A lack of empathy is considered to be a hallmark feature of psychopathy. However, remarkably little research has specifically examined the relationship between psychopathy and empathy, and what does exist relies solely on self-report measures, which is problematic given psychopaths' propensity for dishonesty. This research signifies an attempt to develop an objective measure of empathy, using psychophysiological indices of emotional experience as indicators of an empathic response. Specifically, corrugator and zygomatic facial muscle activity, event-related heart rate, and acoustic startle reflex responses were measured while participants viewed brief films of individuals experiencing happy and sad emotional events.Pilot investigations of the films suggest they are appropriate for eliciting happy and sad empathic emotion, and the pattern of responses by undergraduates with high and low trait levels of empathy indicates the validity of the approach, and the promise of utilizing psychophysiological indices of emotional responding to measure situational empathy in an objective manner. In particular, participants demonstrated patterns of psychophysiological responses to the empathy-inducing films which both matched their self-reported emotional experiences as well as the emotional valence of the stimuli. Moreover, several of the psychophysiological indices were able to discriminate high and low empathy individuals, whereby low empathy participants exhibited less facial expressiveness and less modulation of the acoustic startle reflex to the empathy-inducing stimuli than their high empathy counterparts.The paradigm was then extended to a sample of male undergraduates with high and low levels of psychopathic traits, as assessed by the Psychopathic Personality Inventory-Revised. Results suggest a significant negative relationship between self-reported psychopathy and self-reported dispositional empathy, with high levels of psychopathy associated with low levels of dispositional empathy, particularly with respect to affective empathy. However, despite differences in dispositional empathy, psychopathic and non-psychopathic undergraduates were not easily discriminated on subjective or objective measures of situational empathy, with the exception of an anomalous pattern of zygomatic EMG activity exhibited by the psychopathic participants. The implications of these findings are discussed with respect to the nature of the empathic capabilities of psychopaths, as well as considerations of the methodological limitations of the current study and directions for future research.

I denna studie har tio undersökningsdeltagare sett tio klipp från skräckfilmer som alla innehåller jumpscare-scener, det vill säga scener med plötsliga förändringar som används för att ge tittare en skrämselupplevelse. Undersökningsdeltagarna har för varje klipp fått värdera både i ett bildformulär och i ord hur pass skrämda de blev och vad de tror var anledningarna till det. Jumpscare-scenerna som användartestet bestod av har alla olika komponenter som utmärker dem för att det skulle vara möjligt att se vilka komponenter som bidrog mest till att skrämmas. Begreppen pre-scare, precise-scare och post-scare har utvecklats och använts i undersökningen för att förklara de olika temporala delarna av en jumpscare-scen. Studien utgår bland annat från den så kallade Startle reflexen, som utlöses på grund av plötslig och stark stimuli och audio-visuellinriktad litteratur som exempelvis Chions Audio-Vision. Resultaten visar bland annat att de mest skrämmande jumpscare-scenerna har långa, obehagliga uppbyggnader med lågfrekventa ljud eller musikstycken i prescare- momentet och ett så starkt, oväntat, högfrekvent ljud i precise-scare-momentet som möjligt. De innehåller ofta dissonant och oregelbunden musik och röster samt skrik förstärker även skrämseleffekten eller den så kallade startle-reflexen.

