The sense of self in the aftermath of trauma: Lessons from the default mode network in posttraumatic stress disorder

Abstract

Trauma can profoundly affect the sense of self, where both cognitive and somatic disturbances to the sense of self are reported clinically by individuals with posttraumatic stress disorder (PTSD). These disturbances are captured eloquently by clinical accounts, such as, ‘I do not know myself anymore,’ ‘I will never be able to experience normal emotions again,’ and, ‘I feel dead inside.’ Self-related thoughts and experiences are represented neurobiologically by a large-scale, cortical network located along the brain’s mid-line and referred to as the default mode network (DMN). Recruited predominantly during rest in healthy participants, the DMN is also active during self-referential and autobiographical memory processing – processes which, collectively, are thought to provide the foundation for a stable sense of self that persists across time and may be available for conscious access. In participants with PTSD, however, the DMN shows substantially reduced resting-state functional connectivity as compared to healthy individuals, with greater reductions associated with heightened PTSD symptom severity. Critically, individuals with PTSD describe frequently that their traumatic experiences have become intimately linked to their perceived sense of self, a perception which may be mediated, in part, by alterations in the DMN. Accordingly, identification of alterations in the functional connectivity of the DMN during rest, and during subliminal, trauma-related stimulus conditions, has the potential to offer critical insight into the dynamic interplay between trauma- and self-related processing in PTSD. Here, we discuss DMN-related alterations during these conditions, pointing further towards the clinical significance of these findings in relation to past- and present-centered therapies for the treatment of PTSD.

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European Journal of Psychotraumatology
ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/zept20
The sense of self in the aftermath of trauma:
lessons from the default mode network in
posttraumatic stress disorder
Ruth A. Lanius , Braeden A. Terpou & Margaret C. McKinnon
To cite this article: Ruth A. Lanius , Braeden A. Terpou & Margaret C. McKinnon (2020)
The sense of self in the aftermath of trauma: lessons from the default mode network in
posttraumatic stress disorder, European Journal of Psychotraumatology, 11:1, 1807703, DOI:
10.1080/20008198.2020.1807703
To link to this article: https://doi.org/10.1080/20008198.2020.1807703
© 2020 The Author(s). Published by Informa
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CLINICAL PRACTICE PAPER
The sense of self in the aftermath of trauma: lessons from the default mode
network in posttraumatic stress disorder
Ruth A. Lanius
a,b,c
*, Braeden A. Terpou
a
* and Margaret C. McKinnon
d,e,f
a
Department of Neuroscience, Western University, London, Ontario, Canada;
b
Department of Psychiatry, Western University, London,
Ontario, Canada;
c
Imaging Division, Lawson Health Research Institute, London, Ontario, Canada;
d
Mood Disorders Program, St. Joseph’s
Healthcare, Hamilton, Ontario, Canada;
e
Department of Psychiatry and Behavioural Neurosciences, McMaster University, Hamilton,
Ontario, Canada;
f
Homewood Research Institute, Guelph, Ontario, Canada
ABSTRACT
Trauma can profoundly affect the sense of self, where both cognitive and somatic distur-
bances to the sense of self are reported clinically by individuals with posttraumatic stress
disorder (PTSD). These disturbances are captured eloquently by clinical accounts, such as,
‘I do not know myself anymore,’ ‘I will never be able to experience normal emotions again,’
and, ‘I feel dead inside.’ Self-related thoughts and experiences are represented neurobiolo-
gically by a large-scale, cortical network located along the brain’s mid-line and referred to as
the default mode network (DMN). Recruited predominantly during rest in healthy partici-
pants, the DMN is also active during self-referential and autobiographical memory proces-
sing – processes which, collectively, are thought to provide the foundation for a stable sense
of self that persists across time and may be available for conscious access. In participants
with PTSD, however, the DMN shows substantially reduced resting-state functional connec-
tivity as compared to healthy individuals, with greater reductions associated with heigh-
tened PTSD symptom severity. Critically, individuals with PTSD describe frequently that their
traumatic experiences have become intimately linked to their perceived sense of self, a
perception which may be mediated, in part, by alterations in the DMN. Accordingly,
identification of alterations in the functional connectivity of the DMN during rest, and
during subliminal, trauma-related stimulus conditions, has the potential to offer critical
insight into the dynamic interplay between trauma- and self-related processing in PTSD.
Here, we discuss DMN-related alterations during these conditions, pointing further towards
the clinical significance of these findings in relation to past- and present-centred therapies
for the treatment of PTSD.
