Classical fear conditioning – including acquisition and extinction – is a model for fear learning and memory in health and disease. Moreover, trauma-related disorders can be viewed as comprising fear acquisition under severe stress. Yet, in humans, we know comparatively little about how acute stress affects fear conditioning. Therefore, the first aim of this thesis was to investigate the effect of stress on fear acquisition or extinction. Stress induces multiple hormonal and neurotransmitter changes dynamically developing over time, including a fast first-wave and a slower second-wave stress response. Models derived from avoidance learning and declarative memory studies suggest that stress effects on memory depend on the temporal proximity between learning and stressor: encoding close to the stressor will be enhanced, but encoding and recall later in time (during the second-wave) will be suppressed (e.g., Schwabe, Joëls, Roozendaal, Wolf, & Oitzl, 2012). So far, these predictions were not related to fear conditioning. Therefore, we investigated if the model-based predictions are also valid in human fear conditioning. We used two stressors to investigate first-wave and second-wave stress effects: the cold pressor test (CPT) inducing a strong first-wave but little second-wave activation and a psychosocial stressor, reliably inducing both, first- and second-wave stress responses. Conditioning was measured via skin conductance responses (SCRs). Investigating the first-wave (Experiment 2), we placed fear acquisition and immediate extinction directly after the CPT (n = 20) or after the control treatment (n = 20). We found no group difference in acquisition performance, but significantly increased extinction resistance in the stressed CPT group. In Experiment 3, CPT (n = 20) or control (n = 20) was placed after acquisition but directly prior to extinction training. Here, we found improved extinction and 24h-delayed extinction recall after CPT. Investigating the second-wave (Experiment 1), we placed fear acquisition and immediate extinction 45 min after the psychosocial stressor (i.e., at the peak of salivary cortisol, n = 12) or after control (n = 12). Here, we found no significant stress effects. Sex and female sex hormones also influence fear conditioning: Women are at a higher risk to develop anxiety and stressor-related disorders than men. Interestingly, patients with these disorders show impaired fear extinction and extinction recall, and low levels of the sex hormone 17β-estradiol (E2) are linked to impaired extinction in both, healthy and patient female samples. So far, there is little data on how acute stress and circulating E2-levels might interact in fear acquisition and especially in fear extinction. Therefore, the second aim of this thesis was to explore this possible interaction in healthy women in different cycle phases compared to men. Thus, in Experiment 4, we included hormone status as a quasi-experimental variable and compared free cycling women in the midcycle phase (high E2, low progesterone, n = 24), women in the early follicular phase of the menstrual cycle (low E2, low progesterone, n = 24), and men (n = 24). We placed fear acquisition and extinction 45 min after the psychosocial stressor (n = 36) or control (n = 36), and tested extinction recall after 24 h. In line with Experiment 1, the second-wave stressor did not affect fear acquisition and immediate extinction. However, we found a stress by hormone status interaction within women at the 24h-delayed extinction recall test: in the stressed group, early follicular women showed impaired extinction recall and a higher return of fear compared to midcycle women, whereas there was no difference between early follicular and midcycle women after control treatment. Collectively our results support a different role for the first- and second-wave stress response in human fear conditioning. Fear acquisition near the first-wave stress response results in enhanced fear memory, which is resistant to extinction. Extinction training near the first-wave enhances extinction learning. In contrast, fear conditioning at the peak of the peripheral second-wave cortisol response had no effect on acquisition or extinction performance. However, second-wave stress interacted with the hormone status of women, where only women in a low E2 state were vulnerable to negative stress effects in extinction recall. The last result will encourage further investigation of the interplay between E2 and stress in fear extinction. Enhancement of extinction by the CPT could – if replicated – be translated into strategies for optimizing exposure therapy.

This dissertation extends previous work on the Positive and Negative Affect Test (IPANAT), a test that was developed to measure affective processes at an implicit (supposedly automatic, preconscious) level. In particular, the first manuscript that is included in this dissertation demonstrates that the IPANAT picks up on processes resembling counter-regulation (cf. Rothermund, Voss, & Wentura, 2008) after exposure to negatively valenced emotional stimuli. The IPANAT might therefore be a suitable measure of affect regulation in some contexts. Furthermore, the second manuscript introduces a version of the IPANAT that was designed for the measurement of distinct emotions, the IPANAT for distinct emotions (IPANAT-DE). Finally, the third manuscript and an additional supplemental study not yet submitted for publication provide evidence that the subscales of the IPANAT-DE are sensitive to priming with different emotional expressions, and that facial expressions of the same valence but of different distinct emotions trigger emotion specific changes of IPANAT-DE scores. In the last part of this dissertation, the three manuscripts and the supplemental study are discussed with respect to three overarching issues: 1. The kinds of processes that might drive IPANAT scores and whether these processes are indeed influenced by affective experience or whether they are purely cognitive. 2. How IPANAT scores are related to other aspects of affect. 3. How the IPANAT can be adapted to different research questions as well as to applied contexts.

