Climate change and habitat fragmentation modify the natural habitat of many wetland biota and lead to new compositions of biodiversity in these ecosystems. While the direct effects of climate are often well known, indirect effects due to biotic interactions remain poorly understood. The water meadow grasshopper, Chorthippus montanus, is a univoltine habitat specialist, which is adapted to permanently moist habitats. Land use change and drainage led to highly fragmented populations of this generally flightless species. In large parts of the Palaearctic Ch. montanus occurs sympatrically with its widespread congener, the meadow grasshopper Chorthippus parallelus. Due to their close relationship and their similar songs, hybridization is likely to occur in syntopic populations. Such a species pair of a habitat specialist and a habitat generalist represents an ideal model system to examine the role of ongoing climate change and an accumulation of extreme climatic events on the life history strategies, population dynamics and inter-specific interactions. In Chapter I a laboratory experiment was conducted to identify the impact of environmental factors on intra-specific life-history traits of Ch. montanus. Like other Orthoptera species, Ch. montanus follows a converse temperature size rule. In line with the dimorphic niche hypothesis, which states that sexual size dimorphism evolved in response to the different sexual reproductive roles, both sexes showed different responses to increasing density at lower temperatures. Males attained smaller body sizes at high densities, whereas females had a prolonged development time. This is the first evidence for a sex-specific phenotypic plasticity in Ch. montanus. Females benefit from the prolonged development as their reproductive success depends on the size and number of egg clutches they may produce. By contrast, the reproductive success of males depends on the chance to fertilize virgin females, which increases with faster development. This may become a disadvantage for Ch. montanus as an intraspecific phenology shift may increase hybridization risk with the sibling species. Despite the widespread assumption that hybridization between two sympatric species is rare due to complete reproductive barriers, the genetic analyses of 16 populations (Chapter II) provided evidence for wide prevalence of hybridization between both species in the wild. As no complete admixture was found in the examined population, it is assumed that hybridization only occurs in ecotones between wetlands and drier parts. Reproductive barriers (habitat isolation, behavior, phenology) seem to prevent the genetic swamping of Ch. montanus populations. Although a behavioral experiment showed that mate choice presents an important reproductive barrier between both species, the experiment also revealed that reproductive barriers could be altered by environmental change (e.g. increasing heterospecific frequency). Chapter III analyzes the impact of extreme climatic events on population dynamics and interspecific hybridization. A mark-recapture analysis combined with weather records over five years provides evidence that the embryonic development in Ch. montanus is vulnerable to extreme climatic events. Strong population declines in Ch. montanus lead to a disequilibrium between Ch. montanus and Ch. parallelus populations and increases the risk of hybridization. The highest hybridization risk was found in the first weeks of a season, when both species had an overlapping phenology. Furthermore, hybrids were generally localized at the edge of the Ch. montanus distribution with higher heterospecific encounter probabilities. The hybridization rate reached up to 19.6%. The genetic analyses in Chapter II and III show that hybridization differentially affects specialists and generalists. While generalists may benefit from hybridization by an increasing genetic diversity, such a positive correlation was not found for Ch. montanus. The results underline the importance of reproductive barriers for the co-existence of these sympatric species. However, climate change and other anthropogenic disturbances alter reproductive barriers and promote hybridization, which may threaten small populations by genetic displacement. As anthropogenic hybridization is recognized as a major threat to biodiversity, it should be considered in environmental law and policy. In Chapter IV the role of hybrids and hybridization in three levels of law and the historical backgrounds of hybrids becoming a part of legal instruments is analyzed. Due to legal uncertainties and the complexity of this topic a legal assessment of hybrids is challenging and argues for species-specific approaches. Nonetheless, existing legal norms provide a suitable basis, but need to be specified. Finally, this chapter discusses different opportunities for the management of hybrids and hybridization in a conservation perspective and their necessity.