El sentido del Yo luego de experimentar Trastorno de Estrés
Postraumático: Lesiones desde la Red Neuronal por defecto
El trauma puede afectar profundamente el sentido del Yo, donde tanto perturbaciones
cognitivas como somáticas del sentido del Yo han sido reportadas clínicamente por indivi-
duos con trastorno de estrés postraumático (TEPT). Estas perturbaciones son capturadas de
forma elocuente por relatos clínicos como ‘ya no me conozco a mí mismo(a)’, ‘nunca más
volveré a ser capaz de experimentar emociones normales’, y ‘me siento muerto(a) por
dentro’. Los pensamientos y experiencias relacionadas con el Yo se representan
neurobiológicamente por una red cortical a gran escala, localizada entre la línea media
del cerebro, conocida como la red neuronal por defecto (DMN por sus siglas en ingles).
Reclutada predominantemente durante el reposo en pacientes sanos, la DMN también está
activa durante el procesamiento de memorias autobiográficas y auto-referentes, procesos a
los que comúnmente se les atribuye la función de proveer la base para un sentido estable
del Yo, que persiste a lo largo del tiempo y puede estar disponible para el acceso con-
sciente. En participantes con TEPT, sin embargo, la DMN muestra conectividad de reposo
disminuida comparada con la de individuos sanos, con mayores disminuciones asociadas
con mayor severidad en síntomas de TEPT. De manera crítica, individuos con TEPT describen
con frecuencia que sus experiencias traumáticas se han vuelto íntimamente relacionadas
con su percepción del sentido del Yo, que puede estar mediada, en parte, por alteraciones
en la DMN. Por lo tanto, la identificación de alteraciones en la conectividad funcional de la
DMN en reposo, y durante condiciones subliminales y estímulos relacionados con el trauma,
tiene el potencial de ofrecer una introspección crítica hacia la interacción dinámica entre el
trauma y el procesamiento relacionado al self en el TEPT. En este estudio, discutimos las
alteraciones relacionadas con la DMN durante estas condiciones, apuntando hacia la sig-
nificancia clínica de estos hallazgos en relación a las terapias centradas en el presente y el
pasado, para el tratamiento del TEPT.
ARTICLE HISTORY
Received 23 April 2020
Revised 10 July 2020
Accepted 30 July 2020
KEYWORDS
default mode network;
PTSD; trauma; sense of self;
posterior cingulate cortex;
medial prefrontal cortex;
somatic; present-centred
treatment; past-centred;
treatment; MDMA
PALABRAS CLAVE
Red Neuronal Por Defecto;
TEPT; Trauma; Sentido del
Yo; Corteza Cingulada
Posterior; Corteza Prefrontal
Medial; Somático; Terapia
Centrada en el Presente;
Terapia Centrada en el
Pasado; MDMA
关键词
默认模式网络; PTSD; 创伤;
自我意识; 后扣带回皮层;
前额叶内侧皮层; 躯体; 关
注当下的治疗; 关注过去;
治疗; MDMA
HIGHLIGHTS:
• Trauma may impact
significantly an individual's
sense of self.
• The default mode network
is thought to confer a stable
sense of self.
• We discuss alterations in
connectivity in this network
in PTSD and their clinical
significance.
CONTACT Ruth A. Lanius ruth.lanius@lhsc.on.ca Department of Psychiatry, Western University, University Hospital (Room C3-103), 339
Windermere Road, London, ON N6A 5A5, Canada
*
These authors contributed equally to this work
EUROPEAN JOURNAL OF PSYCHOTRAUMATOLOGY
2020, VOL. 11, 1807703
https://doi.org/10.1080/20008198.2020.1807703
© 2020 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial License (http://creativecommons.org/licenses/by-nc/4.0/),
which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

创伤后应激障碍之后的自我意识: 来自默认模式网络的经验
摘要: 创伤会深刻影响自我意识, 创伤后应激障碍 (PTSD) 患者临床上会报告自我意识的认
知和躯体障碍。这些障碍可被临床报告充分捕捉到, 例如‘我不再了解自己”, “我将再也无法
体验正常的情绪”和“我内心生不如死”。自我相关想法和体验在神经生物学上由位于大脑
中线的大面积皮层网络呈现, 被称为默认模式网络 (DMN) 。 DMN主要是在健康参与者静
息状态下收集的, 在自我参照和自传体记忆处理过程中也很活跃, 而这些过程共同被认为
可为跨时间持续存在的自我意识提供稳定的基础, 并且可用于有意识的访问。但是, 相较
于健康个体, 患有 PTSD 的参与者的 DMN 表现出明显降低的静息态功能连接, 越高的
PTSD 症状严重程度降低程度更大。至关重要的是, PTSD 患者经常描述他们的创伤经历已
经与他们的感知自我意识紧密联系在一起, 这种感知可能被 DMN 的改变部分中介。因此,
识别 DMN 在静息态以及阈下创伤相关刺激条件下功能连接的改变, 有可能为 PTSD 中创伤
和自我相关加工之间动态相互作用提供关键见解。在这里, 我们讨论了在这些情况下与
DMN 相关的改变, 进一步指出了这些发现与关注过去和关注当下的 PTSD 治疗相关的临床
意义。
1. The sense of self in the aftermath of
trauma
Trauma can profoundly affect the sense of self
(Schore, 2003) – leaving a lasting imprint on both
the cognitive and the somatic domains of an indivi-
dual’s sense of self (Lanius, Frewen, Tursich, Jetly, &
McKinnon, 2015; for a review, see Frewen et al., 2008,
2020). Cognitively, individuals who have experienced