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Defence sounds are widespread and diverse amongst insects. Despite their ubiquity and variability, hypotheses explaining their functions and evolutionary origins have been understudied. My thesis focused on these topics using silk and hawkmoth Bombycoidea caterpillars as a model system. In Chapter Two I investigated why defence sounds have evolved in some caterpillars but not others by testing the hypothesis that large body size is a factor in the evolution of defence sounds. To test this hypothesis, I followed the development of defence sounds in four Bombycoidea species from hatching to pupation. I predicted that early instars would not produce defence sounds, and that within sound producing instars defence sounds would be more likely to occur in larger caterpillars. Results showed that defence sounds were absent in the first and second instar, and that they developed in the third through to the fifth instar in all species. Moreover, the onset of sound production occurred when all species were the same relative size (~1.12 g, ~26.37 mm), despite the fact that the species differed in their final instar size. I concluded that early instar caterpillars do not make defence sounds, and that there is a critical size when defence sounds develop. I further tested the hypothesis that smaller caterpillars do not have enough energy to make defence sounds, by analyzing the relationship between size and several temporal characteristics of the sounds. I predicted that smaller caterpillars would signal less than larger caterpillars, and produce shorter signal units and trains, with lower duty cycles. Results partly supported the hypothesis, showing that in two species there was a positive relationship between size and the number of units produced within two seconds following an attack, the mean number of units per train, and the mean duration of the units in one species. I also tested the hypothesis that sounds of small caterpillars are not in the hearing range of predators. I predicted that there would be a relationship between caterpillar size, and the sound pressure levels and dominant frequencies of the sounds. Results showed no significant relationships with dominant frequencies or sound pressure levels and size. I concluded that the caterpillars made sounds that were within the hearing range of major predators from the onset of sound production. In Chapter Three I followed the other antipredator defences of the four species throughout development. I investigated whether the frequency of defences changed with instar. I found that the caterpillars employed up to seven different secondary defences throughout development. In one species the frequency of dropping and major thrashing increased in the late instars, and in a different species the frequency of regurgitation increased. I concluded that in some cases defence sound production accompanies other secondary defences that increase with the size of caterpillars during development. In Chapter Four I tested the hypothesis that the defensive whistle of the walnut sphinx caterpillar, Amorpha juglandis (Sphingidae: Sphinginae), functions to startle birds. I predicted that the birds would startle to the sounds, and habituate upon repeated exposure within a trial. Results showed that play-back recordings of the whistles elicited a startle response in captive red-winged blackbirds (Agelaius phoeniceus) and caused them to hesitate and/or flee from prey. I concluded that the whistles function as a startle display. Together, the experiments conducted within my thesis addressed important outstanding questions regarding the evolutionary origins of defence sounds in caterpillars, and their functions in predator-prey interactions.

Decades of research in quantitative genetics have found substantial heritability for personality traits as well as for mental disorders which formed the basis of the ongoing molecular genetic studies that aim to identify genetic variations that actually contribute to the manifestation of complex traits. With regard to psychological traits, genetic variation impacting neurotransmitter function have been of particular interest. Additionally, the role of environmental factors including gene × environment interactions has been further investigated and the impor-tance of developmental aspects has been stressed. Furthermore, endophenotypes which link complex traits with their respective biological underpinnings and thus bridge the gap between gene and behaviour have begun to be included in research efforts. In accordance with this approach, this thesis aims to further examine the influence of genetic variation impacting serotonergic and dopaminergic functioning on endophenotypes of anxiety-related behaviour. To this end, two well established paradigms – the acoustic startle reflex and the cortisol stress response – were employed. Both show considerable interindividual variation which has been found in quantitative genetic studies to be at least partly based on genetic factors. In addition, the neural circuits underlying these endophenotypes are relatively well understood and thus reveal references for the detection of associated genetic influences. The results of this thesis associate the overall startle magnitude in two independent samples of young adults with a polymorphism in the promoter region of the serotonin transporter (5-HTT) gene (5-HTTLPR): Carriers of the short (S) allele which results in a reduced gene ex-pression showed a stronger startle magnitude which is in line with numerous findings linking the S allele to increased measures of negative emotionality. In addition to 5-HTTLPR, the effects of past stressful life events on the startle response were investigated: Participants who had recently experienced at least one stressful life event exhibited stronger startle responses and reduced habituation of the startle reflex although there was no 5-HTTLPR × environment inter-action effect. A third study revealed independent and joint effects of 5-HTTLPR and a poly-morphism in the dopamine receptor 4 gene (DRD4) in the same sample of young adults with regard to the cortisol stress response with carriers of the DRD4 7R allele which has been associ-ated with higher scores in sensation seeking, showing reduced cortisol responses. In addition, a 5-HTTLPR × DRD4 interaction effect emerged: 5-HTTLPR long (L) allele carriers showed the lowest cortisol response but only when they possessed at least one copy of the DRD4 7R allele. Moreover, in a fourth study a life span approach was taken and the influence of a further important serotonergic polymorphism which impacts the functioning of tryptophan hydroxylase 2 (TPH2), the rate limiting enzyme in the biosynthesis of serotonin, on interindividual differences in the startle response was investigated in three different age samples: children, young adults and older adults. There was a sex × TPH2 genotype interaction effect in a sample of young adults on the overall startle response while there was no effect of TPH2 in children or older adults. The last study of this thesis presents findings regarding the influence of two dopaminergic polymorphisms in genes encoding the enzyme catechol-O-methyltransferase (COMT) and the dopamine transporter (DAT), respectively, which both terminate dopamine signalling and are thus important regulators of dopaminergic neurotransmission, on the startle reflex in older adults. COMT met/met homozygotes showed the strongest and val/val homozygotes displayed the smallest startle magnitude which is in line with findings linking the COMT met allele to increased scores of anxiety related traits and disorders. Regarding DAT, participants homozygous for the 10R allele, which had previously associated with attention-deficit hyperactivity disorder, showed a stronger overall startle response. In sum, this thesis comprises data on interindividual differences in an electrophysiological and a hormonal endophenotype across the life span and their association with serotonergic and dopaminergic function based on genetic variation. One major finding is the clear evidence for the influence of serotonergic polymorphisms on the startle response in young adults while in contrast in older adults genetic variation in the dopaminergic system exerted considerable influence. These differences might be due to developmental processes in the different stages of life although cohort effects and effects of different recruitment strategies can also not be ruled out. Furthermore, there were significant differences regarding the genetic influence on the acoustic startle reflex and cortisol stress response in one and the same sample which might be due to methodological differences of the two paradigms as well as differences in their underlying neuronal circuits. In conclusion, this thesis supports the acoustic startle reflex and the cortisol stress response as valuable endophenotypes and thus indicators for underlying neurobiological circuits although some methodological issues remain. It also highlights the importance of taking developmental factors and changes over the course of life into account. Finally, this thesis emphasizes the necessity to include reliably and validly assessed past experienced events in molecular genetic studies in order to understand the interplay between genetic and environmental factors in shaping (endo)-phenotypes.