The ultimate aim of psychological research is to disentangle everyday human functioning. Achieving this goal has always been limited by the necessity of balancing experimental control and ecological validity. Recent technical advances, however, reduce this trade-off immensely, perhaps even rendering it void: Sophisticated virtual reality (VR) systems provide not only high experimental control but also multidimensional and realistic stimuli, tasks, and experimental setups. Yet prior to applying VR as a standalone experimental method, an empirical foundation for its application needs to be established. To this end, this dissertation aims to shed light on whether and which changes in cognitive-affective standard findings result from increasing the ecological validity by means of VR paradigms. The four empirical studies included in this dissertation focus either on the affective or mnemonic processes and mechanisms occurring under immersive VR conditions compared to conventional laboratory setups. Study 1.1 investigated whether the electrophysiological correlates of the approach/avoidance dimension differ depending on the mode of presentation, i.e., immersive VR footage or a virtual 2D desktop. Study 2 was extended by a behavioral component. Full-body responses were enabled within this paradigm to examine holistic fear responses and to put to the test whether the respective electrophysiological responses translate from keystrokes to natural responses. With respect to the retrieval of such immersive experiences, Study 1.2 aimed to replicate the memory superiority effect found for VR conditions compared to conventional conditions. The generalizability of this effect will be examined using complex, multimodal scenes. Going one step further, Study 3 differentiated the retrieval mechanisms underlying VR-based or conventional laboratory engrams on the electrophysiological level. The well-established theta old/new effect served as a benchmark to check whether cognitive processes obtained under conventional conditions translate to VR conditions. The results of these studies are discussed with respect to whether and how increasing ecological validity alters the standard findings expected on the basis of the previous research background. Special attention will be paid to the differences between conventional laboratory setups and sophisticated VR setups with the aim to identify possible sources of the obtained deviations from standard findings. Such changes in the findings that overlap and exceed all studies beyond their primary focus, whether emotional or mnemonic, are discussed in terms of embodied simulations and the predictive coding hypothesis. A shared mental 3D default space is proposed as a possible source of fundamental differences between conventional and VR-based research outcomes. In particular, it will be demonstrated that conventional research approaches and findings may not only be amplified but fundamentally altered when translated to VR paradigms.

Emotion, as well as cognition, are often understood as a manifestation of brain activity. However, bodily processes are also involved in mental functioning, referring to the concept of embodiment. Embodied emotion, traditionally, implies that experiencing an emotion involves perceptual, somato-visceral, and motor aspects. Within the frame of the Research Training Group “Situated Cognition”, we here extend the concept of embodiment by considering the role of hormones in the processing of emotional content. Importantly, hormones allow a bidirectional body-to-brain and brain-to-body coupling. The endocrine system, e.g., steroid sex hormones, produced in the gonads, send feedback to the brain by binding at their receptors. These receptors are relatively abundant in the brain regions associated with emotional processing, memory, and executive functions (i.e., amygdala, hippocampus, and prefrontal cortex). Moreover, peripheral hormone secretion is modulated via actions from the central nervous system. We intended to characterize the role of sex hormones, and partly also of stress hormones, on different components of emotion as a hormonal embodiment of emotion. Thus, we examined emotional processing in different sex hormone-status groups. To account for different levels of sex hormones, we used a quasi-experimental approach by comparing women in different cycle phases, women using hormonal oral contraceptives (Study 1), and additionally men (in Study 2). The female menstrual cycle is characterized by fluctuating sex hormone levels. On the peripheral gonadal level, these are 17β-estradiol and progesterone. These hormones are low at the beginning of the cycle (early follicular phase). Estradiol rises towards the middle of the cycle (mid-cycle) and stays moderately high until the next cycle. Progesterone levels are high after mid-cycle in the luteal phase until the end of the cycle. Hormonal contraceptives suppress the endogenous production of estradiol and progesterone, keeping the hormone levels low during the whole cycle. Estradiol and progesterone are also present in males, however, at low levels with no sign of cyclical fluctuations. In Study 1, we examined three independent groups of women in the mid-cycle (n = 24), in the luteal phase (n = 24), and women using hormonal oral contraceptives (n = 24). We assessed different measures of emotional processing, i. e. emotional memory, cognitive and affective empathy-related measures (emotion recognition and ratings for feeling with a protagonist´s emotion, respectively), as well as mimic and skin-conductance responses to affective stimuli. Additionally, we addressed interactions of experimental stress (cold pressor test vs. control) with sex hormones in emotional memory. Our data demonstrated the role of hormones in empathy-related measures