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Researchers have made many approaches to study the complexities of the mammalian taste system; however molecular mechanisms of taste processing in the early structures of the central taste pathway remain unclear. More recently the Arc catFISH (cellular compartment analysis of temporal activity by fluorescent in situ hybridisation) method has been used in our lab to study neural activation following taste stimulation in the first central structure in the taste pathway, the nucleus of the solitary tract. This method uses the immediate early gene Arc as a neural activity marker to identify taste-responsive neurons. Arc plays a critical role in memory formation and is necessary for conditioned taste aversion memory formation. In the nucleus of the solitary tract only bitter taste stimulation resulted in increased Arc expression, however this did not occur following stimulation with tastants of any other taste quality. The primary target for gustatory NTS neurons is the parabrachial nucleus (PbN) and, like Arc, the PbN plays an important role in conditioned taste aversion learning.
The aim of this thesis is to investigate Arc expression in the PbN following taste stimulation to elucidate the molecular identity and function of Arc expressing, taste- responsive neurons. Naïve and taste-conditioned mice were stimulated with tastants from each of the five basic taste qualities (sweet, salty, sour, umami, and bitter), with additional bitter compounds included for comparison. The expression patterns of Arc and marker genes were analysed using in situ hybridisation (ISH). The Arc catFISH method was used to observe taste-responsive neurons following each taste stimulation. A double fluorescent in situ hybridisation protocol was then established to investigate possible neuropeptide genes involved in neural responses to taste stimulation.
The results showed that bitter taste stimulation induces increased Arc expression in the PbN in naïve mice. This was not true for other taste qualities. In mice conditioned to find an umami tastant aversive, subsequent umami taste stimulation resulted in an increase in Arc expression similar to that seen in bitter-stimulated mice. Taste-responsive Arc expression was denser in the lateral PbN than the medial PbN. In mice that received two temporally separated taste stimulations, each stimulation time-point showed a distinct population of Arc-expressing neurons, with only a small population (10 – 18 %) of neurons responding to both stimulations. This suggests that either each stimulation event activates a different population of neurons, or that Arc is marking something other than simple cellular activation, such as long-term cellular changes that do not occur twice within a 25 minute time frame. Investigation using the newly established double-FISH protocol revealed that, of the bitter-responsive Arc expressing neuron population: 16 % co-expressed calcitonin RNA; 17 % co-expressed glucagon-like peptide 1 receptor RNA; 17 % co-expressed hypocretin receptor 1 RNA; 9 % co-expressed gastrin-releasing peptide RNA; and 20 % co-expressed neurotensin RNA. This co-expression with multiple different neuropeptides suggests that bitter-activated Arc expression mediates multiple neural responses to the taste event, such as taste aversion learning, suppression of food intake, increased heart rate, and involves multiple brain structures such as the lateral hypothalamus, amygdala, bed nucleus of the stria terminalis, and the thalamus.
The increase in Arc-expression suggests that bitter taste stimulation, and umami taste stimulation in umami-averse animals, may result in an enhanced state of Arc- dependent synaptic plasticity in the PbN, allowing animals to form taste-relevant memories to these aversive compounds more readily. The results investigating neuropeptide RNA co- expression suggest the amygdala, bed nucleus of the stria terminalis, and thalamus as possible targets for bitter-responsive Arc-expressing PbN neurons.
