusions arise when an unfettered left hemi-
“first-order” game was being played by his two sons while a sphere “creative narrator” is allowed to confabulate explana-
“second-order” game pitted him, as he put it later, “in an tions for experiences without the ongoing “monitoring of
ideological conflict between me, personally, and the Jews self, memory, and familiarity” normally offered by the right
collectively.”24,25 frontal lobe. Here, we examine the case for a right hemi-
The deficit in autonomic reactivity observed in Capgras sphere contribution to delusion production by way of four
provides support for a general model of delusions as arising interrelated lines of evidence relating to its roles in non-
out of a core perceptual anomaly, not unlike the model verbal communication; perceptual integration; attentional
proposed by James over a century ago. Maher26 first sug- surveillance and anomaly/novelty detection; and belief
gested that a sufficiently abnormal perception alone might updating.
be enough to produce a delusion. Noting that this model
would not explain individuals with abnormal perceptions
THE RIGHT HEMISPHERE AND DELUSIONS:
who did not develop delusions, Davies and colleagues pro-
A BRIEF HISTORY
posed a two-factor theory10 which accepted this initial
proposition and posited a second abnormality, this time of When structural or functional imaging abnormality can be
reasoning: faced with an unusual perceptual experience demonstrated with delusions, the right hemisphere is fre-
(“abnormal data”27), the patient in this model develops an quently implicated.35–39 Anosognosia has long been ob-
explanation that should be rejected but, due to some second served to occur disproportionately after right hemisphere
failure, erroneously is not. stroke as compared with left, as have asomatognosia, in
While the model’s empiric support comes largely from which the paralyzed limb is disowned, and somatoparaphrenia
studies of Capgras patients and has been applied primarily in which there is a delusional belief about the true identity
to misidentification delusions, its theoretical framework is or source of the limb.20 Delusional supernumerary limbs
consistent with and can incorporate other models of de- have also been reported with right hemisphere lesions.40
lusion. In the “comparator model” of delusions of control, Delusional misidentification syndromes in particular show
the primary anomalous data are proposed to be a deficit in a right hemisphere association,38,41,42 with specific case
using “efference copy” generated by corollary discharges reports and series associating right hemisphere pathology
from a motor command to predict its sensory consequences, with reduplicative paramnesia16,43,44; the Cotard delusion45,46;
yielding a subsequent mismatch of predicted and experi- the Capgras delusion 35 ; mirror agnosia 47–49 ; Fregoli
enced behavior that makes possible a belief that the move- syndrome50,51; and Othello syndrome.52,53 In a recent
ment was externally controlled.28,29 A similar hypothesis review of 61 case reports of delusional misidentifica-
regarding a mismatch between predicted and actual expe- tion syndromes associated with specific lesions, Darby and
rience of inner speech has been advanced to explain audi- Prasad found right hemisphere lesions in 92% of cases, with
tory hallucinations and related delusions of alien thought right frontal lobe lesions present in 63%.54 Preexisting bi-
insertion.30,31 In the “aberrant salience” model of schizo- lateral hemispheric pathology likely accentuates this effect:
phreniform psychosis, built on an understanding of dopa- Levine et al, in a study of 25 right hemisphere injured pa-
mine as central to mediating the experience of stimulus tients, found the existence of preexisting brain atrophy to
significance, a hyperdopaminergic state leads to an in- be significant in predicting a delusional syndrome, with
appropriately heightened degree of salience assigned to no clear significance attributed to lesion size or location

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GURIN AND BLUM

within the right hemisphere.55 Several functional imaging Broca69 localizing speech and language there, the right
studies examining Alzheimer’s disease (AD) patients with hemisphere has since proven itself to be a major mediator of
and without delusions found an association between the human experience, at times by way of more abstract, less
presence of delusion and relative right hemisphere (usu- tangible modulatory effects on cognition, emotion, and ver-
ally right frontal and/or temporal) hypometabolism or bal and behavioral output. Following an initial awareness
hypoperfusion.56–59 of its role in visuospatial orientation beginning in the
One meta-analysis of functional neuroimaging studies ex- 1940s,70,71 what was previously thought of as the “minor”
amining time perception in healthy controls and schizophrenic hemisphere has subsequently become known to play a major
patients showed significantly decreased activation of most role in spatial attention,72 mental manipulation of objects