and skin-conductance responses depending on the stimulus characteristics (valence, the gender of the protagonist). Emotional memory was not affected by hormone status, stressor or salivary hormone levels. In the cognitive empathy-related measure, women in the luteal phase, as well as oral contraceptive users, identified emotions depicted by female protagonists more accurately than those by male protagonists. On the other hand, estradiol correlated positively with recognition of emotions depicted by males in the total sample. In the affective empathy-related measure, oral contraceptive users rated negative emotions higher than the positive ones. Finally, in the luteal phase skin-conductance responses to negative stimuli were heightened, also supported by a positive correlation with the salivary progesterone levels. The mimic responses remained unaffected. None of the remaining associations with the salivary hormone levels were significant. These results indicate that sex hormones modulated emotional processing by interacting with the stimulus features, as evident in the negativity bias under oral contraceptive use and in the luteal phase in the affective empathy-related measure and sympathetic autonomous reactivity, respectively. However, emotional memory and mimic activity to affective stimuli were not affected. In Study 2, we extended the initial scope to examine the role of sex hormones and olfaction in empathy-related measures. Reports of female advantage in empathy-related measures suggest a role for sex hormones, although data are inconsistent. Studies also report similar sex differences in human olfactory perception. In rodents, olfaction is involved in detecting and integrating socially-relevant information and is modulated by the brain-actions of estrogens. Based on this background, we hypothesized that olfaction may untangle the mixed evidence regarding the relationship between sex hormones and empathy-related measures (cognitive, affective). Thus, we measured odor discrimination ability, empathy-related measures, and facial mimic activity (also associated with affective empathy-related measures) in free-cycling women in high sex-hormone phases (n = 20), oral contraceptive users (n = 19), and men (n = 21). Free-cycling women outperformed only men in the recognition of emotions depicted from the eye region. Oral contraceptive users showed higher scores in the affective empathy-related measure towards negative emotions. Free-cycling women exhibited the strongest facial mimicry (viewing female, but not male protagonists), positively associated with progesterone. Finally, the groups differed in odor discrimination, with free-cycling women outperforming men. However, odor discrimination ability and empathy-related performance were not correlated. Our results support the role of sex hormones in odor perception and empathy-related measures, to a certain extent. However, no common underlying mechanism was found. Finally, we conducted a systematic review (Study 3) aiming to elucidate factors contributing to the inconsistent results concerning the role of sex hormones in the two most addressed areas of emotional processing, emotion recognition (empathy-related measure) and emotional memory. Thereby, we extended previous reviews that address single areas of emotion processing. Moreover, we systematically addressed the role of situational features (mainly emotion-type and/or stimulus valence). All studies included healthy women of reproductive age either in stages of their natural menstrual cycle or using oral contraceptives, and measured or at least estimated levels of ovarian sex hormones. We document the methodological diversity in the field, presumably contributing to the heterogeneity of results. We recognized the need for studies explicitly contrasting the early follicular, mid-cycle, and mid-luteal phases, as well as OC-intake and using standardized tasks. Research would take advantage of using within-subject design more frequently and account for the recognition of complex emotions. In sum, our data suggest that sex hormones differentially modulate the cognitive and affective empathy-related performance and skin-conductance responses by interacting with situational variables, such as the emotional valence of the stimuli and the gender of the protagonist. Women in the luteal phase and under oral contraceptive use demonstrated better recognition of emotions depicted by female protagonists. By contrast, estradiol levels positively correlated with the recognition of emotions depicted by male protagonists. Sex-hormone status main effects only manifested in the emotion recognition advantage of free-cycling women over men (Reading the Mind in The Eyes Test; Study 2). In both studies, affective empathy ratings towards negative emotions were higher in the oral contraceptive users. Moreover, although mimic activity was not associated with sex hormones, skin-conductance responses to negative stimuli were heightened in the luteal phase. On the other hand, the performance in empathy-related measures in different hormone-status groups was not related to odor discrimination ability. Additionally, the inconsistencies of the sex hormone and emotion research could be the result of variations of designs and tasks used across studies from a similar field. This is also indicated in our findings from the empathy-related measures differing in tasks and hormone-status groups in two studies. Finally, our findings provide evidence that emotional processes under sex-hormone modulation are situated, i.e., subject to the influence of the stimulus valence. Furthermore, they are embodied via coupling between the endocrine system and the brain as evident in hormone status and valence interactions in empathy-related measures and sympathetic reactivity.