An exploration of activity and therapist preferences and their predictors in German-speaking samples
(2023)
According to current definitions of evidence-based practice, patients’ preferences play an important role for the psychotherapeutic process and outcomes. However, whereas a significant body of research investigated preferences regarding specific treatments, research on preferred activities or therapist characteristics is rare, investigated heterogeneous aspects with inconclusive results, lacked validated assessment tools, and neglected relevant preferences, their predictors as well as the perspective of mental health professionals. Therefore, the three studies of this dissertation aimed to address the most fundamental drawbacks in current preference research by providing a validated questionnaire, focus efforts on activity and therapist preferences and add preferences of psychotherapy trainees. To this end, Paper I reports the translation and validation of the 18-item Cooper-Norcross Inventory of Preference (C-NIP) in a broad, heterogeneous sample of N = 969 laypeople, resulting in good to acceptable reliabilities and first evidence of validity. However, the original factor structure was not replicated. Paper II assesses activity preferences of psychotherapists in training using the C-NIP and compares them with the initial laypeople sample. There were significant differences between both samples, with trainees preferring a more patient-directed, emotionally intense and confrontational approach than laypeople. CBT trainees preferred a more therapist-directed, present-focused, challenging and less emotional intense approach than psychodynamic or -analytic trainees. Paper III explores therapist preferences and tests predictors for specific preference choices. For most characteristics, more than half of the participants did not have specific preferences. Results pointed towards congruency effects (i.e., preference for similar characteristics), especially for members of marginalized groups. The dissertation provides both researchers and practitioners with a validated questionnaire, shows potentially obstructive differences between patients and therapists and underlines the importance of therapist characteristics for marginalized groups, thereby laying the foundation for future applications and implementations in research and practice.
The PhD thesis entitled “Actions through the lens of communicative cues. The influence of verbal cues and emotional cues on action processing and action selection in the second year of life” is based on four studies, which examined the cognitive integration of another person’s communicative cues (i.e., verbal cues, emotional cues) with behavioral cues in 18- and 24-month-olds. In the context of social learning of instrumental actions, it was investigated how the intention-related coherence of either a verbally announced action intention or an emotionally signaled action evaluation with an action demonstration influenced infants’ neuro-cognitive processing (Study I) and selection (Studies II, III, IV) of a novel object-directed action. Developmental research has shown that infants benefit from another’s behavioral cues (e.g., action effect, persistency, selectivity) to infer the underlying goal or intention, respectively, of an observed action (e.g., Cannon & Woodward, 2012; Woodward, 1998). Particularly action effects support infants in distinguishing perceptual action features (e.g., target object identity, movement trajectory, final target object state) from conceptual action features such as goals and intentions. However, less is known about infants’ ability to cognitively integrate another’s behavioral cues with additional action-related communicative cues. There is some evidence showing that in the second year of life, infants selectively imitate a novel action that is verbally (“There!”) or emotionally (positive expression) marked as aligning with the model’s action intention over an action that is verbally (“Whoops!”) or emotionally (negative expression) marked as unintentional (Carpenter, Akhtar, & Tomasello, 1998; Olineck & Poulin-Dubois, 2005, 2009; Repacholi, 2009; Repacholi, Meltzoff, Toub, & Ruba, 2016). Yet, it is currently unclear which role the specific intention-related coherence of a communicative cue with a behavioral cue plays in infants’ action processing and action selection that is, whether the communicative cue confirms, contrasts, clarifies, or is unrelated to the behavioral cue. Notably, by using both verbal cues and emotional cues, we examined not only two domains of communicative cues but also two qualitatively distinct relations between behavioral cues on the one hand and communicative cues on the other hand. More specifically, a verbal cue has the potential to communicate an action intention in the absence of an action demonstration and thus a prior-intention (Searle, 1983), whereas an emotional cue evaluates an ongoing or past action demonstration and thus signals an intention-in-action (Searle, 1983). In a first research focus, this thesis examined infants’ capacity to cognitively integrate another’s intention-related communicative cues and behavioral cues, and also focused on the role of the social cues’ coherence in infants’ action processing and action selection. In a second research focus, and to gain more elaborate insights into how the sub-processes of social learning (attention, encoding, response; cf. Bandura, 1977) are involved in this coherence-sensitive integrative processing, we employed a multi-measures approach. More specifically, we used Electroencephalography (EEG) and looking times to examine how