Available online at www.sciencedirect.
com
ScienceDirect
The innate alarm system in PTSD: conscious and subconscious
processing of threat
Ruth A Lanius1,2,3,4,9, Daniela Rabellino2,9, Jenna E Boyd4,5,6,
Sherain Harricharan1, Paul A Frewen2,7 and Margaret C McKinnon4,5,8
The innate alarm system (IAS), comprised of functionally
connected brain regions including the brainstem, amygdala,
pulvinar, and frontotemporal cortex, is a fast subcortical brain
network facilitating rapid responses to threat. Post-traumatic
stress disorder (PTSD) features subconscious and conscious
threat detection, together contributing to hyperarousal
symptoms. Emerging literature identifies aberrant threat-related
neurocircuitry involved in subconscious and conscious threat
processing in PTSD. We review this literature, focusing on
subconscious threat processing and its relation to the IAS.
Available evidence indicates increased neural activity and
functional connectivity between IAS brain regions (e.g. locus
coeruleus, superior colliculus, amygdala, and prefrontal cortex).
These alterations are observed during both subconscious threat
processing and at rest, suggesting increased defensive
posturing, maintained in the absence of overt threat.
Addresses
1
Department of Neuroscience, University of Western Ontario,
1151 Richmond St, London, ON N6A 3K7, Canada
2
Department of Psychiatry, University of Western Ontario,
1151 Richmond St, London, ON N6A 3K7, Canada
3
Imaging Division, Lawson Health Research Institute, 750 Base Line
Road East, London, ON N6C 2R5, Canada
4
Homewood Research Institute, 150 Delhi St, Guelph, ON N1E 6K9,
Canada
5
Mood Disorders Program, St. Joseph’s Healthcare Hamilton, 100 West
5th St., Hamilton, ON L8S 4K1, Canada
6
Department of Psychology, Neuroscience, and Behaviour, McMaster
University, 1280 Main Street West, Hamilton, ON L8S 4K1, Canada
7
Department of Psychology, University of Western Ontario,
1151 Richmond St, London, ON N6A 3K7, Canada
8
Department of Psychiatry and Behavioural Neurosciences, McMaster
University, 100 West 5th, Hamilton, ON L8N 3K7, Canada
Corresponding author: Lanius, Ruth A (ruth.lanius@lhsc.on.ca)
9
Shared first authorship.
Current Opinion in Psychology 2017, 14:109–115
This review comes from a themed issue on Traumatic stress
Edited by Anka A. Vujanovic and Paula P. Schnurr
For a complete overview see the Issue and the Editorial
Available online 26th November 2016
http://dx.doi.org/10.1016/j.copsyc.2016.11.006
2352-250X/# 2016 Elsevier Ltd. All rights reserved.
Introduction
Threat-related stimuli present in the natural environment
can be detected consciously and subconsciously (i.e. below
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the threshold of consciousness), where subconscious de-
tection of threatening stimuli confers an evolutionary
advantage, promoting the initiation of fight-or-flight re-
sponse in the absence of conscious awareness [1–3]. This
subconscious detection is mediated by the ‘innate alarm
system’ (IAS), a network of inter-connected brain regions
including prominently the brainstem, amygdala, pulvinar,
and fronto-temporal cortex [3–6]. Together, these brain
regions facilitate a ‘fast-track’ response at the level of the
central nervous system to threatening stimuli [3].
Both subconscious rapid defensive responses and con-
scious defensive responses feature in the clinical picture
of post-traumatic stress disorder (PTSD). Whereas trig-
gers processed at the conscious level elicit a recognizable
defensive response pattern (e.g. active avoidance, hyper-
vigilance), subconsciously processed triggers elicit a range
of symptoms that, to date, have been poorly character-
ized. Critically, while both subconscious and conscious
triggers may elicit similar defensive responses (e.g. hy-
pervigilance), subconscious triggers may be more difficult
to target clinically as they occur outside the conscious
awareness of patients.
Here, we review emerging evidence of altered threat-
related neurocircuitry among individuals with PTSD
at the conscious and subconscious processing level,
with a focus on subconscious processing. We also re-
view seminal work highlighting the existence of the
IAS in non-psychiatric populations. Finally, we explore
