Activating memories of depression alters the experience of voluntary action Keywords Agency ·...

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1 3 Exp Brain Res (2013) 229:497–506 DOI 10.1007/s00221-012-3372-5 VOLITION Activating memories of depression alters the experience of voluntary action Sukhvinder S. Obhi · Kristina M. Swiderski · Riley Farquhar Received: 17 September 2012 / Accepted: 5 December 2012 / Published online: 18 December 2012 © Springer-Verlag Berlin Heidelberg 2012 Keywords Agency · Depression · Volitional action · Sense of agency · Authorship over action · Priming · Interval estimation Introduction The experimental psychology of action has mostly con- cerned itself with the processes that produce and control voluntary movements (Passingham 1993; Rosenbaum 1991; Haggard 2001; Obhi and Haggard 2004; Obhi and Goodale 2005; Kalaska 2009). In contrast, the experience of vol- untary action has received less research attention (but see Haggard 2008). The sense of agency is one of the most important and pervasive feelings surrounding voluntary action (Strother et al. 2010; Gallagher 2000). The feeling of control over the actions we make, and the effects they produce, is crucial to our sense of ourselves as purposeful organisms (Obhi and Haggard 2005). Despite this impor- tance, humans do not routinely dwell on this experience; instead, the sense of agency seems to be somehow woven into the processes that produce purposeful behavior and is “just there.” For example, if I realize that it is getting dark outside and I flick the light switch, and the light turns on, although I fully understand, and at some level feel, that my action caused the light to come on, I do not stop to ponder my role as the cause of the illumination (Obhi and Hall 2011a). In cases where something unexpected happens, such as, the light fails to come on, or the light flickers and then goes out, I might pause to consider the potential causes. Nevertheless, routine agency experience, where our actions produce effects that they are “meant” to, is something we do not often reflect upon. In this way, normal agency experience can be conceived as low level and pre-reflective (Synofzik et al. 2008; David et al. 2008). Abstract The sense of agency is a profoundly impor- tant human experience and is strongly linked to volitional action. The importance of this experience is underscored by the fact that many neurological and psychiatric disorders are partially characterized by an abnormal sense of agency (e.g., schizophrenia, anxiety disorders, depression). Healthy participants perceive the temporal interval between a vol- untary action and its effect to be shorter than it actually is, and this illusion has been suggested as an implicit index of agency. Here, we investigated whether activating memories of depression alters perception of this action–effect interval, compared to activating memories of the previous day, or a baseline condition in which specific memories were not acti- vated. Results showed that action–effect interval estimates were significantly longer after remembering a depressing episode than after remembering the previous day, or in the baseline condition. Thus, activating memories of depression alters the experience of voluntary actions and effects. We suggest that interval estimation measures could be useful in clinical settings, to implicitly assess the sense of agency in patients with disorders affecting their sense of control. In this way, obtaining action–effect interval estimates, pre-, during, and post-treatment, could aid in tracking treatment- induced changes in the sense of agency. S. S. Obhi (*) · K. M. Swiderski · R. Farquhar Centre for Cognitive Neuroscience, Wilfrid Laurier University, Waterloo, ON, Canada e-mail: [email protected] S. S. Obhi · K. M. Swiderski · R. Farquhar Department of Psychology, Wilfrid Laurier University, Waterloo, ON, Canada

Transcript of Activating memories of depression alters the experience of voluntary action Keywords Agency ·...

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Exp Brain Res (2013) 229:497–506DOI 10.1007/s00221-012-3372-5

VOLITION

Activating memories of depression alters the experience of voluntary action

Sukhvinder S. Obhi · Kristina M. Swiderski · Riley Farquhar

Received: 17 September 2012 / Accepted: 5 December 2012 / Published online: 18 December 2012 © Springer-Verlag Berlin Heidelberg 2012

Keywords Agency · Depression · Volitional action · Sense of agency · Authorship over action · Priming · Interval estimation

