The Benzodiazepine Alprazolam Dissociates
Contextual Fear from Cued Fear in Humans as
Assessed by Fear-potentiated Startle
Christian Grillon, Johanna M.P. Baas, Daniel S. Pine, Shmuel Lissek, Megan Lawley, Valerie Ellis, and
Jessica Levine
Background: The startle reflex is potentiated by aversive states. It has been proposed that phasic startle potentiation to a threat cue
and sustained startle potentiation to contextual stimuli reflect distinct processes mediated by different brain structures. The present
study tested the hypothesis that alprazolam would reduce the sustained startle potentiation to contextual threats but not the startle
potentiation to a threat cue.
Methods: Sixteen healthy subjects received each of four treatments: placebo, .5 mg of alprazolam, 1 mg of alprazolam, and 50 mg
of diphenhydramine (Benadryl) in a crossover design. Participants were exposed to three conditions, including one in which
predictable aversive shocks were signaled by a cue, a second in which shocks were administered unpredictably, and a third condition
in which no shocks were anticipated. Acoustic startle were delivered regularly across conditions.
Results: Phasic startle potentiation to the threat cue in the predictable condition was not affected by alprazolam. In contrast, the
sustained increase in startle in the predictable and unpredictable conditions was reduced significantly by the high dose of alprazolam.
Conclusions: Startle responses to an explicit threat cue and to an aversive context are psychopharmacologically distinct, suggesting
that they may represent functionally dissociable aversive states.
Key Words: Alprazolam, anxiety, benzodiazepine, context, fear,
startle reflex
T
he startle reflex, a cross-species response to a sudden
intense stimulus, is sensitive to aversive states (Davis et al
1993). Rodents show robust startle potentiation to a
conditioned cue that signals an aversive event (e.g., a shock).
Evidence suggests that this effect is mediated by the central
nucleus of the amygdala (CeA). For example, lesions of the CeA
block fear-potentiated startle to a conditioned cue previously
paired with shock (Davis 1998). Although the CeA responds to
various types of stressors, this structure is not always critical for
fear-potentiated startle (Davis 1998) or for responses to stress in
general (Hammack et al 2004). Davis and his collaborators have
reported a series of studies showing no effect of lesions of the
CeA on startle potentiation caused by various stressors, such as
bright lights (i.e., light-enhanced startle), shock sensitization, and
corticotropin-releasing hormone injection (reviewed in Walker et
al 2003). Rather, another structure, the bed nucleus of the stria
terminalis (BNST), was found to mediate startle potentiation in
these conditions (Walker et al 2003). In an analysis of the
experimental situations that do or do not require the CeA and
BNST, Walker et al (2003) suggested that these structures were
involved in functionally different aversive states in rodents. It was
proposed that the CeA was crucial for the phasic form of
fear-potentiated startle to a predictable threat cue, whereas the
BNST was responsible for the more sustained form of startle
potentiation induced by unpredictable or unconditioned aver-
sive stimuli (Davis 1998; Gewirtz et al 1998; Walker and Davis
1997a). It was suggested further that the two aversive states
mediated by the CeA and the BNST in rodents were reminiscent
of fear and anxiety states in humans, respectively (Davis 1998).
According to this view, fear is a response to a clearly identifiable
danger that subsides shortly after the offset of a threat cue.
Anxiety is a more sustained form of general distress and anxious
apprehension in response to less identifiable cues (Barlow 2000;
Davis 1998; Lang et al 2000).
The distinction between a phasic and a more sustained form
of startle potentiation also has been made in humans (Cuthbert et
al 2003; Grillon et al 1991, 1997; Grillon and Davis 1997; Lang et
al 2000; Pole et al 2003). Startle is potentiated by an explicit threat
cue that signals an impending aversive event (e.g., a shock;
Grillon et al 1993; Hamm and Vaitl 1996), such as when phobic
individuals are confronted with their phobic objects (de Jong et
al 1996; Globisch et al 1999). More sustained forms of startle
potentiation can be found among individuals who are exposed
to stressful experimental settings (Bocker et al 2001; Grillon and
Ameli 1998; Pole et al 2003), when the experimental room is in
complete darkness (Grillon et al 1997), or after context condi-
tioning (Grillon and Davis 1997).
Consistent with the animal literature, phasic and sustained
forms of startle potentiation in humans appear to reflect distinct
processes. For example, individuals with posttraumatic stress
disorder (PTSD) or with panic disorder display normal startle to
explicit threat cues that signal a shock but display enhanced
startle reactivity in the experimental context in which the shocks
are administered (Grillon et al 1994, 1998b; Pole et al 2003).
Patients with PTSD also exhibit increased context conditioning
(Grillon and Morgan 1999) and increased facilitation of startle in
the dark (Grillon et al 1998a).
There currently is little information on the neurobiological
mechanisms that may differentiate phasic cued fear from more
sustained contextual anxiety in humans. Evidence for such a
neurobiological dissociation would be bolstered if one could
demonstrate that these two forms of aversive states are differen-
tially responsive to psychopharmacologic treatments. The main
objective of this study was to obtain such evidence by using the
benzodiazepine alprazolam.
From the Mood and Anxiety Disorder Program, National Institutes of Mental
Health, Bethesda, Maryland.
Address reprint requests to Christian Grillon, Ph.D., National Institutes of
Mental Health–Mood and Anxiety Disorder Program, 15K North Drive,
Building 15K, Room 113, MSC 2670, Bethesda, MD 20892-2670; E-mail:
christian.grillon@nih.gov.
Received July 12, 2005; revised October 27, 2005; accepted November 11,
2006.
BIOL PSYCHIATRY 2006;60:760 –766 0006-3223/06/$32.00
doi:10.1016/j.biopsych.2005.11.027 © 2006 Society of Biological Psychiatry