trauma are often tormented by thoughts that reflect
intensely negative core beliefs about themselves,
which can include, ‘I will never be able to feel normal
emotions again,’ ‘I feel like an object, not like a
person,’ ‘I do not know myself anymore,’ or, ‘I have
permanently changed for the worse’ (Cox, Resnick, &
Kilpatrick, 2014; Foa, Tolin, Ehlers, Clark, & Orsillo,
1999). Somatically, recent research points increas-
ingly towards the notion that trauma can leave a
lasting physical representation, where lower back
pain, general muscle aches and pains, flatulence/
burping, or feeling as though your bowel movement
has not finished have been identified as somatic dis-
turbances that significantly perturb the sense of self
(Graham, Searle, Van Hooff, Lawrence-Wood, &
McFarlane, 2019). Here, Graham et al. (2019) found
that two thirds of cases of military-related posttrau-
matic stress disorder (PTSD) are missed when a
PTSD checklist for the Diagnostic and Statistical
Manual (DSM-5; APA, 2013) (i.e. PCL-5) served as
the only assessment tool; these cases are, however,
captured when physical symptoms are considered in
conjunction with the PCL-5. Moreover, participants
with PTSD report somatically-based alterations in
relation to self-experience, including feelings of dis-
embodiment and related identity disturbances,
revealed by reports like, ‘I feel dead inside,’ ‘I feel as
if I am outside my body,’ ‘I feel like my body does not
belong to me,’ or, ‘I feel like there is no boundary
around my body’ (Bernstein & Putnam, 1986; Foa
et al., 1999; Briere & Runtz, 2002; Frewen & Lanius,
2015; for a review, see Frewen et al., 2008, 2020).
These reports underscore the vulnerability the sense
of self has in the aftermath of trauma, where both
cognitive and somatic disturbances to the sense of self
are thought to reflect remnants of the traumatic past
among individuals with PTSD.
2. Neural underpinnings of the sense of self:
the default mode network
A wide body of evidence suggests that self-referential
processes, as well as past- and future-related autobio-
graphical memory processing, are facilitated by a
large-scale, intrinsic network known as the default
mode network (DMN) (Greicius, Krasnow, Reiss, &
Menon, 2003; for a review, see Raichle, 2015). Self-
referential processes define the various self-related or
social-cognitive functions that allow us to gain insight
and to draw inferences related to our own mental and
physical conditions, as well as to mentalize these alike
conditions in others (Greicius et al., 2003). Although
the DMN is recruited predominantly during rest, it is
also active during internally-directed cognitive pro-
cesses (for a review, see Raichle, 2015). Collectively,
these DMN-mediated processes are thought to pro-
vide the foundation for a continued experience of the
self across time, occasionally referred to as ‘autonoe-
tic consciousness’ (Fransson, 2005; Piolino et al.,
2006; Tulving, 1985), where self-relevant information
and events associate to produce our sense of self
(Conway & Pleydell-Pearce, 2000; Levine, Svoboda,
Hay, Winocur, & Moscovitch, 2002).
The DMN is composed primarily by cortical
regions located across the brain’s mid-line, including
the posterior cingulate cortex, the precuneus, and the
medial prefrontal cortex (Spreng, Mar, & Kim, 2009;
Greicius et al., 2003; Buckner, Andrews-Hanna, &
Schacter, 2008; for a review, see Qin & Northoff,
2011). These DMN-related cortices contribute var-
iously to self-related processes. Whereas the posterior
cingulate cortex and the precuneus are associated
more strongly with our experience of having an
embodied self that exists in space, the medial pre-
frontal cortex is associated more strongly with our
awareness of thoughts and emotions related to the
2R. A. LANIUS ET AL.

self (for a review, Qin & Northoff, 2011). Critically,
the DMN displays widespread functional alterations
in individuals with PTSD during rest (Akiki et al.,
2018; Brown et al., 2014; for a review, see Moore &
Zoellner, 2007; St Jacques, Kragel, & Rubin, 2013;
Nazarov et al., 2014, 2015; Wang et al., 2016; Koch
et al., 2016; Barredo et al., 2018; Akiki, Averill, &
Abdallah, 2017) and during trauma-related stimulus
conditions (Terpou et al., 2019a; for a review, see St
Jacques et al., 2013; Freeman, Hart, Kimbrell, & Ross,
2009), where these alterations are likely to mediate
the clinical disturbances underlying self-related pro-
cesses observed in PTSD.