Introduction

The experimental psychology of action has mostly con-cerned itself with the processes that produce and control voluntary movements (Passingham 1993; Rosenbaum 1991; Haggard 2001; Obhi and Haggard 2004; Obhi and Goodale 2005; Kalaska 2009). In contrast, the experience of vol-untary action has received less research attention (but see Haggard 2008). The sense of agency is one of the most important and pervasive feelings surrounding voluntary action (Strother et al. 2010; Gallagher 2000). The feeling of control over the actions we make, and the effects they produce, is crucial to our sense of ourselves as purposeful organisms (Obhi and Haggard 2005). Despite this impor-tance, humans do not routinely dwell on this experience; instead, the sense of agency seems to be somehow woven into the processes that produce purposeful behavior and is “just there.” For example, if I realize that it is getting dark outside and I flick the light switch, and the light turns on, although I fully understand, and at some level feel, that my action caused the light to come on, I do not stop to ponder my role as the cause of the illumination (Obhi and Hall 2011a). In cases where something unexpected happens, such as, the light fails to come on, or the light flickers and then goes out, I might pause to consider the potential causes. Nevertheless, routine agency experience, where our actions produce effects that they are “meant” to, is something we do not often reflect upon. In this way, normal agency experience can be conceived as low level and pre-reflective (Synofzik et al. 2008; David et al. 2008).

Abstract The sense of agency is a profoundly impor-tant human experience and is strongly linked to volitional action. The importance of this experience is underscored by the fact that many neurological and psychiatric disorders are partially characterized by an abnormal sense of agency (e.g., schizophrenia, anxiety disorders, depression). Healthy participants perceive the temporal interval between a vol-untary action and its effect to be shorter than it actually is, and this illusion has been suggested as an implicit index of agency. Here, we investigated whether activating memories of depression alters perception of this action–effect interval, compared to activating memories of the previous day, or a baseline condition in which specific memories were not acti-vated. Results showed that action–effect interval estimates were significantly longer after remembering a depressing episode than after remembering the previous day, or in the baseline condition. Thus, activating memories of depression alters the experience of voluntary actions and effects. We suggest that interval estimation measures could be useful in clinical settings, to implicitly assess the sense of agency in patients with disorders affecting their sense of control. In this way, obtaining action–effect interval estimates, pre-, during, and post-treatment, could aid in tracking treatment-induced changes in the sense of agency.

S. S. Obhi (*) · K. M. Swiderski · R. Farquhar Centre for Cognitive Neuroscience, Wilfrid Laurier University, Waterloo, ON, Canadae-mail: [email protected]

S. S. Obhi · K. M. Swiderski · R. Farquhar Department of Psychology, Wilfrid Laurier University, Waterloo, ON, Canada

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Understanding how the sense of agency arises is a key challenge for the psychological and brain sciences (Obhi and Hall 2011a, b), not least because a loss of the normal sense of agency is a product of various psychiatric and neu-rological disorders. For example, individuals with schizo-phrenia experience delusions of control (Moore et al. 2012). Thus, they often feel like the author of actions and events that they are not actually responsible for, and conversely, sometimes feel no control over actions they do in fact make (Voss et al. 2010; Moore and Obhi 2012).

Studying the sense of agency in the laboratory is often done by asking participants to make movements that pro-duce effects and then asking them to report the degree of control or responsibility they feel for producing the outcome (e.g., Preston and Newport 2010; Yomogida et al. 2010). This self-report approach is useful and has led to insights into the factors that affect the conscious sense of agency (Nahab et al. 2011). However, this self-report approach is limited because it accesses the conscious, reflective sense of agency (Synofzik et al. 2008; David et al. 2008; Moore and Obhi 2012). Given that everyday agency experience is often not accompanied by conscious reflection, it is hard to relate research findings to the type of agency experiences humans have much of the time. In addition, and importantly, the limits of self-report are well known, in terms of the effects of demand characteristics and the use of explicit strategies (Podsakoff and Organ 1986; McCroskey 1984).

Over the last decade, there has been increasing inter-est in a temporal illusion that surrounds voluntary action, and more specifically, the notion that it could serve as an implicit index of the sense of agency (Haggard et al. 2002; Moore and Obhi 2012). Briefly, when an individual makes an action that is followed by an effect such as an auditory or visual event, they perceive the time interval between their action and the onset of the effect to be shorter than it actu-ally is (Haggard et al. 1999, 2002). This perceived compres-sion of the time interval between action and effect is only present for voluntary actions and does not typically mani-fest for passive actions, or movements that are the result of transcranial magnetic stimulation applied over the motor cortex (Tsakiris and Haggard 2003, but see also Strother et al. 2010). Owing to this apparent dependence on voli-tion, the perceived compression of the action–effect interval has been termed the “intentional binding effect” (see Moore and Obhi 2012 for a full discussion of intentional bind-ing). More generally, changes in the perceived duration of action–effect intervals may correspond to varying degrees of agency experience.