right hemisphere regions during timing tasks as compared with in space73,74; and body image.18,75 Previously dismissed as
controls, suggesting a role for right hemisphere dysfunction lacking language, the right hemisphere is crucial for social
in the time perception abnormalities of schizophrenia.60 In a communication, mediating comprehension of emotional con-
series of small studies examining the P300 component of au- tent through interpretation of prosody, facial expressions
ditory event-related potentials (thought to reflect conscious and gestures.76 It is thought to control spontaneous facial
attention to a stimulus), patients with psychotic depression and expression of emotion, with the left face shown to express
delusional misidentification disorders showed reduced right emotion more intensely in healthy people 77 and right
frontal P300 amplitude, and the delusional misidentification hemisphere-damaged individuals demonstrating relative
patients showed an additional P300 amplitude reduction in reductions in spontaneous facial expressivity.78 Individuals
the right parietal region as well as increased P300 latency in with right hemisphere injuries have difficulty understanding
the central midline brain region.61 verbal humor,79 idioms,80 and metaphors.81,82 They perform
Some authors have reasoned that if right hemisphere un- worse than left-hemisphere aphasic patients on tests of
deractivity allows delusions to occur, then stimulating the connotations of words, even while their understanding of
right hemisphere might suppress them. Studies of cold water word denotation is generally preserved.83,84 Some func-
caloric vestibular stimulation (CVS) suggest that this may be tional imaging studies support a role for the right hemi-
true, at least to some extent. CVS produces widespread, sphere in the interpretation of metaphor,85,86 although
largely contralateral hemispheric activation. Left-sided others have disputed this.87,88 Beyond interpreting content
CVS can resolve, transiently, left hemispatial neglect,62,63 at the sentence level, the right hemisphere is thought to
somatoparaphrenia,64 and anosognosia65 after right-sided play a key role in organizing complex narrative material:
stroke. Levine and colleagues reported improvement in right hemisphere-injured individuals have difficulty mak-
delusions and anosgnosia in schizoaffective disorder and ing correct inferences89,90; distilling central themes (the
schizophrenia66 after left ear as opposed to right ear CVS, “gist” of a narrative) from complex linguistic material91–93;
and another group reported a case of improvement in con- integrating elements of a story into a coherent narrative94,95;
version disorder following left CVS.67 selecting appropriate endings to jokes79; and assessing
But even with multiple lines of inquiry providing highly plausibility of individual story elements.94 Right hemisphere
suggestive circumstantial evidence for a right hemisphere injured individuals have difficulty distinguishing lies from
role in delusion production, direct evidence and specific jokes and have demonstrated deficits in theory of mind.96
neurophysiologic models of the relationship are lacking. Their speech, described by the aphasiologist Myers in
Right hemisphere dysfunction remains a slippery suspect: 1977 as “copious and inappropriate… confabulatory, ir-
present at the scene of delusions too often to be chalked up relevant, literal, and occasionally bizarre,” is adequate at
to chance, but not often enough to be implicated directly, and the sentence level of language but fails at its pragmatic
at times occurring without any delusion at all. Here, we function.90,97
explore four of the right hemisphere’s purported functions While schizophrenia has been associated at various times
in depth and suggest that these functions, taken together, with both left-98 and right-sided99 dysfunction, numerous
subserve a right hemisphere-dominated grip on reality that studies have identified deficits in the pragmatic aspects of
becomes increasingly tenuous the more impaired these func- communication and understanding nonliteral speech100–102
tions become. While right hemisphere lesions likely do not and facial expressions,103 and it has been suggested that this
“create” delusions per se, and while there is no clear single deficit in discourse-level communication may in fact be a
anatomic location or network to blame when delusions core feature of the illness.104
arise, we suggest that these four right hemisphere func-
tions, when intact, work in tandem to provide at least a partial Perceptual Integration
barrier against delusional belief. When one or a combi- The right hemisphere is thought to play a dominant role in
nation of these functions fails, delusions may arise. our ability to integrate disparate perceptions into an overall
“gist” or “gestalt” comprehension.93,105 Studies of visuo-
Pragmatic Communication spatial processing using hierarchical visual stimuli—e.g.,