the cues’ coherence influenced the compound of attention and encoding, and imitation (including latencies to first-touch and first-action) to address the compound of encoding and response. Based on the action-reconstruction account (Csibra, 2007), we predicted that infants use extra-motor information (i.e., communicative cues) together with behavioral cues to reconstruct another’s action intention. Accordingly, we expected infants to possess a flexibly organized internal action hierarchy, which they adapt according to the cues’ coherence that is, according to what they inferred to be the overarching action goal. More specifically, in a social-learning situation that comprised an adult model, who demonstrated an action on a novel object that offered two actions, we expected the demonstrated action to lead infants’ action hierarchy when the communicative (i.e., verbal, emotional) cue conveyed similar (confirming coherence) or no additional (un-related coherence) intention-related information relative to the behavioral cue. In terms of action selection, this action hierarchy should become evident in a selective imitation of the demonstrated action. However, when the communicative cue questioned (contrasting coherence) the behaviorally implied action goal or was the only cue conveying meaningful intention-related information (clarifying coherence), the verbally/emotionally intended action should ascend infants’ action hierarchy. Consequently, infants’ action selection should align with the verbally/emotionally intended action (goal emulation). Notably, these predictions oppose the direct-matching perspective (Rizzolatti & Craighero, 2004), according to which the observation of another’s action directly resonates with the observer’s motor repertoire, with this motor resonance enabling the identification of the underlying action goal. Importantly, the direct-matching perspective predicts a rather inflexible action hierarchy inasmuch as the process of goal identification should solely rely on the behavioral cue, irrespective of the behavioral cue’s coherence with extra-motor intention-related information, as it may be conveyed via communicative cues. As to the role of verbal cues, Study I used EEG to examine the influence of a confirming (Congruent) versus contrasting (Incongruent) coherence of a verbal action intention with the same action demonstration on 18-month-olds’ conceptual action processing (as measured via mid-latency mean negative ERP amplitude) and motor activation (as measured via central mu-frequency band power). The action was demonstrated on a novel object that offered two action alternatives from a neutral position. We expected mid-latency ERP negativity to be enhanced in Incongruent compared to Congruent, because past EEG research has demonstrated enhanced conceptual processing for stimuli that mismatched rather than matched the semantic context (Friedrich & Friederici, 2010; Kaduk et al., 2016). Regarding motor activation, Csibra (2007) posited that the identification of a clear action goal constitutes a crucial basis for motor activation to occur. We therefore predicted reduced mu power (indicating enhanced motor activation) for Congruent than Incongruent, because in Congruent, the cues’ match provides unequivocal information about the model’s action goal, whereas in Incongruent, the conflict may render the model’s action goal more unclear. Unexpectedly, in the entire sample, 18-month-olds’ mid-latency ERP negativity during the observation of the same action demonstration did not differ significantly depending on whether this action was congruent or incongruent with the model’s verbal action intention. Yet, post hoc analyses revealed the presence of two subgroups of infants, each of which exhibited significantly different mid-latency ERP negativity for Congruent versus Incongruent, but in opposing directions. The subgroups differed in their productive action-related language skills, with the linguistically more advanced infants exhibiting the expected response pattern of enhanced ERP mean negativity in Incongruent than Congruent, indicating enhanced conceptual processing of an action demonstration that was contrasted rather than confirmed by the verbal action context. As expected, central mu power in the entire sample was reduced in Congruent relative to Incongruent, indicating enhanced motor activation when the action demonstration was preceded by a confirming relative to a contrasting verbal action intention. This finding may indicate the covert preparation for a preferential imitation of the congruent relative to the incongruent action (Filippi et al., 2016; Frey & Gerry, 2006). Overall, these findings are in line with the action-reconstruction account (Csibra, 2007), because they suggest a coherence-sensitive attention to and encoding of the same perceptual features of another’s behavior and thus a cognitive integration of intention-related verbal cues and behavioral cues. Yet, because the subgroup constellation in infants’ ERPs was only discovered post hoc, future research is clearly required to substantiate this finding. Also, future research should validate our interpretation that enhanced motor activation may reflect an electrophysiological marker of subsequent imitation by employing EEG and imitation in a within-subjects design. Study II built on Study I by investigating the impact of coherence of a verbal cue and a behavioral cue on 18- and 24-month-olds’ action selection in an imitation study. When infants of both age groups observed a confirming (Congruent) or unrelated (Pseudo-word: action demonstration was associated with novel verb-like