If perceptual estimates of action–effect intervals are indeed related to agency, it should be possible to demonstrate that judged intervals are affected in conditions where a per-son’s sense of control is pathologically abnormal, or experi-mentally perturbed. Some evidence supporting pathological

effects on interval estimates has been provided by studies using schizophrenic patients as experimental participants, although the exact relationship between interval estimates and explicit experiences of agency in these patients remains to be explained (Voss et al. 2010; Hauser et al. 2011; Moore and Obhi 2012).

We recently investigated whether experimentally induced feelings of social power would affect the perceived interval between a voluntary action and a visual effect. We used an established episodic priming technique to prime healthy par-ticipants to feel powerful, or relatively powerless (Galinsky et al. 2003; Obhi et al. 2012). We found that, compared to a baseline, no-priming condition, interval estimates were indeed longer when participants felt powerless. Based on this result, we speculated that measures of interval estima-tion might be a useful tool in clinical settings, for use with patients who suffer from conditions affecting their sense of agency. Specifically, we suggested that interval estimates might be useful for implicitly assessing the sense of agency before, during, and after a treatment intervention, with the prediction being that binding would increase (i.e., interval estimates would decrease) after a successful intervention.

Two patient groups who commonly report reduced feel-ings of control are individuals with anxiety disorders and individuals with depression (Keeton et al. 2008). Depres-sion in particular has long been associated with feelings of loss of control, or helplessness. As has been highlighted, perhaps most widely by Seligman et al. over the years, the conditions for depression can ensue when individu-als feel a chronic lack of control over outcomes in their lives (Peterson and Seligman 1984; Seligman 1989). In his theory of helplessness and depression, Seligman sug-gests that, when people perceive no control over failures in their lives, helplessness ensues. In turn, if feelings of helplessness persist, full-blown depression can often follow (Seligman 1989). Specifically, attribution of a lack of con-trol to internal, stable, and global factors is what sets up the fundamental conditions for helplessness and depres-sion to arise (Seligman et al. 1979). These ideas provide a strong theoretical framework with clear conceptual links to the sense of agency. In fact, the suggestion from Seligman’s work is that a loss of the sense of agency is actually causally linked to the development of depression.

Similarly, the phenomenon of “depressive realism” high-lights the link between the sense of agency and depression. This phenomenon refers to the finding that depressed indi-viduals are more likely to make accurate judgments regard-ing the contingency between responses and outcomes, even when they experience a relatively high frequency of desired outcomes. In contrast, non-depressed individuals fall prey to an illusion of control whereby a higher frequency of desired outcomes is associated with an enhanced perception of con-tingency between action and effect (Alloy and Abramson

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1979; see also Allan et al. 2007). This finding has led to the claim that depressed individuals are sadder, but perhaps wiser than their non-depressed counterparts (Alloy and Abramson 1979).

In view of this previous research, the motivation for assessing the relationship between depression and interval estimation is clear. A first step toward validating the sug-gestion that estimates of action–effect intervals could be used to assess treatment-induced “agency improvement” is to determine whether or not perceived intervals are dif-ferent in individuals who feel anxious or depressed. In addition, if, for example, interval estimates were found to be longer for individuals who feel depressed, it would bol-ster the very notion that this measure relates to the sense of agency. While it would be ideal to assess interval esti-mation in depressed patient samples and compare them to their non-depressed counterparts, there are many challenges with such an approach. First, depression is often co-morbid with a host of other conditions, like anxiety, pain, chronic illness, or other mental health issues (Mclntyre et al. 2012). Second, it is difficult to account for the effects of medica-tion and drugs on performance in cognitive tasks such as interval estimation. Both these concerns diminish the ease with which the effects of depression per se can be isolated in an experimental study. Thus, as a first step toward study-ing the potential link between the experience of voluntary action and effects, and depression, we opted to use a prim-ing procedure, to activate memories of a depressing episode in psychiatrically intact individuals.