While the left hemisphere enjoyed early celebrity status in a large letter made up of smaller letters—have long sug-
the mid- to late 19th century thanks to the work of Dax68 and gested a model of lateralized function in which the left

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DELUSIONS AND THE RIGHT HEMISPHERE

hemisphere preferentially attends to an item’s component Attentional Surveillance, Self-Monitoring, and Novelty/
parts (“local” processing) while the right hemisphere at- Anomaly Detection
tends to the item’s overall contour and gestalt impression The right hemisphere provides ongoing attentional surveil-
(“global” processing).106–108 A similar phenomenon is lance of both hemifields in the visuospatial realm72 and is
demonstrated in music, with the right hemisphere pro- largely responsible for vigilance and detecting novel or in-
posed to play a role in global appreciation of melodic con- congruent stimuli across all perceptual modalities.116–119 It is
tour and meter while the left deals with the more local believed that it serves the same function at a heteromodal,
features of pitch intervals and rhythm.109,110 Some authors, conceptual level as well, providing ongoing monitoring of
noting parallels in right hemisphere patients’ visuospatial the self and its relationship to the environment and func-
and verbal deficits, have speculated that these may be two tioning as what Ramachandran has called an “anomaly
faces of a single central failure of perceptual and ideational detector.”120 The left hemisphere, focused on processing at
integration—i.e., global processing—based in the right hemi- the local level, seeks to establish order and consistency be-
sphere. Wapner and colleagues suggested that their right tween individual features; it is, as Gazzaniga writes, “con-
hemisphere patients’ difficulties organizing and compre- stantly looking for order and reason, even when there is
hending narratives might reflect a broader deficit in handling none—which leads it to make mistakes.”121 These mistakes
complex ideational materials.94 Benowitz and colleagues, create inconsistencies within the explanatory model and
finding a strong correlation between deficits in verbal story between the model and reality which Ramachandran argues
recall and visuospatial organization in right hemisphere in- are explained away by the left hemisphere until some anom-
jured patients, considered the same.91 Myers showed a cor- aly threshold is reached, at which point the right hemi-
relation between visuospatial integration and interpretive sphere “forces a Kuhnian paradigm shift”120,122 in order to
language ability, hypothesizing that the tangential, overinclusive develop an alternate, more workable hypothesis.
speech seen in right hemisphere injury might reflect a There is evidence that the right hemisphere plays a role in
higher level difficulty with conceptualizing situations and novelty detection, and problems with novelty detection have
using contextual cues to distinguish relevant from irrele- been linked to delusions.123 Novelty detection is a function
vant details.97 attributed to the hippocampus, specifically dopamine-related
Devinsky, in a discussion of the spectrum of disorders of gating at CA1; now, emerging evidence shows that dysfunc-
body image and ego boundaries found in right hemisphere tion in this novelty detection mechanism is related to
injury, argued for a dominant right hemisphere role in the psychosis and delusion.124,125 Notably, delusions correlated
most fundamental synthesizing task of all: construction of positively with the difference of the functional connectivity
the corporeal and psychological self.75 Bogousslavsky and of the right hippocampus with the frontal lobe, suggesting
Regli described a “response-to-next-patient-stimulation” that alterations of fronto-limbic novelty processing may
phenomenon in 11 right hemisphere stroke patients in contribute to the pathophysiology of delusions in patients
which these patients were observed to follow commands with acute psychosis.123
directed to other patients as though they were directed to Without appropriate salience given to novel and
them; interpreted by the authors as a variant of persevera- anomalous stimuli, right hemisphere injured patients are
tion, this behavior might also suggest the presence of inappropriately blasé about bizarre occurrences and con-
impaired ego boundaries in which self and other are fabulate explanations for how these might fit into a pre-
not clearly demarcated.111 Here, too, we find parallels in viously established framework. In a story retelling task,
schizophrenia. Patients with schizophrenia have difficul- while controls and left hemisphere injured patients looked
ties with complex visuospatial processing112,113 similar to puzzled on hearing nonsensical story elements and left them
those seen right hemisphere patients. Authors have long out on retelling, right hemisphere patients not only readily
suggested a primary role for heteromodal perceptual in- accepted these odd elements but added justifications for