3. The default mode network at rest in the
aftermath of trauma
Altered functionality across large-scale, resting-state
networks can be evaluated using functional connec-
tivity – an endeavour that has proven vital to identi-
fying the neural correlates underlying PTSD.
Functional connectivity estimates the degree to
which a particular brain region (i.e. seed) reveals
neuronal coupling across the brain by correlating
the spontaneous low-frequency activity of the seed
across the remaining whole-brain voxels (Friston,
1994). In PTSD, the DMN displays reduced func-
tional connectivity during rest as compared to
healthy controls (Figure 1(a)) (Bluhm et al., 2009;
Sripada et al., 2012). In particular, reduced resting-
state functional connectivity has been observed for
each DMN-related hub as exhibited with other
DMN-related brain regions in participants with
PTSD as compared to healthy individuals (posterior
cingulate cortex: Lanius et al., 2010; Wu et al., 2011;
Sripada et al., 2012; Miller et al., 2017; precuneus:
Bluhm et al., 2009; Di et al., 2012; Reuveni et al.,
2016; medial prefrontal cortex: DiGangi et al.,
2016). Reduced functional connectivity between
DMN hubs and the DMN more generally points
strongly towards a diminished coherence across the
network, where stronger reductions in resting-state
functional connectivity across the DMN are asso-
ciated with greater PTSD symptom severity (Bluhm
et al., 2009; Di et al., 2012; Shang et al., 2014; Sripada
et al., 2012). Critically, reduced resting-state func-
tional connectivity across the DMN among partici-
pants with PTSD has been replicated by multiple
research groups and across varying trauma exposures
(combat exposure: DiGangi et al., 2016; Reuveni
et al., 2016; King et al., 2016; Kennis, Van Rooij,
Van Den Heuvel, Kahn, & Geuze, 2016; interpersonal
trauma: Lanius et al., 2010; Bluhm et al., 2009; acute
trauma: Wu et al., 2011; Patriat, Birn, Keding, &
Herringa, 2016; Lu et al., 2017).
Although PTSD symptoms can emerge following
trauma(s) that are experienced well into adulthood,
Daniels, Frewen, McKinnon, and Lanius (2011) have
proposed that early-life trauma(s) may interfere par-
ticularly with the developmental trajectory of the
DMN – beginning in childhood and maturing well
into adolescence and early adulthood (Supekar et al.,
2010; Sripada, Swain, Evans, Welsh, & Liberzon,
2014; Sherman et al., 2014; for a review, see Fair
Figure 1. Images show the functional connectivity of the DMN in healthy controls (left) and in participants with PTSD (right)
under different conditions. Top and bottom images depict within-group patterns in functional connectivity during rest and
during trauma-related stimulus processing, respectively. Whereas resting-state functional connectivity is depicted in relation to
the time series of the posterior cingulate cortex (PCC), trauma-related functional connectivity is depicted in relation to the time
series of the periaqueductal grey (PAG). Figure 1 is an adaptation from two previous findings, where resting-state and threat-
related functional connectivity are related to results by Bluhm et al. (2009) and Terpou et al. (2019a), respectively.
EUROPEAN JOURNAL OF PSYCHOTRAUMATOLOGY 3

et al., 2008). Specifically, resting-state functional con-
nectivity between the posterior cingulate cortex and
the medial prefrontal cortex is thought to play an
important role in the development of self-related
and social-cognitive functions. Both DMN hubs
reveal reduced resting-state functional connectivity
in individuals with PTSD (Bluhm et al., 2009; Miller
et al., 2017; Sripada et al., 2012) – an observation that
has been linked, in part, to childhood maltreatment
or neglect. For example, Sripada et al. (2014) found
that adults reporting a history of childhood abuse are
more likely to demonstrate not only decreased DMN
functional connectivity, but also greater cortisol levels
in response to a perceived social stress. Accordingly,
these results have been taken as evidence for the
psychopathological vulnerability the DMN shows in
relation to adverse environmental conditions, parti-
cularly in childhood, where reduced resting-state
functional connectivity across the DMN may harbour
the vestiges of an individual’s trauma history, or,
alternatively, may serve as a predisposing factor
towards the development of PTSD.
The DMN can be divided into two dissociable
subsystems – the medial prefrontal and the medial-
temporal subsystem. These subsystems are recruited
during autobiographical memory processes, where
the medial prefrontal and the medial-temporal sub-
system are thought to govern different autobiogra-
phical memory processes, namely construction and
elaboration, respectively (Buckner & Carroll, 2007;
Cabeza & St Jacques, 2007; Svoboda, McKinnon, &
Levine, 2006). Autobiographical memory-related
construction refers to early memory formation pro-
cesses, where the semantic and contextual informa-
tion that provide the setting for a memory to be
recalled are brought to mind – a process mediated
mostly by the medial-prefrontal subsystem.