In dieser Arbeit entwickle ich den Begriff "habitualisierte affektive Intentionalität", um erklären zu können, auf welche Weise Emotionen in der sozio-kulturell spezifischen Lebenswelt epistemisch problematisch oder förderlich wirken angesichts des epistemischen Ziels des Verstehens. Dieser Begriff erlaubt die Weltbezüglichkeit von Emotionen aus multidimensional situierter Perspektive und mit Fokus auf die lebensformspezifisch habitualisierte Dimension des Fühlens auszuleuchten. Erstens verstehe ich affektive Intentionalität als irreduzibel komplexe Weise des Weltbezugs (Kapitel 2), zweitens als strukturiert durch die je sozio-kulturell spezifische Einbettung (Kapitel 3) und drittens als strukturiert durch einen individuellen aber in sozio-kultureller Umgebung gegossenen Habitualisierungsprozess (Kapitel 4). Der resultierende Begriff lässt verstehen, inwiefern das, was durch Emotionen verstanden wird, strukturiert ist durch a) präreflexive, fungierende Intentionalität und Leiblichkeit, b) situative Faktoren, c) Emotionsrepertoire und habitualisierte Angänglichkeit. Die epistemische Relevanz des Fühlens erscheint dadurch in einer Perspektive, die insbesondere die Möglichkeitsbedingungen spezifischen affektiven Verstehens auslotet. Gefühle beeinträchtigen Verstehensprozesse u.a. dadurch, dass Subjekte a) affektiv an Lebensformen anhaften und sich b) bestimmte Weisen der Wahrnehmung verknöchern. Dies einzusehen ist die notwendige Bedingung, um über Lösungen nachdenken zu können – u. a. Weisen der (Ver)Störung, die ein Verstehen jenseits von erstarrten Wahrnehmungsmustern möglich machen können (Kapitel 5).

Using action orientation after failure as a measure of individual differences in emotion regulation abilities (ERA), this thesis’ studies investigated the impact of ERA on cognition, behavior, and own versus imposed goals differentiation. The first study used cortisol as a physiological stress marker to replicate the link between ERA and the ability to make intuitive judgments under stress. High ERA were associated with increased performance in an intuition task under stress. In contrast, when feeling no stress, low ERA were associated with increased performance in an intuition task. The second study showed that ERA can compensate for depression-associated biased processing of negative stimuli. This effect was present even at mild to moderate depression levels. Replicating earlier findings, the third study showed that ERA are associated with an increased ability to distinguish self-chosen from imposed goals. Most importantly, the study identified activation in the right medial prefrontal cortex as a neural correlate of identifying self-chosen goals, and activation in the anterior cingulate cortex, as a correlate of falsely identifying imposed goals as self-chosen ones. Altogether, these studies show the necessity to consider individual differences in ERA in stress, clinical, and motivational research. The findings are discussed with respect to three theories that relate to motivation and personality from behavioral and neurobiological perspectives, namely, Personality Systems Interaction Theory, Predictive and Reactive Control Systems Theory, and Self-Determination Theory.