Decades of research in quantitative genetics have found substantial heritability for personality traits as well as for mental disorders which formed the basis of the ongoing molecular genetic studies that aim to identify genetic variations that actually contribute to the manifestation of complex traits. With regard to psychological traits, genetic variation impacting neurotransmitter function have been of particular interest. Additionally, the role of environmental factors including gene × environment interactions has been further investigated and the impor-tance of developmental aspects has been stressed. Furthermore, endophenotypes which link complex traits with their respective biological underpinnings and thus bridge the gap between gene and behaviour have begun to be included in research efforts. In accordance with this approach, this thesis aims to further examine the influence of genetic variation impacting serotonergic and dopaminergic functioning on endophenotypes of anxiety-related behaviour. To this end, two well established paradigms – the acoustic startle reflex and the cortisol stress response – were employed. Both show considerable interindividual variation which has been found in quantitative genetic studies to be at least partly based on genetic factors. In addition, the neural circuits underlying these endophenotypes are relatively well understood and thus reveal references for the detection of associated genetic influences. The results of this thesis associate the overall startle magnitude in two independent samples of young adults with a polymorphism in the promoter region of the serotonin transporter (5-HTT) gene (5-HTTLPR): Carriers of the short (S) allele which results in a reduced gene ex-pression showed a stronger startle magnitude which is in line with numerous findings linking the S allele to increased measures of negative emotionality. In addition to 5-HTTLPR, the effects of past stressful life events on the startle response were investigated: Participants who had recently experienced at least one stressful life event exhibited stronger startle responses and reduced habituation of the startle reflex although there was no 5-HTTLPR × environment inter-action effect. A third study revealed independent and joint effects of 5-HTTLPR and a poly-morphism in the dopamine receptor 4 gene (DRD4) in the same sample of young adults with regard to the cortisol stress response with carriers of the DRD4 7R allele which has been associ-ated with higher scores in sensation seeking, showing reduced cortisol responses. In addition, a 5-HTTLPR × DRD4 interaction effect emerged: 5-HTTLPR long (L) allele carriers showed the lowest cortisol response but only when they possessed at least one copy of the DRD4 7R allele. Moreover, in a fourth study a life span approach was taken and the influence of a further important serotonergic polymorphism which impacts the functioning of tryptophan hydroxylase 2 (TPH2), the rate limiting enzyme in the biosynthesis of serotonin, on interindividual differences in the startle response was investigated in three different age samples: children, young adults and older adults. There was a sex × TPH2 genotype interaction effect in a sample of young adults on the overall startle response while there was no effect of TPH2 in children or older adults. The last study of this thesis presents findings regarding the influence of two dopaminergic polymorphisms in genes encoding the enzyme catechol-O-methyltransferase (COMT) and the dopamine transporter (DAT), respectively, which both terminate dopamine signalling and are thus important regulators of dopaminergic neurotransmission, on the startle reflex in older adults. COMT met/met homozygotes showed the strongest and val/val homozygotes displayed the smallest startle magnitude which is in line with findings linking the COMT met allele to increased scores of anxiety related traits and disorders. Regarding DAT, participants homozygous for the 10R allele, which had previously associated with attention-deficit hyperactivity disorder, showed a stronger overall startle response. In sum, this thesis comprises data on interindividual differences in an electrophysiological and a hormonal endophenotype across the life span and their association with serotonergic and dopaminergic function based on genetic variation. One major finding is the clear evidence for the influence of serotonergic polymorphisms on the startle response in young adults while in contrast in older adults genetic variation in the dopaminergic system exerted considerable influence. These differences might be due to developmental processes in the different stages of life although cohort effects and effects of different recruitment strategies can also not be ruled out. Furthermore, there were significant differences regarding the genetic influence on the acoustic startle reflex and cortisol stress response in one and the same sample which might be due to methodological differences of the two paradigms as well as differences in their underlying neuronal circuits. In conclusion, this thesis supports the acoustic startle reflex and the cortisol stress response as valuable endophenotypes and thus indicators for underlying neurobiological circuits although some methodological issues remain. It also highlights the importance of taking developmental factors and changes over the course of life into account. Finally, this thesis emphasizes the necessity to include reliably and validly assessed past experienced events in molecular genetic studies in order to understand the interplay between genetic and environmental factors in shaping (endo)-phenotypes.