Autobiographical memory-related elaboration refers
to subsequent memory formation processes, where
self-referential perspectives and visual imagery pro-
cesses allow a memory to be re-experienced by recal-
ling its more salient characteristics (i.e. emotional,
sensory) – a process mediated by the medial-tem-
poral subsystem (along with the posterior parietal
cortex) (Buckner et al., 2008; Kim, 2012; Philippi,
Tranel, Duff, & Rudrauf, 2013).
In PTSD, reduced resting-state functional connec-
tivity between DMN hubs and the DMN more gen-
erally appears concentrated predominantly in the
medial-temporal subsystem (Bluhm et al., 2009;
DiGangi et al., 2016; Miller et al., 2017; Sripada
et al., 2012) – a pattern related likely to the altered
autobiographical memory-related re-experiencing
observed among individuals with PTSD (for a review,
see Brewin, 2015). Moreover, reduced functional con-
nectivity between DMN-related hubs and the medial-
temporal subsystem correlates negatively to symptom
severity in avoidance and numbing measures admi-
nistered to participants with PTSD (Miller et al.,
2017). Accordingly, reduced DMN functional con-
nectivity may promote clinical disturbances in self-
related processes during rest in PTSD, which can
include guilt, shame, or remorse emotional experi-
ences, as well as alterations in perceptual experiences,
including self-perceptions of body awareness that are
perturbed during moments of depersonalization and/
or derealization (Barredo et al., 2018; Bluhm et al.,
2009; Cloitre, Scarvalone, & Difede, 1997; Frewen
et al., 2008, 2020; van der Kolk, Roth, Pelcovitz,
Sunday, & Spinazzola, 2005). To date, however,
these hypotheses remain to be tested directly in indi-
viduals with PTSD.
4. The default mode network under threat in
the aftermath of trauma
In PTSD, the DMN also demonstrates altered func-
tional connectivity during threat- or trauma-related
conditions – a pattern that contrasts sharply with the
DMN deactivation observed usually in healthy indi-
viduals. Threat- or trauma-related information is
processed generally by an inter-connected network
comprised of brainstem, midbrain, and thalamic
structures, jointly referred to as the innate alarm
system (IAS) (Liddell et al., 2005). The IAS detects
threat-related information at the level of the mid-
brain, where the information is then transmitted to
the frontolimbic neural cortices (Tamietto & de
Gelder, 2010). Critically, the IAS bypasses the pri-
mary sensory processing cortices, thus allowing threat
information to be transmitted rapidly and permitting
the network to respond to a stimulus that has been
presented subliminally (i.e. below conscious thresh-
old) (Tamietto & de Gelder, 2010; Williams et al.,
2006). In PTSD, the IAS demonstrates increased
activity during subliminal, trauma-related stimulus
conditions as compared to healthy individuals
(Felmingham et al., 2008; Bryant et al., 2008;
Rabellino, Densmore, Frewen, Théberge, & Lanius,
2016; Terpou et al., 2019b; for a review, see Lanius
et al., 2017), where a trauma-related stimulus refers to
self-generated material in relation to an individual’s
trauma memory, or, in healthy individuals, a highly
aversive/stressful memory. In PTSD, subliminal,
trauma-related stimulus conditions evoke stronger
activity across the IAS, including the amygdala
(Bryant et al., 2008; Felmingham et al., 2008; Steuwe
et al., 2014), the midbrain periaqueductal grey (PAG)
(Rabellino et al., 2016; Terpou et al., 2019b), as well as
the brainstem more generally (Felmingham et al.,
2008). In our own research, we have postulated that
the IAS overactivation occurs in association with the
increased hypervigilance and hyperarousal symptoms
reported among individuals with PTSD – symptoms
4R. A. LANIUS ET AL.

that are coordinated largely by the midbrain and, in
particular, by the PAG (for a review, see Terpou et al.,
2019c).
More recent works points towards the PAG as
central to threat-related processing in PTSD. Here,
the PAG refers to the grey matter located around the
cerebral aqueduct of the midbrain, which, when acti-
vated, can engage evolutionarily conserved defensive
responses that function to resist or to avoid an
impending threat (e.g. fight, flight, faint; De Oca,
DeCola, Maren, & Fanselow, 1998; Brandão,
Zanoveli, Ruiz-Martinez, Oliveira, & Landeira-
Fernandez, 2008; Fenster, Lebois, Ressler, & Suh,
2018; for a review, see Keay & Bandler, 2014).