cue) coherence, they selectively imitated the demonstrated action over the not demonstrated, alternative action, with no difference between these two conditions. These findings suggest that, as expected, infants’ action hierarchy was led by the demonstrated action when the verbal cue provided similar (Congruent) or no additional (Pseudo-word) intention-related information relative to a meaningful behavioral cue. These findings support the above-mentioned interpretation that enhanced motor activation during action observation may reflect a covert preparation for imitation (Study I). Interestingly, infants did not seem to benefit from the intention-highlighting effect of the verbal cue in Congruent, suggesting that the verbal cue had an unspecific (e.g., attention-guiding) effect on infants’ action selection. Contrary, when infants observed a contrasting (Incongruent) or clarifying (Failed-attempt: model failed to manipulate the object but verbally announced a certain action intention) coherence, their action selection varied with age and also varied across the course of the experiment (block 1 vs. block 2). More specifically, the 24-month-olds made stronger use of the verbal cue for their action selection in block 1 than did the 18-month-olds. However, while the 18-month-olds’ use of the verbal cue increased across blocks, particularly in Incongruent, the 24-month-olds’ use of the verbal cue decreased across blocks. Overall, these results suggest that, as expected, infants’ action hierarchy in Incongruent (both age groups) and Failed-attempt (only 24-month-olds) drew on the verbal action intention, because in both age groups, infants emulated the verbal intention about as often as they imitated the demonstrated action or even emulated the verbal action intention preferentially. Yet, these findings were confined to certain blocks. It may be argued that the younger age group had a harder time inferring and emulating the intended, yet never observed action, because this requirement is more demanding in cognitive and motor terms. These demands may explain why the 18-month-olds needed some time to take account of the verbal action intention. Contrary, it seems that the 24-month-olds, although demonstrating their principle capacity to take account of the verbal cue in block 1, lost trust in the model’s verbal cue, maybe because the verbal cue did not have predictive value for the model’s actual behavior. Supporting this interpretation, research on selective trust has demonstrated that already infants evaluate another’s reliability or competence, respectively, based on how that model handles familiar objects (behavioral reliability) or labels familiar objects (verbal reliability; for reviews, see Mills, 2013; Poulin-Dubois & Brosseau-Liard, 2016). Relatedly, imitation research has demonstrated that the interpersonal aspects of a social-learning situation gain increasing relevance for infants during the second year of life (Gellén & Buttelmann, 2019; Matheson, Moore, & Akhtar, 2013; Uzgiris, 1981). It may thus be argued that when the 24-month-olds were repeatedly faced with a verbally unreliable model, they de-evaluated the verbal cue as signaling the model’s action intention and instead relied more heavily on alternative cues such as the behavioral cue (Incongruent) or the action context (e.g., object affordances, salience; Failed-attempt). Infants’ first-action latencies were higher in Incongruent and Failed-attempt than in both Congruent and Pseudo-word, and were also higher in Failed-attempt than in Incongruent. These latency-findings thus indicate that situations involving a meaningful verbal cue that deviated from the behavioral cue are cognitively more demanding, resulting in a delayed initiation of a behavioral response. In sum, the findings of Study II suggest that both age groups were highly flexible in their integration of a verbal cue and behavioral cue. Moreover, our results do not indicate a general superiority of either cue. Instead, it seems to depend on the informational gain conveyed by the verbal cue whether it exerts a specific, intention-highlighting effect (Incongruent, Failed-attempt) or an unspecific (e.g., attention-guiding) effect (Congruent, Pseudo-word). Studies III and IV investigated the impact of another’s action-related emotional cues on 18-month-olds’ action selection. In Study III, infants observed a model, who demonstrated two actions on a novel object in direct succession, and who combined one of the two actions with a positive (happy) emotional expression and the other action with a negative (sad) emotional expression. As expected, infants imitated the positively emoted (PE) action more often than the negatively emoted (NE) action. This preference arose from an increase in infants’ readiness to perform the PE action from the baseline period (prior to the action demonstrations) to the test period (following the action demonstrations), rather than from a decrease in readiness to the perform the NE action. The positive cue thus had a stronger behavior-regulating effect than the negative cue. Notably, infants’ more general object-directed behavior in terms of first-touch latencies remained unaffected by the emotional cues’ valence, indicating that infants had linked the emotional cues specifically to the corresponding action and not the object as a whole (Repacholi, 2009). Also, infants’ looking times during the action demonstration did not differ significantly as a function of emotional valence and were characterized by a predominant attentional focus to the action/object rather than to the model’s face. Together with the findings on infants’ first-touch