tegration deficits in driving what Borda and Sass have them.94 Rather than being totally insensitive to incongruities
called a disorder of “basic-self experience” or “ipseity in the narrative, right hemisphere patients seemed “at least
disturbance”: here, failure to adequately integrate the tangentially aware that something does not fit and yet, are
multimodal sensory experience of existing as a “self ” either unwilling or unable to frankly label the anomalous
in reality disrupts a patient’s “grip” or “hold” on that element as such.”94
reality.114 Where the hold on reality has been disrupted, Anosodiaphoria, the inappropriate lack of concern about
delusions can seep in. Postmes and colleagues suggest that one’s illness that can occur with (and often outlasts)
such perceptual incoherence creates a “sensory vacuum” anosognosia after right hemisphere stroke, may similarly be
into which the brain pours imagined or remembered ex- understood as a failure to be adequately impressed by the
periences in an effort to reinstate coherence: “thus, sensory very salient fact of one’s own new neurologic deficit. Studies
coherence will be restored at the expense of reality moni- of insight in Alzheimer’s Disease have shown a correlation
toring,” and delusions and hallucinations “can be regarded between impaired insight and decreased right temporo-
as a ‘solution’ for incomprehensible, incoherent multisen- occipital perfusion on SPECT imaging126 and right lateral
sory experiences.”115 and dorsolateral frontal cortical perfusion on FDG-PET.127

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Furthermore, AD patients with lower right insula volume when switching tasks.147 On the Wisconsin Card Sorting
have worse awareness of memory (metamemory).128 In Test (WCST), an executive function task known to produce
frontotemporal dementia (FTD), patients with right frontal strong activation in the DLPFC, particularly on the right,148
dominant disease present with the behavioral variant of FTD patients with right frontal lobe tumors and patients with
which is characterized with reduced symptom awareness schizophrenia made perseverative errors at a rate that was
as compared with the nonfluent, agrammatic aphasia FTD equal to each other and significantly greater than that of
patients who have left frontal predominant disease and in normal controls and patients with left frontal and nonfrontal
whom symptom awareness is more frequently intact.129 tumors.149 Another functional imaging study examining
WCST performance after head trauma showed an inverse
Belief Updating relationship between perseverative responses and metabo-
The related tasks of recognizing that an explanatory model lism in the right, but not left, dorsolateral prefrontal cortex
has become outdated and shifting allegiance to a new, more and caudate nucleus.150
workable one comprise the function of belief updating. The
frontal lobes facilitate changing cognitive set, with the right
THE RIGHT HEMISPHERE AT THE INTERFACE OF
frontal lobe in particular dominant for updating beliefs and
SELF, ENVIRONMENT, AND REALITY
avoiding repetitive responses.130,131 The right dorsolateral
prefrontal cortex (DLPFC) is thought to play a major role in Woven together, these threads reveal a picture of a right
problem-solving in complex, “ill-structured” situations.132 hemisphere that is essential for our ability to create and
Drake and colleagues, in a series of studies in healthy indi- maintain accurate appraisals of mental objects holistically
viduals, showed that counter-attitudinal messages were and in context—be they simple visuospatial figures, complex
more persuasive, and disagreeing statements more readily narratives, or the self. Returning to the two-factor theory of
recalled, when heard from the left133,134; they hypothesized delusions, it follows that a primary somatic/perceptual ab-
this represented increased openness to cognitive set ad- normality creates an inconsistency in a previously functional
justment with relatively increased right hemisphere acti- explanatory model, which the relatively preserved left
vation. In studies using mixed-handedness as a marker for hemisphere does its best to explain while keeping the model
relatively stronger nondominant hemisphere influence,135 intact. Overly drawn to the left hemisphere task of con-
mixed-handers were found to be more gullible and easily necting individual dots, the right hemisphere injured patient
persuaded136; more apt to experience sensory illusions137; is unable to appreciate that the picture thus created is bi-
more prone to magical thinking138; and more likely to in- zarre and incoherent. If anomalies are noted, they lack the
ternalize false personality trait characterizations139 than cognitive and emotional valence usually accorded to strange
strong left- or right-handers. Strong-handers, meanwhile, or surprising occurrences and do not capture the attention
are suggested to be less sensation-seeking140; more likely to the way they should.