Terpou et al. (2019a, 2019b) recently investigated
the functional characteristics of the PAG during sub-
liminal, trauma-related stimulus conditions in parti-
cipants with PTSD as compared to healthy
individuals. Initially, Terpou et al. (2019b) assessed
subcortical activity using improved spatial normal-
ization procedures to attain greater resolution across
brainstem, midbrain, and cerebellar brain structures
(Diedrichsen, 2006). The approach revealed the PAG
to display stronger activity in participants with PTSD
as compared to healthy controls during subliminal,
trauma-related stimulus conditions (Terpou et al.,
2019a). These findings are in keeping with a putative
bias towards evolutionarily conserved defensive
responses in PTSD, which may be expressed more
actively in participants with PTSD than in healthy
controls (for a review, see Fragkaki, Thomaes, &
Sijbrandij, 2016; Kozlowska, Walker, McLean, &
Carrive, 2015; Terpou et al., 2019c).
In a subsequent paper, Terpou et al. (2019a)
sought to assess functional connectivity of the PAG
during subliminal, trauma-related stimulus condi-
tions in participants with PTSD as compared to
healthy individuals using the same participant sample
and paradigm as described in Terpou et al. (2019b).
Here, Terpou et al. (2019a) found significantly stron-
ger functional connectivity between the PAG and the
DMN in individuals with PTSD as compared to
healthy controls during subliminal, trauma-related
stimulus conditions. In particular, the DMN hubs
(i.e. posterior cingulate cortex, precuneus, medial
prefrontal cortex) were connected functionally to
the PAG, but only during subliminal, trauma-related
stimulus conditions and not during subliminal, neu-
tral stimulus conditions (Figure 1(b)). Terpou et al.
(2020) also evaluated the directed functional connec-
tivity, or effective connectivity, between the PAG and
the DMN to determine whether the PAG or the
DMN were driving patterns of functional connectivity
in participants with PTSD and healthy controls. In
PTSD as compared to controls, Terpou et al. found
the PAG showed stronger excitatory effective connec-
tivity to the DMN during subliminal, trauma-related
stimulus processing. Taken together, these findings
contribute to our understanding of why individuals
with PTSD may describe experientially links between
trauma- and self-related processing states.
Specifically, the PAG, which mediates innate, defen-
sive responses, demonstrates strong, directed func-
tional connectivity to the DMN responsible, in part,
for mediating higher-order, self-related processing;
these patterns are only present during subliminal,
trauma-related stimulus conditions but not during
rest more generally.
These findings shed light on the central role that
trauma plays in personal identity among participants
with PTSD – a relation that may be mediated, in part,
by these functional alterations. A stronger under-
standing of PTSD-related asymmetries in functional
connectivity across the DMN during rest, as well as
during subliminal, trauma-related stimulus condi-
tions may yield important clinical knowledge sur-
rounding the sense of self in the aftermath of
trauma. Going forward, such efforts will be necessary
to grasp more thoroughly the experience of indivi-
duals with PTSD who do describe an inextricable link
between trauma and the sense of self in the aftermath
of trauma (for a review, see Olff et al., 2019).
5. The DMN under threat: how might DMN-
related alterations present clinically?
Thus far, we have characterized the DMN-related
alterations observed in individuals with PTSD as
compared to healthy individuals, both during rest,
as well as during subliminal, trauma-related stimulus
conditions, where the former and the latter contexts
are associated with greatly different functional con-
nectivity patterns – that is, decreased and increased
DMN functional connectivity, respectively.
Additionally, we have reviewed an important func-
tion of the DMN in generating a perceived sense of
self, where an interplay across interacting DMN-
related processes associate to produce the sense of
self, which, critically, shows disturbances in PTSD.
For example, it is well known clinically that indivi-
duals with PTSD – particularly when associated with
early childhood maltreatment – report frequently a
rudimentary sense of self, or a sense of self that does
not exist entirely, illustrated eloquently through state-
ments, such as, ‘I do not know who I am,’ or, ‘I feel
like I have stopped existing.’ Our research, as well as
that of other research groups, suggests that these
experiences may relate, in part, to the reduced func-
tional connectivity observed during rest among indi-
viduals with PTSD, where these patterns contrast
strikingly with the enhanced DMN functional con-
nectivity observed under threat- or trauma-related
stimulus conditions in PTSD. Here, data pointing to
an increased DMN functional connectivity under
EUROPEAN JOURNAL OF PSYCHOTRAUMATOLOGY 5

conditions of threat (i.e. subliminal, trauma-related
processing) are just emerging. Given the multi-func-
tional properties of the DMN, we now offer several
interpretations on why the DMN may display recruit-
ment under conditions of threat from a clinical per-
spective. We hope the following discussion will serve
as a starting point for future investigation of the
DMN under threat in traumatized individuals.
Autobiographical memory represents an important
function mediated by the DMN, which may assist our
understanding on why the DMN demonstrates
increased functional connectivity in PTSD during sub-
liminal, trauma-related stimulus conditions.