latencies, these results indicate a sensitivity for the notion that emotions can have very specific referents (referential specificity; Martin, Maza, McGrath, & Phelps, 2014). Together, Study III provided evidence for selective imitation based on another’s intention-related (particularly positive) emotional cues in an action-selection task, and thus indicates that infants’ action hierarchy flexibly responds to another’s emotional evaluation of observed actions. According to Repacholi (2009), we suggest that infants used the model’s emotional evaluation to re-appraise the corresponding action (effect), for instance in terms of desirability. Study IV followed up on Study III by investigating the role of the negative emotional cue for infants’ action selection in more detail. Specifically, we investigated whether a contrasting (negative) emotional cue alone would be sufficient to differentially rank the two actions along infants’ action hierarchy or whether instead infants require direct information about the model’s action intention (in the form of a confirming action-emotion pair) to align their action selection with the emotional cues. Also, we examined whether the absence of a direct behavior-regulating effect of the negative cue in Study III was due to the negative cue itself or to the concurrently available positive cue masking the negative cue’s potential effect. To this end, we split the demonstration of the two action-emotion pairs across two trials. In each trial, one action was thus demonstrated and emoted (PE, NE action), and one action was not demonstrated and un-emoted (UE action). For trial 1, we predicted that infants, who observed a PE action demonstration, would selectively imitate the PE action, whereas infants, who observed a NE action demonstration would selectively emulate the UE action. As to trial 2, we expected the complementary action-emotion pair to provide additional clarifying information as the model’s emotional evaluation of both actions, which should either lead to adaptive perseveration (if infants’ action selection in trial 1 had already drawn on the emotional cue) or adaptive change (if infants’ action selection in trial 1 signaled a disregard of the emotional cue). As to trial 1, our findings revealed that, as expected, infants imitated the PE action more often than they emulated the UE action. Like in Study III, this selectivity arose from an increase in infants’ propensity to perform the PE action from baseline to trial 1. Also like in Study III, infants performed the NE action about equally often in baseline and trial 1, which speaks against a direct behavior-regulating effect of the negative cue also when presented in isolation. However, after a NE action demonstration, infants emulated the UE action more often in trial 1 than in baseline, suggesting an indirect behavior-regulating effect of the negative cue. Yet, this indirect effect did not yield a selective emulation of the UE action, because infants performed both action alternatives about equally often in trial 1. Unexpectedly, infants’ action selection in trial 2 was unaffected by the emotional cue. Instead, infants perseverated their action selection of trial 1 in trial 2, irrespective of whether it was adaptive or non-adaptive with respect to the model’s emotional evaluation of the action. It seems that infants changed their strategy across trials, from an initial adherence to the emotional (particularly positive) cue, towards bringing about a salient action effect (Marcovich & Zelazo, 2009). In sum, Studies III and IV indicate a dynamic interplay of different action-selection strategies, depending on valence and presentation order. Apparently, at least in infancy, action reconstruction as one basis for selective action performance reaches its limits when infants can only draw on indirect intention-related information (i.e., which action should be avoided). Overall, our findings favor the action-reconstruction account (Csibra, 2007), according to which actions are flexibly organized along a hierarchy, depending on inferential processes based on extra-motor intention-related information. At the same time, the findings question the direct-matching hypothesis (Rizzolatti & Craighero, 2004), according to which the identification (and pursuit) of action goals hinges on a direct simulation of another’s behavioral cues. Based on the studies’ findings, a preliminary working model is introduced, which seeks to integrate the two theoretical accounts by conceptualizing the routes that activation induced by social cues may take to eventually influence an infant’s action selection. Our findings indicate that it is useful to strive a differentiated conceptualization of communicative cues, because they seem to operate at different places within the process of cue integration, depending on their potential to convey direct intention-related information. Moreover, we suggest that there is bidirectional exchange within each compound of adjacent sub-processes (i.e., between attention and encoding, and encoding and response), and between the compounds. Hence, our findings highlight the benefits of a multi-measures approach when studying the development of infants’ social-cognitive abilities, because it provides a more comprehensive picture how the concerted use of social cues from different domains influences infants’ processing and selection of instrumental actions. Finally, this thesis points to potential future directions to substantiate our current interpretation of the findings.. Moreover, an extension to additional kinds of coherence is suggested to get closer to infants’ everyday-world of experience.