prefer authoritarianism and conservative politics140,141; and Sass and Byrom have described this phenomenon in
more likely to retain beliefs in creationism from childhood schizophrenia as an “anything-goes orientation” in which
despite extensive scientific evidence for evolution.140 Sharot patients “quickly identify, accept and take in stride phe-
and colleagues showed an increased ability to incorporate nomena that most people would find anomalous.”151 In the
new negative information into preexisting belief frameworks recent past, the default mode network (DMN) has gained
after transient disruption of the left—but not right—inferior attention as a possible mediator of function and dysfunction
frontal gyrus with repetitive transcranial magnetic stimula- of ‘real-time’ thought and belief monitoring and constraints.
tion (rTMS).142 Cacioppa, Petty and Quintanar, using elec- The DMN is associated with daydreaming, imaginative
troencephalography (EEG) to monitor cortical activity during planning, and stimulus-independent reflection, perhaps
exposure to pro- and counter-attitudinal beliefs, showed acting as a threshold between consciousness and behavior.
a relative shift in activity from left to right as subjects con- Studies of patients with schizophrenia consistently demon-
sidered issues for longer periods of time.143 strate DMN hyperactivation (i.e., impaired suppression) on a
Patients with right hemisphere injuries, predictably, have variety of cognitive tasks152 as well as decreased connectivity
difficulty updating their beliefs. In 16 unilateral anterior to task-positive right frontal networks.153,154 Sass and Byrom
temporal lobectomy patients given a problem solving task, proposed that an overactive default mode network (DMN)
Rausch found that while all patients had difficulty solving might be partly responsible for the “hyposalience” attributed
problems as compared with controls, patients with left to experiences by schizophrenic patients - experiences that
temporal lobectomies were more likely to shift from a hy- should trigger alarm bells for strangeness but nevertheless
pothesis even when it was correct, while right temporal do not.151 Gerrans, drawing on studies demonstrating an
lobectomy patients tended to maintain a hypothesis even anticorrelation between DMN activity and activity in task-
when told it was not.144 Right hemisphere patients persever- focused networks, has proposed a model in which the task-
ate more than left hemisphere patients on measures of de- negative DMN is inhibited (“supervised”) by right prefrontal
sign fluency145 and number fluency146 and they have more task-positive networks; when these fail, a person with a dis-
difficulty suppressing previously learned cognitive sets inhibited DMN is free to generate a range of beliefs across

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DELUSIONS AND THE RIGHT HEMISPHERE

a broad spectrum of likelihood, without the constraints deficit, whereas the latter might be a normal response to the
usually imposed by the reality-based, environment-surveying experience of impaired memory.159 In a subsequent study
right prefrontal cortex.155 Unimpressed by major inconsis- of 11 brain injured patients, Bajo, Kopelman and col-
tencies and unable to revise beliefs, patients cling to old ex- leagues found an association between severity of confabu-
planatory models while simultaneously acknowledging, either lation and severity of memory impairment and executive
explicitly or implicitly, the existence of contradictory in- dysfunction.160 Schnider notes that spontaneous confabu-
formation. This may explain, in part, the widely observed lation “constitutes a syndrome of profound derangement
phenomenon in delusional patients referred to as “double of thought,” rather than of memory per se, “in which the
bookkeeping,” in which the delusion is upheld even while concept of ongoing reality in thinking and planning is
other statements or behaviors suggest that the patient, on dominated by a patient’s past experiences and habits rather
some level, knows it is not true.156 than true ongoing reality; the confabulations are simply the
verbal manifestation of the thought disorder.”161 Noting
that cases of spontaneous confabulation reported in the lit-
DISCUSSION
erature invariably involve lesions of the anterior limbic
It is probably not the case that right hemisphere lesions structures, and specifically the posteromedial orbitofrontal
directly cause false beliefs; more likely, without the complex cortex (OFC) and its connections, he suggests a role for
cognitive skill set normally offered by the right frontal lobe, this set of structures in monitoring ongoing reality and
there may be fewer barriers preventing their occurrence. “constantly suppressing activated, but currently irrelevant,