Specifically, to investigate whether feeling depressed modulates interval estimation, we used a similar episodic priming approach to that which we used (and others have used extensively) to prime social power (Obhi et al. 2012; Galinsky et al. 2003). Healthy participants wrote about a depressing episode from their own life, or they wrote about what they did the previous day (“previous day priming”), or they did not write about anything (baseline), prior to completing an interval estimation task in which they judged intervals between actions and effects. Due to the fact that depression is associated with a perceived loss of control, we predicted that after recalling a depressing episode, partici-pants would report action–effect intervals as being longer than in a baseline condition with no priming, or the “pre-vious day” priming condition. In addition, we predicted no difference between interval estimates in the “previous day” priming condition and the baseline (i.e., no priming) condition.

Methods

Thirty-six healthy participants (21 females, 15 males), with a mean age of 25 years, gave informed consent and

took part in the experiment for monetary compensation, or partial course credit. The study conformed to local ethical guidelines for conducting research with human participants.

Apparatus and stimuli

The experimental task involved participants sitting approxi-mately 65 cm in front of a Dell 19-inch LCD computer monitor on which was displayed an image of an avatar with their right index finger in an extended position (image appeared in the center of the screen and was 8.5 × 11 cm in size). The avatar was shown sitting at a desk, and a fixation cross was superimposed in the middle of the image. Par-ticipants fixated the cross throughout the trial and, on each trial, they were required to press a key on a Cedrus RB-730 response pad with their right index finger, at a time of their own choosing. The key press was registered by Superlab 4.5 (Cedrus corporation, CA) software and caused the previ-ously displayed photographic image of the avatar with their index finger in raised (extended) to transition to a photo-graphic image of the avatar with their index finger lowered (flexed). The interval between the participant’s key press and the transition of the image from finger extended to fin-ger flexed was either 100, 400 or 700 ms (Obhi et al. 2012), and intervals were pseudo-randomly selected and equally probable throughout each experimental block. Participants gave verbal reports of the perceived interval between their key press (i.e., the action) and the transition to the image with the index finger flexed (i.e., the effect). Priming pro-cedures were carried out on a desktop computer, into which participants typed their recollection of a depressing episode (depression priming), or a chronological description of their previous day (previous day priming). In a baseline condi-tion, they did not write about anything.

Procedure

Participants arrived at the lab under the impression they were taking part in two separate studies, one on “action aware-ness” and the other on “adjustment states.” Upon arrival, an experimenter informed participants, that, instead of com-pleting the experiments sequentially, they could complete parts of the adjustment states experiment in between blocks of the action awareness experiment to “save time.” All par-ticipants agreed to complete the experiments in this manner to save time.

To avoid contamination by any form of priming proce-dure, participants always performed the baseline condi-tion of the interval estimation task first, followed by either a post-depression priming block or previous day priming block (Obhi et al. 2012; order of depression and previous day blocks was counterbalanced across participants).

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In all experimental blocks, participants sat in front of a computer monitor, with their right index finger resting lightly on a response pad key hidden from view under an occlusion platform. Prior to the baseline block, participants completed a block of 5 practice trials, to familiarize them-selves with the procedure. In the experimental blocks (base-line, depression priming, and previous day priming), each trial began with the appearance, in the center of the screen, of a third person image of a model (i.e., an avatar) seated at a desk with their right index finger raised (extended). The avatar was shown from the neck down, thus avoiding any effects related to seeing the avatar’s face. As shown in Fig. 1, on each trial, participants pressed the response pad key at a time of their own choosing, causing the image on screen to change to an image of the same avatar with their index finger lowered after one of three intervals (100, 400, 700 ms). Participants were instructed that their task was to estimate the interval in milliseconds between their key press and the transition to the second image. They were informed that the interval could range between 0 and 999 ms and they were reminded that 1 s is 1,000 ms. To participants it appeared that their key press caused the ava-tar to lower their index finger. After the presentation of the second image, participants gave a verbal report of the per-ceived action–effect interval duration, which was recorded

by the experimenter. There were 30 trials in each experi-mental block.