Die Studie untersucht die Wirksamkeit eines neuen Zieltypen (Motto-Ziel), welcher im Rahmen der theoretischen Überlegungen und praktischen Erfahrungen mit dem Selbstmanagement-Training nach dem Zürcher Ressourcen Modell ZRM (Storch & Krause, 2007) entwickelt wurde (Storch, 2009). Durch die Verankerung im Unbewussten ermöglichen es Motto-Ziele Menschen Handlungen in der Selbstregulation auszuführen, für welche normalerweise Selbstkontrolle benötigt wird (unangenehme Pflichten). In der Studie werden die Motto-Ziele zwei anderen Zieltypen gegenübergestellt, welche im Selbstmanagementbereich zur Erzeugung von Motivation eingesetzt werden: hohe spezifische Ziele nach Locke & Latham (1990, 2007) und Schwelgen in positiver Zielerreichungsphantasien (ein Teil des mentalen Kontrastieren nach Öttingen, 1999). Um die Zieltypen hinsichtlich ihres Stellenwertes für das innerpsychische Zielsystem einordnen zu können, dient als theoretische Grundlage die Theorie der Persönlichkeits-System-Interaktion (PSI-Theorie) von Kuhl (2001). In einem kontrollierten randomisierten Untersuchungsdesign mit 67 Probanden werden die Zieltypen hinsichtlich verschiedener Konstrukte untersucht und verglichen: Willensbahnung, intrinsische Motivation, Selbstregulations- und Affektregulationskompetenz, Handlungskontrolle, implizite und explizite Affektlage, Entschlossenheit/Bindung/Realisierbarkeit des Ziels, Selbst- und Fremdinfiltration, Einfluss auf Blutglukosewert, Verankerung im Unbewussten. Dazu wurden nebst mehreren Fragebögen der Emoscan (Kazén & Kuhl, 2005) und der Panter (Baumann, Kuhl & Kazén, 2005) eingesetzt und der intraindividuelle Glukosewert (Galliot & Baumeister, 2007) erhoben. Die ersten Ergebnisse zeigen auf, dass Motto-Ziele im Vergleich mit den beiden anderen Gruppen signifikant den Optimismus erhöhen und die Affektregulationskompetenz nach Misserfolg verbessern. Des weiteren kann die Hypothese bestätigt werden, dass Motto-Ziele durch ihre Verankerung im Unbewussten nicht überexplizite Kontrolle (wie die hohen spezifischen Ziele) sondern intuitiv umgesetzt werden.

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Though a major illness in modern society, depression is still not completely understood. A number of empirical observations point to the importance of basic cognitive processes as well as personality variables as antecedents of a depressive disorder. In this work it is argued that “state orientation”, a personality style characterized by the inability to actively influence one’s focus of thought, plays an important role in the development of at least some forms of major depressive disorder. In the present work, it is suggested that (1) state-oriented cognitions are equivalent to sustained information processing, that (2) depressed individuals are characterized in particular by state-oriented cognitions related to prior failure experiences, that (3) sustained processing of affective information will interfere with normal executive cognitive functioning in depressed individuals resulting in impairments of normal behavior, and that (4) both sustained information processing and “affective interference” will be associated with specific dysfunctional patterns of brain activity in depressed individuals. In the first chapter of this thesis, theorizing pertaining to “action control” and the relationship between action control and state orientation are reviewed. After having established the potential functional significance of state-oriented cognitions, their possible link to depression is developed by introducing the “degenerated-intention hypothesis”. Afterwards, the role of state orientation in the advent of the depressive state is discussed against the background of the “functional helplessness” model of depression. Next, recent empirical findings related to executive dysfunction associated with state-oriented cognitions in major depressive disorder and related dysfunctional patterns of brain activity are reviewed. By considering evidence from studies on executive functioning, brain imaging, and neurophysiological studies, support is found for a possible frontocingulate dysfunction associated with a state-oriented cognitive style underlying a major depressive disorder. Consistent with the proposed link between depression and state orientation, in the second chapter of the thesis, Studies 1a – 1c demonstrate that subclinically and clinically depressed individuals are specifically characterized by failure-related state orientation. Moreover, the results of Study 2, described in Chapter 3, reveal that sustained processing of affectively valenced information may indeed interfere with subsequent executive cognitive functioning, especially in individuals demonstrating relatively high levels of depression. Finally, in line with the idea that sustained information processing and affective interference will be related to an individual’s level of state orientation and will be reflected in specific patterns of neural activity, Study 3, presented in Chapter 4, provides considerable evidence for disturbed brain function in clinically depressed individuals during processing of affective information as well as subsequent executive cognitive functioning and its relation to state-oriented thought. The current research supports the idea that state orientation, in particular its failure-focused form, is a crucial process involved in the development and maintenance of a depressive disorder. Specifically, the present findings suggest that certain forms of major depressive disorder are associated with sustained processing of affective information and with the resulting affective interference with executive cognitive functioning. Findings further suggest that sustained information processing is experienced by affected individuals as ruminative, state-oriented thought on past aversive experiences, and that both sustained information processing and affective interference are associated with distinct patterns of brain activity, which are related to early stimulus evaluation, conflict monitoring, and conflict resolution. The processes possibly underlying some forms of depression, as proposed in this thesis, comprise what may be called “the spinning mind”, whose important functional significance is to hinder an individual from adaptive behavior by impairing the ability to direct thought. Although state orientation may therefore appear to be maladaptive per se, it may be argued instead that this mode of action control is also an adaptive process as long as critical limits of certain parameters are met and the spinning mind is prevented. These and similar considerations are addressed in the concluding discussion in Chapter 5.