the implications of these findings for future research
and clinical practice.
Neural correlates of conscious and
subconscious threat processing
Subconscious processing of threat-related stimuli in
non-psychiatric populations
A subcortical route for fear processing in response to
subconscious fear stimuli has been characterized among
healthy individuals [3–6,16]. Here, Liddell and collea-
gues published pivotal work, describing activation of the
locus coeruleus (LC), superior colliculus (SC), and pulvi-
nar, as well as the amygdala and prefrontal regions, in
response to subconscious fear stimuli among healthy
controls [3]. This mechanism is thought to represent an
evolutionarily adaptive response, facilitating rapid activa-
tion of alerting mechanisms and defensive responses prior
to conscious awareness [17], and thus increasing the
probability of survival [5,18,19].
Current Opinion in Psychology 2017, 14:109–115
110 Traumatic stress
Recent work using intracranial electrophysiological meth-
ods among epilepsy patients identified rapid activation
of the amygdala preceding activation of the visual cortex
in response to fearful, but not neutral or happy low-
frequency faces [20]. Similarly, in healthy individuals,
a subcortical amygdala–pulvinar pathway involved in
early (but not late) visual processing of fearful and happy
or neutral faces has been identified when measured by
magnetoencephalography [21]. These studies support
the notion of a fast, subcortical fear pathway, allowing
rapid responding to (or evaluation of) threat, prior to
conscious awareness (e.g. processing via the visual cor-
tex). It remains to be determined whether differences in
emotional valence (e.g. fearful vs happy) differentially
affect rapid visual processing.
Conscious processing of threat-related stimuli in PTSD
Our understanding of PTSD at the neurobiological level
relies largely on observations of conscious threat proces-
sing, where during experimental inductions, individuals
with PTSD who present primarily with symptoms of
hyper-reactivity (e.g. hypervigilance) demonstrate de-
creased neural activity in prefrontal regions [e.g. the
medial prefrontal cortex (mPFC)] and concomitant in-
creased activity in regions including the perigenual ante-
rior cingulate cortex (pgACC), amygdala, hippocampus,
and posterior regions involved in memory recall (i.e. mid-
line retrosplenial cortex, precuneus) [7–11]. These find-
ings are consistent with a neurocircuitry model of PTSD
suggesting reduced top-down inhibition of limbic regions
[7,12]. By contrast, patients exhibiting prominent symp-
toms of depersonalization and derealization (as outlined
in the DSM-5 dissociative subtype of PTSD
(PTSD DS) [13]) in response to threatening stimuli
exhibit increased neural activity in prefrontal regions [e.g.
mPFC, dorsal anterior cingulate cortex (dACC)], accom-
panied by reduced activity in limbic regions (e.g. amyg-
dala, anterior insula (AI)) [14,15].
Subconscious processing of threat-related stimuli in
PTSD
The innate alarm system (IAS) is identified as a key
contributor to the development and maintenance of
PTSD symptomatology. Increased activity in areas asso-
ciated with the IAS is reported during subconscious
processing of threat stimuli, including in brainstem
regions [22], the amygdala [22–24], mPFC [22,23], para-
hippocampus [25], and visual cortex [26]. Together, these
findings suggest hyperreactivity of the IAS to fear or
trauma-related stimuli in PTSD. A critical early study
by Bryant and colleagues [23] reported increased amyg-
dalar-mPFC connectivity during subconscious fear pro-
cessing among individuals with PTSD when compared to
controls. Similarly, a recent study described hypersyn-
chrony within fear circuit brain regions when implicitly
(subconsciously) viewing fearful faces, including
the mPFC and amygdala, among individuals with
Current Opinion in Psychology 2017, 14:109–115
combat-related PTSD as compared to combat-exposed
controls [27].
Differential responding to subconscious threat stimuli in
PTSD (PTSD DS) and its dissociative subtype
(PTSD + DS) has also been described. Specifically, as
compared to patients with PTSD DS, patients with
PTSD + DS showed enhanced amygdala and parahippo-