After completing the baseline block, participants were told they could take a break from the awareness study and do a portion of the adjustment states study. At this point, they were asked to write about either a depressing episode from their own lives (depression priming) or give a descrip-tion of the events on the previous day (previous day prim-ing). The depression priming required participants to focus on how sad and depressed the episode had made them feel. Half the participants did the depression priming procedure immediately after the baseline block and the other half did the previous day priming procedure immediately after the baseline block. After performing another block of 30 trials, participants were asked to complete a mundane word search task for 5 min. After this task, participants were exposed to the other priming procedure (depression or previous day, depending on which they had done after the baseline block). The word search task was included to allow some time for any effects of the previous priming procedure to decay, at least partially. Words in the word search task were mundane, non-emotional words such as “desk,” “chair,” etc. After the word search task, participants completed the alternate prim-ing procedure and finally completed another 30 trials of the interval estimation task.

Fig. 1 Trial procedure in the experiment. Participants pressed a response pad key at a time of their own choosing which triggered a transition from an image of an avatar with their right index finger extended, to an image of an avatar with their right index finger flexed. The second image stayed on screen for 1,250 ms. At the end of each trial, participants verbally reported the perceived interval between their action and the on-screen effect in milliseconds (0–999)

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Manipulation check

To verify that depression priming resulted in episodic recol-lections that were indeed different in terms of depressive content than the previous day descriptions, off line coding of the recalled episodes was performed. Specifically, after the experiment, two independent coders coded the recollec-tions for presence of depression and emotion. Coding was performed using a scale from −3 (no depression or sadness, no emotional content) to +3 (a strong degree of depression or sadness, strong emotional content).

Trait measure of depression

After the experiment, participants completed the CES-D scale (Radloff 1977). This scale is a depression measure-ment instrument for use in the general population. After completing this scale, participants were debriefed and asked whether they had any hypotheses about what the experi-menter’s were investigating.

Control experiment 1: Ruling out general arousal effects

In order to rule out the possibility that general differences in the degree of arousal in the depression condition and the pre-vious day priming condition could account for any observed differences between conditions, we conducted an additional control experiment with ten new participants (ages 18–25). In this experiment, participants again underwent the depres-sion priming and previous day priming procedures (order counterbalanced across the participants). However, we took the additional step of measuring participants’ heart rate while they wrote about each episode. Any differences in arousal between the two conditions would then be apparent. Heart rate data were measured with a finger pulse transducer connected to an AD instruments psychophysiological data acquisition system, and data were analyzed in LabChart software (AD instruments, Colorado Springs, CO, USA). Average heart rate for the entire time taken to write each rec-ollection was calculated offline and compared to determine whether any differences in arousal were present between the two priming conditions.

Control experiment 2: Ruling out effects on general time perception

In the main experimental procedure, any differences in interval estimation could theoretically be due to a general effect of depression priming on time perception and may not be solely linked to interval estimation in tasks involv-ing an agentic component. To determine whether there were any general effects on time perception of depression priming, we engaged ten new participants (ages 18–21)

in a scaled down version of the main experiment. We included only a depression priming condition and a no-priming condition, and instead of judging action–effect intervals as in the main experiment, participants simply judged the time interval between two auditory tones, pre-sented on a computer. Tone separation was either 100, 400 or 700 ms, and there were 30 trials in each block. The baseline condition was always run prior to the depression priming condition.

Results

Outlier exclusion

The main dependent variable in this experiment was the action–effect interval judgment. In the data-processing stage, any judgments that were more than 3 standard deviations away from the participant’s mean judgment for that condition were excluded from further analysis. This procedure resulted in removal of less than 0.3 % of the data.

Manipulation check

Two independent raters coded each episodic recollec-tion from the priming procedures for depressive content and emotional content. Coding was done on a scale from −3 to +3, and inter-rater correlations were significant for both recollection types (depression recollection condition: depressive content correlation = .503, p < .05; emotional content correlation = .358, p < .05; previous day condition: depressive content correlation = 0.488, p < .05; emotional content correlation = .540, p < .05). Having established these correlations, for the depression priming and previ-ous day priming conditions, the two sets of rater scores for depressing and emotional content were averaged and subjected to statistical analysis. As Table 1 shows, t tests on this coding data revealed that recollections of a depressing episode differed significantly from recollections of the previ-ous day on both types of content. Specifically, recollections

Table 1 Mean depression and emotion-related content in the recol-lections of depressing episodes and the previous day

Content Depressing episode

Previous day description

t test

Depressive M = 2.0, SD = 0.78

M = − 2.80, SD = 0.34

t(36) = − 35.95, p < .001

Emotional M = 2.0, SD = 1.0

M = − 2.50, SD = 0.55

t(36) = − 26.64, p < .001

Scores are from a −3 to +3 scale. t tests reveal that the recollections differed on both types of content

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of depressive episodes contained more depressive content and more emotional content than recollections of events on the previous day. Post-experiment debriefing revealed that none of the 36 participants had any idea of the true purpose of the experiment.