Autobiographical memory helps us learn from previous
experiences (Qin & Northoff, 2011), where we can
evaluate past experiences in the present to guide actions
more adaptively. Autobiographical memory must then
intrinsically represent our past experiences, which, in
turn, would shape how we perceive the present – as the
present exists in constant relation to the past. Through
this lens, we can observe how detrimental childhood
maltreatment may be towards the development of the
DMN, which begins maturation during childhood and
does not end until early adulthood (Sherman et al.,
2014; for a review, see Fair et al., 2008). Where the
maltreatment and/or trauma(s) are repeated often, we
would suspect then that the developing DMN will be
biased towards threat- or trauma-related conditions.
This perspective may also hold for trauma experienced
as a singular event during childhood, but then re-
experienced, or relived internally whilst the trauma
remains unprocessed (van der Kolk, 2015). Although
the present perspective would be more difficult to apply
for trauma(s) experienced later in adulthood, recent
research suggests that individuals with combat-related
PTSD are more likely to report childhood adversity or
maltreatment as compared to combat-exposed indivi-
duals who do not go on to develop PTSD post-combat
(Blosnich, Dichter, Cerulli, Batten, & Bossarte, 2014;
Bremner, Southwick, Johnson, Yehuda, & Charney,
1993; Dannlowski et al., 2012). Here, DMN-related
alterations may be developed during childhood and, in
turn, act as a vulnerability factor towards the develop-
ment of PTSD post-combat. Accordingly, we present
the case where autobiographical memory-related pro-
cesses may promote biases towards trauma-related pro-
cessing among individuals with PTSD, which may assist
to explain the stronger DMN functional connectivity
during subliminal, trauma-related stimulus conditions
reported by Terpou et al. (2019a). Future research will
also need to determine whether such biases towards
trauma-related processing may provoke certain trau-
matic re-enactments, or automatic repetitions of the
past – a phenomenon observed frequently among indi-
viduals with PTSD. Here, van der Kolk (1989) has
suggested that ‘assaults lead to hyperarousal states for
which the memory can be state-dependent, or
dissociated, where the memory only returns fully dur-
ing renewed terror.’ It is therefore possible that some
individuals with PTSD under certain conditions may
seek situations involving threat or terror in order to
engage the DMN. This may in turn afford the experi-
ence of a semblance of a sense of self and a related sense
of agency, which may be lacking in the absence of
extreme hyperarousal states.
Additionally, the ability to mentalize the perspective
of others is another critical function mediated by the
DMN, which may serve a central role in survival, parti-
cularly during conditions of interpersonal violence,
where the awareness of the emotions and the malicious
intents of others may prove life-saving. Under such
conditions, a more intact DMN may serve as an indis-
pensable aid to survival by assisting a repeatedly trau-
matized individual to act and to make decisions in the
present, with consideration to the past and to the future,
as well as to facilitate an ability to assess the mental
states of others. This hypothesis awaits further exam-
ination, where, for example, relative to healthy controls,
women with a history of developmental trauma are
delayed in their response to sad, fearful, and happy,
but not angry (i.e. threatening) voices (Nazarov et al.,
2015), an effect that was enhanced also among survivors
with an increased severity to childhood abuse.
Interestingly, more severe symptoms of dissociation
were associated with reduced accuracy in discriminat-
ing between emotions in the same sample.
Another important question raised by the findings
of enhanced DMN connectivity under threat con-
cerns the possibility that individuals with PTSD may
be drawn towards fear- or terror-inducing conditions
in an attempt to heighten emotional experience and
to perceive at least the semblance of a sense of self.
Here, individuals who suffer from PTSD report fre-
quently that they do not ‘feel alive’ unless they engage
in sensory seeking or reckless behaviours. The follow-
ing quote from an individual with a history of sig-
nificant developmental trauma describes eloquently
how inducing fear and terror through engaging in
shoplifting helped to create the experience of a rudi-
mentary sense of self. She notes, ‘I started shoplifting
when I was five. I’d pretend to add the quarter my
mother gave me to the collection plate, then sink it
deep and hot into the pocket of my Sunday dress. On
the long walk home, I’d pass a pharmacy where I’d
steal a Clark Bar or a Milky Way, pantomime leaving
the quarter for my coke and with a mix of terror and
thrill leave the store, sugar happy and known to
myself … I shoplifted well into my adulthood, at
great risk to me were I to be caught … It was always
confusing why I did this. It was so, so risky. I knew
that. But, I think the adrenaline organized me, rising
it seemed from my belly through my brain, from the
back to the front. I felt my feet; I knew my hands and
fingers; I had eyes. I was agency. It lit me up. It was
6R. A. LANIUS ET AL.

essential. At five and still at fifty, I didn’t exist to
myself except as the artful dodger [pickpocket] – at
these moments, I existed; all of me, in the act of
stealing, I would “come online”.’