Recognizing, understanding, and responding to quantities are considerable skills for human beings. We can easily communicate quantities, and we are extremely efficient in adapting our behavior to numerical related tasks. One usual task is to compare quantities. We also use symbols like digits in numerical-related tasks. To solve tasks including digits, we must to rely on our previously learned internal number representations.
This thesis elaborates on the process of number comparison with the use of noisy mental representations of numbers, the interaction of number and size representations and how we use mental number representations strategically. For this, three studies were carried out.
In the first study, participants had to decide which of two presented digits was numerically larger. They had to respond with a saccade in the direction of the anticipated answer. Using only a small set of meaningfully interpretable parameters, a variant of random walk models is described that accounts for response time, error rate, and variance of response time for the full matrix of 72 digit pairs. In addition, the used random walk model predicts a numerical distance effect even for error response times and this effect clearly occurs in the observed data. In relation to corresponding correct answers error responses were systematically faster. However, different from standard assumptions often made in random walk models, this account required that the distributions of step sizes of the induced random walks be asymmetric to account for this asymmetry between correct and incorrect responses.
Furthermore, the presented model provides a well-defined framework to investigate the nature and scale (e.g., linear vs. logarithmic) of the mapping of numerical magnitude onto its internal representation. In comparison of the fits of proposed models with linear and logarithmic mapping, the logarithmic mapping is suggested to be prioritized.
Finally, we discuss how our findings can help interpret complex findings (e.g., conflicting speed vs. accuracy trends) in applied studies that use number comparison as a well-established diagnostic tool. Furthermore, a novel oculomotoric effect is reported, namely the saccadic overschoot effect. The participants responded by saccadic eye movements and the amplitude of these saccadic responses decreases with numerical distance.
For the second study, an experimental design was developed that allows us to apply the signal detection theory to a task where participants had to decide whether a presented digit was physically smaller or larger. A remaining question is, whether the benefit in (numerical magnitude – physical size) congruent conditions is related to a better perception than in incongruent conditions. Alternatively, the number-size congruency effect is mediated by response biases due to numbers magnitude. The signal detection theory is a perfect tool to distinguish between these two alternatives. It describes two parameters, namely sensitivity and response bias. Changes in the sensitivity are related to the actual task performance due to real differences in perception processes whereas changes in the response bias simply reflect strategic implications as a stronger preparation (activation) of an anticipated answer. Our results clearly demonstrate that the number-size congruency effect cannot be reduced to mere response bias effects, and that genuine sensitivity gains for congruent number-size pairings contribute to the number-size congruency effect.
Third, participants had to perform a SNARC task – deciding whether a presented digit was odd or even. Local transition probability of irrelevant attributes (magnitude) was varied while local transition probability of relevant attributes (parity) and global probability occurrence of each stimulus were kept constantly. Participants were quite sensitive in recognizing the underlying local transition probability of irrelevant attributes. A gain in performance was observed for actual repetitions of the irrelevant attribute in relation to changes of the irrelevant attribute in high repetition conditions compared to low repetition conditions. One interpretation of these findings is that information about the irrelevant attribute (magnitude) in the previous trial is used as an informative precue, so that participants can prepare early processing stages in the current trial, with the corresponding benefits and costs typical of standard cueing studies.
Finally, the results reported in this thesis are discussed in relation to recent studies in numerical cognition.