campal activity during non-conscious fear processing [15].
These findings suggest enhanced activity of the IAS in
PTSD + DS thought to preceed prefrontal dampening of
limbic regions, as observed during conscious fear proces-
sing in PTSD + DS.
In addition to key findings of aberrant connectivity
between the amygdala and prefrontal regions in PTSD
[28–30], a recent study identified increased connectivity
between the basolateral amygdala (BLA) and prefrontal
cortical regions in individuals with PTSD compared to
controls at rest [31]. This finding suggests strongly hyper-
connectivity within the IAS during rest. Work from our
laboratory mirrors these findings, where increased resting
state functional connectivity of the BLA and the centro-
medial amygdala (CMA) with medial prefrontal region
was found in PTSD + DS as compared to PTSD DS
[32]. Increased connectivity between the BLA and cere-
bellar culmen was also found [32], where the cerebellum
has been conceptualized recently as central to a cerebel-
lar-limbic-thalamo-cortical network involved in rapid
responses to threat and in the fight-or-flight response [33].
The insula is also associated with the IAS. This region is
thought to play a key role in response to subconscious
stimuli by monitoring homeostasis of the internal milieu
[3,34]. For example, individuals with PTSD DS and
PTSD + DS showed increased connectivity of insula sub-
regions with the BLA at rest [35]. This pattern of activa-
tion was most prominent among the PTSD + DS group,
and converges with previous findings of heightened
amygdala-insular connectivity in PTSD [36,37]. In an-
other recent study, individuals with PTSD DS and
PTSD + DS demonstrated dorsolateral periaqueductal
gray (PAG) connectivity with IAS brain regions, including
the dorsal ACC and insula [38]. Critically, the PAG is
thought to mediate, in part, motor patterns of fight or
flight response [39]. Moreover, both the dorsolateral and
ventrolateral PAG showed connectivity with the fusiform
gyrus, an area involved in detection of faces, voices, and
movement to determine their environmental threat [17].
These findings again point strongly to hyperconnectivity
within the IAS that is associated with the maintenance
of defensive posturing, even at rest, particularly in
PTSD + DS.
Recent work has suggested further hyperconnectivity of
amygdalar subregions within the IAS during subconscious
threat processing. Here, increased prefrontal (e.g. medial
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superior frontal gyrus (SFG)) connectivity with the right
CMA was reported among PTSD patients [40]. Interest-
ingly, decreased connectivity of the BLA with the SC was
also noted. This pattern of aberrant responding may
contribute to reduced integration of sensory information
from the SC to the BLA, and thus reduced emotional
evaluation of incoming information [40]. Interestingly,
increased connectivity between the LC (involved in
arousal modulation [41] and in the IAS [3,42]) and the
left BLA was also noted during conscious processing,
suggesting further heightened functional connectivity
within the IAS during conscious threat processing [40].
Notably, increased right cerebellar lobule VI and posteri-
or cingulate cortex activation among individuals with
PTSD has been found during subconscious processing
of trauma and threat-related stimuli when compared to
controls [43]. Increased cerebellar activation may indi-
cate an attempt to integrate subconscious threat cues into
conscious experience [43]. Here, re-experiencing and
hyperarousal symptoms were associated with hyperactiva-
tion in key components of the IAS, including the amyg-
dala during subconscious processing and the PAG and SC
during conscious processing [43]. Conversely, avoid-
ance symptoms were associated with increased activation
in regions associated with emotional down-regulation
(e.g. the inferior frontal gyrus [44,45]), consistent with
emotional overmodulation observed in PTSD + DS.
Moreover, reduced activation in cerebellar regions (left
lobule IV/V) during subconscious fear processing was also
observed [43]. Given the role of cerebellar lobule V in
sensorimotor perception [46], it is probable that this
region contributes to sensorimotor integration during