Our experience of reality is mediated in part by the stories memories.”161 Confabulation, then, becomes a frontally-
we tell ourselves to explain it. With deficits in comprehen- mediated disorder of distinguishing “now” from “not-
sion of metaphor, difficulty interpreting nonverbal conver- now,”162 in which memories and associations are allowed to
sational cues, and impaired attention to unexpected events, bubble to the surface of conscious experience in a disorga-
right hemisphere patients are at a disadvantage when it nized, “incoherent and context-free” way.159 Gilboa has
comes to collecting the evidence they need to build a proposed an overarching failure of “strategic retrieval” of
workable explanatory hypothesis for any unusual or novel memories in confabulation, of which temporal confusion is
experience; and having constructed a hypothesis, they are one symptom.163 He describes two interacting memory
unable to evaluate its validity in context of their preexisting evaluation systems: an intuitive “feeling of rightness” at-
knowledge. Their tangential, over-inclusive speech and dif- tached to retrieved memories based on how well they fit
ficulty with organizing complex narrative materials likely with an overall cognitive schema; and a conscious monitor-
reflects a core deficit in filtering out irrelevant data – even ing process that checks these memories for internal and
when those data are their own memories. Because of their current contextual coherence. The first judgment, “rapid,
difficulties with nonliteral communication and affective automatic, and relatively impenetrable to reasoning,” is
regulation, their friends and loved ones really do behave thought to be housed in the ventromedial prefrontal cortex
differently around them, further widening the gap between (VMPFC); the latter system, in the dorsolateral prefrontal
the patient’s expectation of the world and what is actually cortex (DLPFC). When this system breaks down, confabu-
experienced and making it even more difficult for the patient lation may occur.163 Cabeza and colleagues, showing in-
to keep up with reality. creased left prefrontal cortex (PFC) activity on PET imaging
It is important to mention here, as part of a complete of recall tasks and increased right PFC activity on recall
discussion of false beliefs arising after brain injury, the tasks, similarly hypothesized a “production-monitoring”
phenomenon of confabulation. Notably described by Korsakoff framework in which the left PFC generates semantically
as “pseudoreminiscences” occurring in patients with guided information “whereas the right PFC is more involved
chronic alcohol use, seemingly out of proportion to the de- in monitoring operations, including the evaluation and ver-
gree of cognitive impairment otherwise present, confabula- ification of recovered information.”164
tion was historically thought of in terms of memory Whether or not delusions and confabulation in neuro-
dysfunction with false or distorted memories arising to fill logic patients are in fact two distinct processes, or if they are
amnestic patients’ gaps in recall.157 Berlyne, in 1972, defined merely variations along a single spectrum, has yet to be
confabulation as “a falsification of memory occurring in clear agreed upon in the literature. At minimum, it is likely that
consciousness in association with an organically derived their mechanisms overlap, and they may interact with each
amnesia.”158 other. Coltheart takes this approach in a discussion of pro-
Not all amnestic patients confabulate, however, and sub- voked confabulation—instances in which the confabulated
sequent treatments of the topic have introduced the im- content is offered not spontaneously, but in response to some
portance of ongoing reality-monitoring for effective use question or task—noting that delusional patients, amnestic
of memories relevant to the individual’s current situation. patients, and healthy controls all can be shown to confabu-
Kopelman first distinguished between “spontaneous” and late when asked to provide explanations for their own be-
“provoked” confabulation, arguing that the former likely havior in instances where a good explanation is lacking.165
required some frontal dysfunction in addition to a memory Using Capgras as an example, he suggests that the patient,

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deeply attached to a belief in an imposter but with no enough to produce delusions. The DLPFC is typically im-
plausible conscious explanation for this belief, might con- plicated in measures of executive function and the right
fabulate evidence to explain it. Linking this to Gopnik’s DLPFC in particular has been shown to play a role in sup-
discussion of the human “drive for causal knowledge” (the pressing perseverative behaviors, navigating complex situa-