Differences in interval judgments across the experimental conditions

For the main analysis, for each condition (baseline, depres-sion priming, previous day priming), interval judgments were collapsed across the three different (100, 400, and 700 ms) action–effect intervals. Mean judged intervals for each participant in each condition were entered into a repeated measures ANOVA with three levels of the condi-tion factor (levels: baseline (no priming), depression prim-ing, and “previous day” priming). This analysis revealed a significant main effect of condition (F(1.6, 56.7) = 3.574, p = .043—since Mauchley’s test of sphericity was signifi-cant, we report the Greenhouse–Geisser statistic).

Given our directional predictions that depression prim-ing would produce interval judgments that were longer than baseline and previous day priming judgments and that previ-ous day priming and baseline priming judgments would not differ, we conducted a series of one tailed t tests. These tests revealed that the depression priming condition did indeed yield interval judgments that were significantly longer than those obtained in the baseline condition (depression condi-tion = 424 ± 160 ms, baseline condition = 380 ± 158 ms; t(35) = −2.332, p = .013), and the previous day priming condition (depression condition = 424 ± 160 ms, previous day priming = 404 ± 151 ms; t(35) = −1.722, p = .047). In contrast, interval judgments in the previous day priming con-dition were not significantly different from baseline (previ-ous day condition = 404 ± 151, baseline = 380 ± 158 ms; p > .05).

Figure 2 shows the mean interval judgments for the three conditions.

Trait measures of depression

The CES-D scale is a self-report depression scale for research in the general population. Different scores corre-spond to different severities of trait depression, with a score of less than 15 indicating that participants are not depressed, a score of 15–22 indicating that participants are moderately depressed and a score of over 22 indicating that partici-pants may be very depressed (Radloff 1977). For the pur-poses of the present study, we split participants into those who scored below 15 and those who scored above 15 (for a similar approach, see St. John and Montgomery 2009). We then determined the change in interval estimate duration (%) in the depression condition compared to the baseline

condition for each participant in each sub-group (Δestimate) (high trait depression, n = 18; low trait depression, n = 18). An independent samples t test was conducted to determine whether high and low trait depression sub-groups differed on the Δestimate measure. This test revealed that there was no significant difference between high and low trait depression sub-groups on the Δestimate measure (CES-D score >15, Δestimate = 30 ± 63); CES-D score <15, Δestimate = 13 ± 29, p > .05). Despite this lack of significance, there was a pat-tern in which the high trait depression sub-group showed a larger Δestimate than the low trait depression sub-group. However, the t test result suggests that, in the main analy-sis, differences in interval estimates between the baseline condition and the depression priming condition were driven primarily by the priming manipulation, and not by differ-ences in trait levels of depression.

Control experiment 1: Assessing differences in arousal between priming conditions

In order to rule out the possibility that differences in arousal during the two priming procedures (depression priming and previous day priming) contributed to the observed effects on interval estimation, average heart rate data taken from ten new participants while they wrote each episode type was entered into a t test. This test revealed that there was no difference in average heart rate between the two prim-ing procedures and thus provides some confidence that our effects were not due to basic differences in the degree of arousal caused by the priming procedures (depression priming HR = 88 ± 13 bpm versus previous day priming HR = 89 ± 11 bpm; t(9) = 0.994, p = .346).