subconscious processing, a process likely disrupted in
PTSD [43].
Finally, direct eye contact may be conceived as approxi-
mating subconscious threat processing in PTSD, given
that in healthy individuals, direct eye contact is first
detected implicitly by areas involved in the IAS, such
as the SC, PAG, pulvinar, and amygdala [47,48]. Further,
direct eye contact is purported to be involved in the
detection of threat signals [49], before engaging higher
cortical areas involved in social cognition. Functional
neuroimaging studies of fearful vs happy or neutral facial
expressions in PTSD have identified hypoactivation in
prefrontal regions associated with the IAS (e.g. mPFC)
[10,50] and increased amygdalar reactivity [10,24,50] in
response to masked [24] and overt facial stimuli [10,50].
Importantly, as compared to controls, individuals with
PTSD related to interpersonal trauma show altered reac-
tivity within the IAS, including increased activity in the
SC, PAG, and LC accompanied by decreased activity in
cortical regions (e.g. dorsomedial prefrontal cortex, tem-
peroparietal junction) associated with social cognition, in
response to direct vs averted eye gaze [51]. A follow-up
study revealed increased connectivity between the SC
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The innate alarm system in PTSD Lanius et al. 111
and LC with IAS regions, including the insula, inferior
frontal cortex, amygdala, ACC, and striatum [52], findings
thought related to sustained activation of the subcortical
pathway of eye contact processing. These findings sug-
gest that direct eye contact may be perceived subcon-
sciously as threat by individuals with PTSD related
to interpersonal trauma, where these patients show
prominent activation of neural regions within the IAS
rather than higher cortical regions involved in social
cognition.
Clinical and research implications
Our understanding of the neurobiology underlying sub-
conscious processing of threat-related cues among indi-
viduals with PTSD remains in a nascent stage. Alterations
in threat reactivity are prominent in PTSD; thus, it will be
critical to increase this understanding. Existing knowl-
edge surrounding the broader neurobiological underpin-
nings of PTSD may be leveraged to design and test
neuroimaging and behavioural paradigms aimed specifi-
cally at identifying the neurocircuitry and associated
patterns of behaviour involved in subconscious threat
processing in PTSD (see Figure 1).
The emerging literature suggests substantial alterations
in threat-related neurocircuitry among individuals with
PTSD. Earlier work identified alterations in the IAS,
particularly among brainstem regions, the amygdala, para-
hippocampus, and mPFC, during subconscious threat
processing [22–26]. In keeping with these findings, recent
studies point further towards increased resting state con-
nectivity between IAS regions (e.g. amygdala, prefrontal
regions and insula) [31,32,35], a pattern indicating defen-
sive posturing even at rest that may be particularly
prominent in PTSD + DS [32,35]. This pattern of re-
sponse is maintained during subconscious threat proces-
sing, where increased connectivity between amygdalar
regions and other regions associated with the IAS (e.g. SC,
SFG) has been reported [40]. Moreover, during direct eye
gaze, as compared to controls [51], women with PTSD
related to interpersonal trauma show heightened activa-
tion within and increased connectivity between IAS
regions (e.g. SC and LC with amygdala, frontal cortex,
ACC) [52]. Here, IAS may be associated with aberrant
social responding observed in this patient population
[51,52].
In summary, an emerging literature contributes to our
understanding of the IAS among individuals with PTSD.
Perhaps most strikingly, the evidence reviewed here
highlights the urgent need to look beyond the amygdala
to regions such as brainstem and midbrain structures (e.g.
the LC, PAG, SC), and cerebellum in order to better
understand threat reactivity in this population (see
Figure 1). Future work should aim to increase our under-
standing of the time course of threat reactivity in individ-
uals with PTSD, where findings of increased connectivity
Current Opinion in Psychology 2017, 14:109–115
112 Traumatic stress