<p>Bilder av potentiella hot upptäcks snabbt och leder till en snabbare reaktionstid än bilder av icke-potentiella hot. Potentiella hot stjäl också fokus och kognitiva resurser, med en långsammare reaktionstid som följd. Rädsla, som kan mätas genom startlereflexen, leder till antingen ökad handlingsberedskap (snabbare reaktionstid) eller kognitiv distraktion (långsammare reaktionstid). Fem försöksdeltagare fick reagera på en visuell probe som lades ovanpå djurbilder, både potentiella hot och icke-potentiella hot i ett kombinerat reaktionstids- och startleprobeexperiment, där ljudstötar utdelades i hörlurar. Startlereaktion, hjärtfrekvens och reaktionstid mättes, varefter korrelationskoefficienter räknades ut. Resultatet visade på en svag negativ korrelation mellan reaktionstid och startlepotentiering på gruppnivå, men inga signifikanta samband påvisades på individnivå. Resultatet på gruppnivå kan bero på att de individuella resultaten tenderar samlas i kluster och därför skapar ett skensamband. Även en positiv korrelation mellan hjärtfrekvensförändring och startlereaktion påvisades, men ingen korrelation mellan hjärtfrekvens och reaktionstid.</p> / <p>Pictures of fear relevant animals are detected fast and lead to shorter reaction time, compared to pictures of fear irrelevant animals. Fear relevant animals also capture attention and steals focus, which leads to slower reaction time. Fear leads to either preparedness for action (faster reaction time) or cognitive distraction (slower reaction time). In a combined startle probe and reaction time experiment, five participants were requested to react to a visual probe, superimposed on pictures of animals, both relevant and irrelevant to fear. Startle probes were given as noise bursts in headphones. Startle reaction, heart rate and reaction time were measured. A negative correlation between reaction time and startle potentiation was found on group level, whereas no significant correlations were found on the individual level. However, the individual results tend to gather in clusters, which might give a significant result on group level even if there is none.</p>

Bilder av potentiella hot upptäcks snabbt och leder till en snabbare reaktionstid än bilder av icke-potentiella hot. Potentiella hot stjäl också fokus och kognitiva resurser, med en långsammare reaktionstid som följd. Rädsla, som kan mätas genom startlereflexen, leder till antingen ökad handlingsberedskap (snabbare reaktionstid) eller kognitiv distraktion (långsammare reaktionstid). Fem försöksdeltagare fick reagera på en visuell probe som lades ovanpå djurbilder, både potentiella hot och icke-potentiella hot i ett kombinerat reaktionstids- och startleprobeexperiment, där ljudstötar utdelades i hörlurar. Startlereaktion, hjärtfrekvens och reaktionstid mättes, varefter korrelationskoefficienter räknades ut. Resultatet visade på en svag negativ korrelation mellan reaktionstid och startlepotentiering på gruppnivå, men inga signifikanta samband påvisades på individnivå. Resultatet på gruppnivå kan bero på att de individuella resultaten tenderar samlas i kluster och därför skapar ett skensamband. Även en positiv korrelation mellan hjärtfrekvensförändring och startlereaktion påvisades, men ingen korrelation mellan hjärtfrekvens och reaktionstid. / Pictures of fear relevant animals are detected fast and lead to shorter reaction time, compared to pictures of fear irrelevant animals. Fear relevant animals also capture attention and steals focus, which leads to slower reaction time. Fear leads to either preparedness for action (faster reaction time) or cognitive distraction (slower reaction time). In a combined startle probe and reaction time experiment, five participants were requested to react to a visual probe, superimposed on pictures of animals, both relevant and irrelevant to fear. Startle probes were given as noise bursts in headphones. Startle reaction, heart rate and reaction time were measured. A negative correlation between reaction time and startle potentiation was found on group level, whereas no significant correlations were found on the individual level. However, the individual results tend to gather in clusters, which might give a significant result on group level even if there is none.