Fig. 2 Mean interval estimates for each of the three experimental conditions. Note that mean judged intervals were significantly longer after depression priming than after baseline or previous day priming (denoted by asterisk, see text for statistics). In contrast, there was no significant difference between interval estimates from previous day priming and baseline conditions. Error bars are SEM

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Control experiment 2: Ruling out effects of priming on general time perception

In order to rule out that possibility that depression prim-ing in particular causes general differences in time percep-tion that could explain our effects, we entered data from ten new participants’ judgments of tone intervals into a paired samples t test. This test revealed a significant difference in estimates between the baseline and the depression condition (t(9) = 2.401, p = .040). Critically, however, judgments of the interval between two auditory tones were shorter after depression priming compared to baseline (judged interval after depression priming: 245 ± 107 ms, versus judged inter-val, baseline: 277 ± 98 ms). Thus, the difference between conditions when participants judged intervals between two tones is in the opposite direction to the difference found in the main analysis, wherein estimates of intervals between an action and effect were longer after depression priming than after baseline or previous day priming. This control experi-ment effectively rules out the possibility that general dif-ferences in time perception associated with depression can account for our main findings.

Discussion

The sense of agency is a profoundly important human expe-rience, which is altered in several psychiatric conditions including schizophrenia, anxiety disorders, and depression (Hauser et al. 2011; Mirowsky and Ross 1990; Rapee et al. 1996; Seligman 1989). The compression of the perceived interval between an action and effect (i.e., intentional bind-ing) is thought to reflect the sense of agency (Haggard et al. 2002; Moore and Obhi 2012). The present experi-ment sought to determine whether activating memories of a depressive episode affected the perceived interval between a voluntary action and the consequent movement of an avatar on a computer screen. To investigate this, in a within sub-jects design, we asked participants to write about a personal experience where they felt depressed or about the events of the previous day in chronological order, before engaging in an interval estimation task. In a baseline condition, which always occurred prior to the priming conditions, partici-pants simply made judgments about the temporal interval between a self-paced voluntary key press and the movement of an avatar on a computer screen. We found that interval estimates after depression priming were significantly longer than interval estimates obtained in the previous day priming condition, or the baseline condition. Inclusion of the pre-vious day priming condition is important as it rules out a general effect of recalling any personal episode on interval estimation and allows us to somewhat isolate the effect of depressed feelings on the temporal experience of actions

and effects. Because we used movement of an on-screen avatar as the movement “effect” in this experiment, it is tempting to conclude that activating memories of depression causes differences in the experience of control over others in social contexts. While this may be the case, it is important to point out that we did not compare this scenario with another scenario involving non-social events such as the appearance of a color patch or the sounding of a tone. Comparing these situations could form the basis for interesting future studies on how activating depressive memories changes the experi-ence of voluntary action in social and non-social contexts.

A manipulation check confirmed that the recollections provided by participants in the depression condition and the previous day condition did contain different levels of depression and emotion. Furthermore, differences in trait depression scores could not account for the effect. Two control experiments also ruled out that basic differences in arousal between the priming conditions might account for the observed effect and that our effects are simply a spe-cific case of a depression-related change in time perception. Specifically, when asked to judge the interval between two tones (i.e., a judgment about two external events with no agentic component), participants in our control experiment showed a reversed pattern of estimates, with post-depression priming resulting in shorter estimates than baseline esti-mates (see also Gil and Droit-Volet 2009). Finally, the use of a within subjects design provided a strong test of the effects of activating memories of depression on interval estimation. Specifically, because all participants were exposed to both prime types, there was likely to have been some carry over between blocks, which would have worked to reduce the difference between conditions. Despite considerable effort to control or rule out alternative explanations, other possi-ble causes for the observed effects should be considered in future studies. In particular, we did not rule out that differ-ences in the degree of cognitive load between the various conditions could contribute to our findings or that emotional priming per se could cause the effects we observed. Despite these possibilities, there is reason to predict the effects we found on the basis of the extant literature on depression and the perception of control. In this vein, ours is the first dem-onstration (to our knowledge) that action–effect interval estimates are affected by activating memories of depression.

Major depression is a common and debilitating psychiatric illness with prevalence in the United States of between 6 and 9 % and an annual economic cost of $83.1 billion (Richards 2011; Greenberg et al. 2003). The figures in Europe are not much different, and the World Health Organization has shown that depressive disorders are one of the major causes of disease worldwide (Richards 2011). The condition is characterized by a constellation of symptoms, including depressive mood, a lack of interest in most activities and feelings of worthlessness and guilt (APA 2000). A key

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psychological component relating to depression is the locus of control. Studies have found that patients with a low inter-nal locus of control experience higher levels of depression (Ausland 2012). Correspondingly, a greater external locus of control has been associated with more depression (Benassi et al. 1988). More generally, studies have also shown that a high internal locus of control and a low external locus of control are associated with positive health outcomes (e.g., Brown et al. 2000; Benassi et al. 1988). Indeed, in a recent paper Keeton et al. (2008) highlighted studies showing that an impaired sense of control is associated with depression, stress and other anxiety related disorders (Abramson et al. 1989; Chorpita and Barlow 1998; Mirowsky and Ross 1999; Shapiro et al. 1996). Relatedly, the sense of control and the feeling of powerlessness sit at opposite ends of a continuum and are associated with positive and negative mental health outcomes, respectively (Keeton et al. 2008; Mirowsky and Ross 1991).