successful culmination of which she likens to orgasm),166 tions, and exerting “inhibitory cognitive control on affective
Coltheart comments that the satisfaction derived from impulses, being therefore particularly critical to limiting the
reaching an explanation for a previously unexplained influences of impulses in decision making behavior.”175 It is
behavior may override any explanatory implausibility: possible that the DLPFC provides the “top-down” experi-
“it is better to have an explanation for a piece of one’s ential monitoring counterpart to the VMPFC’s “bottom-up”
behaviour—any explanation, no matter how bizarre—than ‘intuitive’ sense that is not accessible to consciousness, with
to have none.”165 delusions being allowed to arise when there are lesions to
One theme common to discussions of both delusions and both of these cortices or to essential connections between the
confabulation is that of a two-step process in which there two. Papageorgiou’s findings of reduced right frontal P300
is an initial automatic, preconscious, autonomic or affective amplitudes in both patients with delusional misidentification
experience; and a secondary conscious mechanism in which syndrome (DMS) and psychotic depression, but increased
this experience is evaluated with reference to a larger con- midline P300 latency only in DMS, would seem to support
text. Linking these two steps, in most instances, is a thought; this multifocal model.176 When this injury or disconnection
but whether the affective experience generates the thought occurs on the right, the right hemisphere-specific deficits
(as in Ellis and Young6; Kapur32; and Davies et al.10), or detailed above may make it difficult for patients to compen-
the thought is experienced as entering consciousness al- sate when interpreting the world around them.
ready somatically or affectively tagged (as in Gilboa167 and Delusions then, are neither a necessary outcome of right
Damasio168) remains yet to be determined. hemisphere injury nor solely dependent on the right hemi-
The ventromedial prefrontal cortex (VMPFC) looms sphere for their production; but it is significantly easier for
large in all of these conversations. The idea that ‘covert’, them to emerge when failures of pragmatic communication,
affective, autonomic, unconscious bodily processes might perceptual integration, attentional surveillance, and belief
surreptitiously influence conscious decision-making has updating are superimposed on impairments of executive
been advanced most notably by Damasio, whose “somatic function and preconscious autonomic processing. It is not
marker hypothesis” proposes that emotion, as registered in clear whether any one deficit is more important for the
the brain by its association with transient autonomic and production of delusions than the others, and it may be the
visceral changes, alters conscious cognition at an un- case that different combinations of deficits produce different
conscious level.168 Damasio argues that the VMPFC is es- delusional presentations. Future experimental work in this
sential for linking unconscious, affective awareness with area will continue to clarify specific right hemisphere net-
conscious cognition, likely due to its strong reciprocal con- works and their contribution to confabulated, delusional,
nections with the hippocampus and amygdala.168,169 Ex- and nonpathologic belief, and further extend our under-
perimentally, patients with VMPFC injury do not generate standing of the fragile nature of our relationship to reality and
appropriate skin conductance responses (SCRs) when the remarkable resourcefulness of the injured brain.
shown emotionally charged stimuli.170 On a gambling task,
they cannot adjust their behavior to account for biases that AUTHOR AND ARTICLE INFORMATION
are too subtle to identify overtly but that nevertheless drive From the Departments of Neurology and Psychiatry, New York Uni-
normal controls and patients without frontal injury to alter versity Langone Medical Center, New York (LG); and the Departments of
their behavior.171 Outside of the laboratory, these patients Neurology and Rehabilitation Medicine, New York University Langone
Medical Center, New York (SB).
cannot link preconscious emotional awareness and intuition
Send correspondence to Dr. Gurin; e-mail: Lindsey.Gurin@nyumc.org
with conscious decision-making, leading to poor choices
particularly in the social and interpersonal realm as well as The authors report no financial relationships with commercial interests.

in risk assessment and outcome prediction; in the lab and in Received June 14, 2016; revisions received Oct. 29, and Jan. 5, 2017;
accepted Jan. 16, 2017.
life, they continue to play from “bad decks” long after ev-
eryone else has perceived a bias and changed course. The
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