La musique a la capacité d’induire et moduler les émotions, décomposées en deux dimensions : le niveau d’activation (relaxant-stimulant) et la valence émotionnelle (déplaisant-plaisant). Une façon de mesurer objectivement la valence musicale est par le réflexe acoustique de sursaut, une réaction de défense qui consiste en un clignement de l’oeil provoqué par un bruit fort et inattendu. Le réflexe est renforcé par la musique déplaisante et inhibée par la musique plaisante. Cependant, l’effet du niveau d’activation émotionnelle lors de l’écoute musicale demeure inconnu. Cette étude a donc pour objectif d’examiner la modulation du réflexe acoustique de sursaut par la musique stimulante et relaxante jugée plaisante. Basée sur les résultats d’études antérieures avec des images, notre hypothèse était que le réflexe serait plus faible dans la condition stimulante que dans la condition relaxante. Dans un devis intrasujet, 47 participants ont écouté de la musique relaxante et stimulante. Des bruits blancs courts et forts ont été rajoutés par-dessus les extraits afin de provoquer le réflexe de sursaut, dont son amplitude et sa latence ont été mesurées par électromyographie. Les résultats ont ensuite été comparés à ceux d’une condition non-musicale, constituée de sons environnementaux plaisants, afin d’explorer si la musique est plus efficace pour inhiber le réflexe. Finalement, des caractéristiques acoustiques, telles que la clarté de la pulsation, la densité acoustique, la dissonance et l’énergie, ont été extraites puis comparées entre les trois conditions pour explorer leur relation avec les paramètres du réflexe. Les résultats rapportent une modulation de la latence du réflexe de sursaut, dans laquelle celle-ci est plus longue dans la condition stimulante comparée à la condition relaxante. Cependant, aucune différence au niveau de l’amplitude n’a été observée. Seule la latence serait donc sensible au niveau d’activation des émotions musicales lorsque la musique est plaisante. Ensuite, la latence dans la condition non-musicale était aussi longue que celle dans la condition stimulante, suggérant que la musique n’est pas plus efficace que les sons non-musicaux pour inhiber le réflexe de sursaut. Finalement, comme l’amplitude et la latence n’ont pas le même patron de réponses, cette étude suggère que le réflexe de sursaut est aussi modulé par le traitement des caractéristiques acoustiques et que ceux-ci ont un effet différent sur ces deux paramètres. En conclusion, la latence du réflexe acoustique de sursaut est une bonne méthode pour mesurer le niveau d’activation des émotions musicales. De futures recherches pourront utiliser le paradigme de la modulation affective du réflexe de sursaut pour mesurer les effets des émotions musicales selon des facteurs individuels tels que l’âge et la dépression. / Music has the capacity to evoke and modulate emotions, divided by two dimensions: arousal (relaxing-stimulating), and valence (unpleasant-pleasant). Musical valence can be objectively measured by the acoustic startle reflex, a defensive reaction consisting of an eye blink provoked by a short and loud noise. This reflex is facilitated by unpleasant music and inhibited by pleasant music. However, the arousal effect while listening to music on the startle reflex remains unknown. This study therefore aims to explore the affective startle modulation by stimulating and relaxing music. In a within-subjects design, 47 participants listened to stimulating music, relaxing music and non-musical sounds. White noises (50 ms, 105 dB(A)) were added over the excerpts to induce startle while eyeblink magnitude and latency were measured by electromyography. Excerpts’ acoustic features were then extracted and compared through experimental conditions to explore their effect on startle modulation. Startle latency was longer in the stimulating condition compared to the relaxing one, but no differences in magnitude were found, partially confirming our predictions. Exploratory analyses suggest that startle modulation is also attributed to bottom-up processes of acoustic features, and that these latter impact differently magnitude and latency. In conclusion, this study highlights startle latency measure efficiently emotional arousal while listening to music, allowing future research to use the paradigm of affective startle reflex modulation to evaluate the effect of music on emotions considering individual factors, such as age and depression. It also paves the way for comparisons of the effect of emotions and acoustic features processes on the startle reflex modulation.

No description available.