It is interesting to note that reckless behaviour is
exhibited by many individuals with PTSD and, accord-
ingly, has been included as a symptom of PTSD in the
most recent version of the DSM, the DSM-5 (APA,
2013). As described in the quote above, reckless beha-
viour may assist the traumatized individual not only to
feel more alive and embodied by helping to overcome
intense symptoms of emotional numbing, but may also
aid in bringing online a sense of agency that is lacking
sorely in the aftermath of their personal trauma. It will
therefore be critical for future research to examine the
relation between reckless behaviour, fear/terror, and the
integrity of the DMN. On balance, the findings
reviewed here point further towards the urgent need
to target therapeutically the sensory seeking and the
reckless behaviour that may be evoked as a last resort
to recruit the DMN, thus leading the participant with
PTSD to experience a veneer of a sense of self they may
have lost in the aftermath of trauma.
What are other implications for treatment that we
need to consider? Given the findings described above,
understanding the importance of clinical treatment to
the restoration of the self, particularly in the absence of
threat, will be of the utmost importance. Here, re-estab-
lishing, or establishing for the first time, a sense of self
that is continuous across time and into the future has
been a focus of treatment for trauma-related disorders
for decades (Allen, 1995; Herman, 1992; van der Kolk,
2015). More specifically, overcoming the fragmentary,
or the timeless nature of traumatic memories, increas-
ing emotional awareness, and helping individuals with
PTSD reclaim his/her body are critical components of
both present- and past-centred therapies (i.e. Cognitive
Processing Therapy (CPT); Resick and Schnicke (1992);
Prolonged Exposure Therapy; Foa, Hembree, and
Rothbaum (2007); Eye Movement Desensitization and
Reprocessing (EMDR); Shapiro (2018); Mindfulness
Therapy; Boyd, Lanius, and McKinnon (2018); Lanius,
Bluhm, and Frewen (2011); Frewen and Lanius (2015))
and mirror eloquently the functions of an intact DMN.
Moreover, several present- and past-centred therapies,
including mindfulness training (King et al., 2016), neu-
rofeedback (Kluetsch et al., 2014), EMDR, cognitive
behavioural therapy (CBT), and prolonged exposure
(for a review, see Malejko, Abler, Plener, & Straub,
2017) that are used to treat PTSD have shown a restora-
tion across the DMN while at rest.
Recently, psychotherapeutic interventions for indi-
viduals with PTSD have been combined with various
psychoactive substances, including the stimulant/psy-
chedelic hybrid 3,4-methylenedioxymethamphetamine
(MDMA) (Mithoefer, Grob, & Brewerton, 2016;
Mithoefer et al., 2018). Here, Carhart-Harris et al.
(2014) have shown MDMA to significantly activate
the DMN during the recall of favourite and worst auto-
biographical memories in healthy controls. Strikingly,
whereas positive memories were experienced as more
vivid and emotionally intense, negative memories were
experienced as less negative following administration of
MDMA versus a placebo in healthy individuals. Future
research examining the effects of MDMA in conjunc-
tion with psychotherapy in participants with PTSD will
therefore need to elucidate DMN functional connectiv-
ity and integrity and associated DMN-related functions
pre- and post-MDMA treatment.
In summary, we have attempted to provide a strong
case for the involvement of the DMN towards an altered
sense of self observed among individuals with PTSD,
where, critically, both the DMN, neurobiologically, and
the sense of self, clinically, demonstrate dramatic altera-
tions among participants with PTSD. Future research
examining treatment outcomes associated with trauma-
related disorders will need to incorporate not only
measures to assess the continuous sense of self into
the future, but also an assessment of the distributed
functions related to the DMN, including self-referential
and autobiographical memory processes, theory of
mind, and embodiment, as well as to investigate speci-
fically how these processes may be influenced by pre-
sent- and past-centred therapies. It will also be critical
to examine DMN integrity and its related functions in
interpersonal versus non-interpersonal trauma, as well
as the vulnerability factors to the development of PTSD.
Moreover, assessing traumatic re-enactments and reck-
less behaviour and their potential relation with the
DMN will be crucial to further our understanding of
trauma-related disorders. Only by targeting specifically
critical DMN dysfunctions in the aftermath of trauma
will we be able to assist further in the restoration of the
self in PTSD, allowing these individuals to reclaim a
sense of self that had previously been lost.
Disclosure statement
No potential conflict of interest was reported by the
authors.
Funding
RAL and MCM are co-funded by the Canadian Institutes
for Health Research (Grant Numbers: 171647). RAL is
funded by the Canadian Institute for Military and
Veteran Health Research (Grant Number: W7714-145967/
001/SV T27).
ORCID
Ruth A. Lanius http://orcid.org/0000-0002-3758-1393
Braeden A. Terpou http://orcid.org/0000-0002-6770-
2715
EUROPEAN JOURNAL OF PSYCHOTRAUMATOLOGY 7

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