We recently performed an experiment to assess whether different levels of power were associated with differences in the temporal experience of actions and effects (Obhi et al. 2012). We found that recalling an episode of pow-erlessness was associated with longer interval estimates (i.e., reduced intentional binding) compared to recall-ing an episode of powerfulness, or a baseline interval estimation condition. The work reported in the current paper extends this result by demonstrating that activat-ing memories of depression also results in longer interval estimates of the duration between a voluntary action and subsequent movement of an on-screen avatar, compared with activating memories related to a chronology of the previous day’s events, or a baseline condition. These results are the first to demonstrate a direct link between depression and the temporal experience of actions and effects.

The current results bolster the idea that changes in perception of action–effect intervals are tied to agency experience, and also reinforce the tentative suggestion we made in our recent study on power and agency (Obhi et al. 2012). In that study, we showed that feeling relatively powerless results in longer action–effect interval estimates. Because several psychiatric conditions including depression are characterized by a loss of control, which may be similar to feeling powerless, we hypothesized that patients who feel a lack of control may also show a similar pattern of interval estimates. The results from the present study underscore the potential of using interval estimation as a measure of agency in individuals who suffer from an abnormal sense of control. Given that interval estimates are lengthened for individuals who have just activated memories of a depressing episode, but not for the same individuals when they simply recall the events of the previous day, we suggest that such estimates

could be a useful index of agency in patients, pre-, during, and post-treatment.

For example, a common treatment option for depressed individuals is a course of cognitive behavioral therapy (Sudak 2012; Williams 1984). Among other diagnos-tics, the interval estimation task could be administered to such patients prior to their first therapeutic session and then again at intermittent times during the treatment pro-cess, and of course, after treatment is completed. Indeed, another approach to the treatment of depression is the use of non-invasive brain stimulation to reduce the excitabil-ity in specific brain regions (Pallanti et al. 2012). Interval estimates could be obtained before and after a course of brain stimulation therapy, to implicitly assess changes in the sense of agency. The task could also be administered in the weeks/months following the cessation of treatment to track any subsequent fluctuations in the sense of agency during this time. Of course, this notion needs to be further investigated, and it remains for future research to do this; efforts are underway in our laboratory to explore this idea more comprehensively.

One question is why such an implicit measure of agency is potentially valuable in the clinical domain? In this regard, it is important to note that much more work is needed to assess the reliability of the potential relationship between interval estimate measures and depression suggested in the current study. This will obviously necessitate conducting careful and well-controlled experiments on depressed par-ticipant samples. However, if a robust relationship is found, all the benefits of an implicit versus an explicit assessment become relevant. That is, due to the relatively indirect man-ner in which interval estimation might reveal information about a depressive state, it becomes difficult for factors such as impression management, demand characteristics and attempts to provide expected or socially normative responses to play a role (as could be the case in explicit assessments) (McCroskey 1984).

In sum, we have shown that recalling a depressing episode produces estimates of action–effect intervals that are longer than judged intervals in a baseline condition and also longer than interval estimates made after recalling the previous day’s events. Thus, activating memories of depression affects the subjective experience of voluntary actions and their effects. Our results support the notion of a relationship between per-ceived action–effect intervals and agency and also highlight the potential link between depression and the experience of voluntary action. Finally, we also suggest that, after further careful validation employing actual depressive samples, there could be a potential use for interval estimation tasks in the implicit assessment of the sense of agency pre-, during, and post-treatment, in individuals with disorders that affect their sense of control.

505Exp Brain Res (2013) 229:497–506

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Acknowledgments This work was funded by a Natural Sciences and Engineering Research Council discovery grant awarded to S. S. O.

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