The startle response in psychophysiological research: modulating effects of contextual parameters
(2013)
Startle reactions are fast, reflexive, and defensive responses which protect the body from injury in the face of imminent danger. The underlying reflex is basic and can be found in many species. Even though it consists of only a few synapses located in the brain stem, the startle reflex offers a valuable research method for human affective, cognitive, and psychological research. This is because of moderating effects of higher mental processes such as attention and emotion on the response magnitude: affective foreground stimulation and directed attention are validated paradigms in startle-related research. This work presents findings from three independent research studies that deal with (1) the application of the established "affective modulation of startle"-paradigm to the novel setting of attractiveness and human mating preferences, (2) the question of how different components of the startle response are affected by a physiological stressor and (3) how startle stimuli affect visual attention towards emotional stimuli. While the first two studies treat the startle response as a dependent variable by measuring its response magnitude, the third study uses startle stimuli as an experimental manipulation and investigates its potential effects on a behavioural measure. The first chapter of this thesis describes the basic mechanisms of the startle response as well as the body of research that sets the foundation of startle research in psychophysiology. It provides the rationale for the presented studies, and offers a short summary of the obtained results. Chapter two to four represent primary research articles that are published or in press. At the beginning of each chapter the contribution of all authors is explained. The references for all chapters are listed at the end of this thesis. The overall scope of this thesis is to show how the human startle response is modulated by a variety of factors, such as the attractiveness of a potential mating partner or the exposure to a stressor. In conclusion, the magnitude of the startle response can serve as a measure for such psychological states and processes. Beyond the involuntary, physiological startle reflex, startle stimuli also affect intentional behavioural responses, which we could demonstrate for eye movements in a visual attention paradigm.
Stress is a common phenomenon for animals living in the wild, but also for humans in modern societies. Originally, the body's stress response is an adaptive reaction to a possibly life-threatening situation, and it has been shown to impact on energy distribution and metabolism, thereby increasing the chance of survival. However, stress has also been shown to impact on mating behaviour and reproductive strategies in animals and humans. This work deals with the effect of stress on reproductive behavior. Up to now, research has only focused on the effects of stress on reproduction in general. The effects of stress on reproduction may be looked at from two points of view. First, stress affects reproductive functioning by endocrine (e.g. glucocorticoid) actions on the reproductive system. However, stress can also influence reproductive behavior, i.e. mate choice and mating preferences. Animals and humans do not mate randomly, but exhibit preferences towards mating partners. One factor by which animals and humans choose their mating partners is similarity vs. dissimilarity: Similar mates usually carry more of one's own genes and the cooperation between similar mates is, at least theoretically, less hampered by expressing diverse behaviors. By mating with dissimilar mates on the other hand one may acquire new qualities for oneself, but also for one's offspring, useful to cope with environmental challenge. In humans we usually find a preference for similar mates. Due to the high costs of breeding, variables like cooperation and life-long partnerships may play a greater role than the acquaintance of new qualities.The present work focuses on stress effects on mating preferences of humans and will give a first answer to the question whether stress may affect our preference for similar mates. Stress and mating preferences are at the centre of this work. Thus, in the first Chapter I will give an introduction on stress and mating preferences and link these topics to each other. Furthermore, I will give a short summary of the studies described in Chapter II - Chapter IV and close the chapter with a general discussion of the findings and directions for further research on stress and mating preferences. Human mating behavior is complex, and many aspects of it may not relate to biology but social conventions and education. This work will not focus on those aspects but rather on cognitive and affective processing of erotic and sexually-relevant stimuli, since we assume that these aspects of mating behaviour are likely related to psychobiological stress mechanisms. Therefore, a paradigm is needed that measures such aspects of mating preferences in humans. The studies presented in Chapter II and Chapter III were performed in order to develop such a paradigm. In these studies we show that affective startle modulation may be used to indicate differences in sexual approach motivation to potential mating partners with different similarity levels to the participant. In Chapter IV, I will describe a study that aimed to investigate the effects of stress on human mating preferences. We showed that stress reverses human mating preferences: While unstressed individuals show a preference for similar mates, stressed individuals seem to prefer dissimilar mates. Overall, the studies presented in this work showed that affective startle modulation can be employed to measure mating preferences in humans and that these mating preferences are influenced by stress.