Figure 1

Sensory Input

Prefrontal
Cortex/ACC

Pulv

SC
Cerebellum
PAG

Amg
LC

Imminent
Threat

Visceral Organs

Current Opinion in Psychology

The innate alarm system. The ‘innate’ alarm system is an evolutionarily conserved neural circuit that relies on lower-order brain structures to help
facilitate an immediate response to a perceived threat rather than engagement of somatosensory and prefrontal cortices to appraise the source of
threat and develop an appropriate response. Mobbs et al. Science 2007 317:1079–1083 suggest that when a distal threat is detected, the
prefrontal cortices facilitate an appropriate escape plan. However, as the threat advances closer, the role of the midbrain and limbic-driven ‘innate
alarm system’ is enhanced. The superior colliculus (SC) is critical for receiving visual, auditory, and somatosensory input, and helps to orient one’s
own body in response to sensory stimuli. The SC works in tandem with the periaqueductal gray (PAG) to help process external stressors at the
midbrain level. Eventually, the SC/PAG provides input to the pulvinar nuclei of the thalamus (Pulv) about the source of threat, which in turn
communicates with the amygdala to help drive the immediate response to threat. Although the amygdala communicates threat information to the
prefrontal cortex, it is nevertheless able to initiate rapid responses to threat independently. The amygdala innervates the locus coeruleus (LC),
another midbrain structure responsible for carrying out physiological responses to threat, and the periaqueductal gray (PAG), where both of these
structures work together to elicit ‘fight-or-flight’ responses to threat. In doing so, the LC eventually projects to the visceral organs to facilitate
sympathetic activation of these organs. The IAS converges with several other neural models proposed to explain physiological and behavioural
responses to conscious and subconscious threat, including Porges Cleve Clin J Med 2009 76:S86–S90 model of ‘neuroception’ as well as
Schutter and colleagues cerebello-limbic-thalamo-cortical network, which we would direct the reader to review in the literature. In the latter model,
the cerebellum plays a homeostatic role in the ‘innate alarm system’ where it is in constant communication with midbrain structures and is thought
important for processing rapid responses to threat. In the figure, red arrows represent unidirectional drives, whereas black arrows depict
bidirectional interactions. Vector clipart images were retrieved and adapted from a free public domain (clker.com, Rolera LLC).

within the IAS during conscious threat processing
[40,43] suggest this system may be maintained (i.e.
continue to be active) during proceeding conscious pro-
cessing in this population. Future work should also aim to
elucidate brain regions involved in the transition from
subconscious to conscious threat processing in PTSD to
better distinguish these two forms of threat reactivity.
This will be critical to our understanding of how over-
lapping brain regions involved in both subconscious
and conscious threat processing (e.g. amygdala,
frontotemporal cortex) are involved differentially in both
systems. Further, it will be crucial to examine the beha-
vioural correlates of subconscious and conscious proces-
sing in order to better delineate these two states among
individuals with PTSD. It may be that subconscious fear
processing relying on subcortical circuits may lead to
behavioural reaction (e.g. fight or flight), but not emotional
reactions (e.g. subjective feelings of fear), which rely on

Current Opinion in Psychology 2017, 14:109–115 www.sciencedirect.com


cortical processing as in LeDoux and Pine’s two-system
framework [53]. In addition, researchers should aim to
understand further differences in subconscious threat
processing between individuals with PTSD DS and
PTSD + DS, given that these patient groups display
distinct patterns of neural activation during conscious
threat processing and that individuals with PTSD + DS
often display profound detachment and lack of awareness
of the internal and external environments [14], which may
lead to inappropriate responses to threat stimuli. Critical-
ly, given the nascent stage of this subconscious threat-
processing research in PTSD, the findings reported here
must be replicated in future studies. Clinically, these
findings highlight the need to assist patients in gaining
awareness of physiological, behavioural, and psychologi-
cal responses to traumatic stimuli that may be perceived
below the level of consciousness.
Funding
This work was supported by the Canadian Institute for
Military and Veteran Health Research, and the Canadian
Institutes of Health Research. Daniela Rabellino was
supported by fellowship from MITACS and Homewood
Research Institute.
Conflict of interest statement
Nothing declared.
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Current Opinion in Psychology 2017, 14:109–115