Genetic and environmental influences on human responses to odors

Olfaction, the sense of smell, has many important functions in humans. Human responses to odors show substantial individual variation. Olfactory receptor genes have been identified and other genes may also influence olfaction. However, the proportion of phenotypic variation in odor response due to genetic variation remains largely unknown. Little is also known about which genes modify specific responses to odors. This study aimed to elucidate genetic and environmental influences on human responses to odors. Individuals from Finnish families (n=146) and Australian (n=413), British (n=163), Danish (n=336), and Finnish (n=399) twins rated intensity and pleasantness of a set of 12 (families) or 6 (twins) odors and tried to identify the odors. In addition, the participants rated their own sense of smell and annoyance experienced with different environmental odors. The odor stimuli of a commercial smell test (The Brief Smell Identification Test; banana, chocolate, cinnamon, gasoline, lemon, onion, paint thinner, pineapple, rose, smoke, soap, and turpentine) were presented in the family study. Based on the results of the family study and a literature survey, a new set of odor stimuli (androstenone, chocolate, cinnamon, isovaleric acid, lemon, and turpentine) was designed for the twin studies. In the family sample, heritabilities of the traits were estimated and underlying genomic regions were searched using a genome-wide linkage scan. In the pooled twin sample, variation in the measured traits was decomposed into genetic and environmental components using quantitative genetic modeling. In addition, associations between nongenetic factors (e.g., sex, age, and smoking) and olfactory-related traits were explored. Suggestive evidence for a genetic linkage for pleasantness of cinnamon at a locus on chromosome 4q32.3 emerged from the family sample. High heritability for the pleasantness of cinnamon was found in the family but not the twin study. Heritability of perceived intensity of androstenone odor was determined to be ~30% in the twin sample. A strong genetic correlation between perceived intensity and pleasantness of androstenone, in the absence of any environmental correlation, indicated that only the genetic correlation explained the phenotypic correlation between the traits (r=-0.27) and that the traits were influenced by an overlapping set of genes. Self-rated olfactory function appeared to reflect the odor annoyance experienced rather than actual olfactory acuity or genetic involvement. Results from nongenetic analyses supported the speculated superiority of females' olfactory abilities, the age-related diminishing of olfactory acuity, and the influences of experience-dependent factors on odor responses. This was the first study to estimate heritabilities and perform linkage screens for individual odors. A genetic effect was detected for only a few responses to specific odors, suggesting the predominance of environmental effects in odor perceptions.

Antipredator behaviour of Baltic planktivores

Predation is an important source of mortality for most aquatic animals. Thus, the ability to avoid being eaten brings substantial fitness benefits to individuals. Predator detection abilities and antipredator behaviour were examined in various planktivores, i.e. the littoral mysids Neomysis integer and Praunus flexuosus, three-spined stickleback Gasterosteus aculeatus larvae, pelagic mysids Mysis mixta and M. relicta, and the predatory cladoceran Cercopagis pengoi, with cues from their respective predators European perch Perca fluviatilis and Baltic herring Clupea harengus membras. The use of different aquatic macrophytes as predation refuges by the littoral planktivores was also examined. All pelagic planktivores and stickleback larvae were able to detect the presence of their predator by chemical cues alone. The littoral mysids N. integer and P. flexuosus responded only when chemical and visual predator cues were combined. The responses of stickleback larvae were stronger to the combined cues than the chemical cue alone. A common antipredator behaviour in all of the planktivores studied was decreased ingestion rate in response to predator cues. N. integer and stickleback larvae also decreased their swimming activity. Pelagic mysids and C. pengoi altered their prey selectivity patterns in response to predator cues. The effects of predator cues on the swarming behaviour of N. integer were examined. Swarming brings clear antipredator advantages to N. integer, since when they feed in a swarm, they do not significantly decrease their feeding rate. However, the swarming behaviour of N. integer was not affected by predation risk, but was instead a fixed strategy. Despite the presence or absence of predator cues, N. integer individuals attempted to associate with a swarm and preferred larger to smaller swarms. In studies with aquatic macrophytes, stickleback larvae and P. flexuosus utilized vegetation as a predation refuge, spending more time within vegetation when under predation threat. The two macroalgal species studied, bladderwrack Fucus vesiculosus and stonewort Chara tomentosa, were preferred by P. flexuosus, whereas Eurasian watermilfoil Myriophyllum spicatum was strongly avoided by N. integer and stickleback larvae. In fact, when in dense patches in aquaria, M. spicatum caused acute and high mortality (> 70%) in littoral mysids, but not in sticklebacks, whereas C. tomentosa and northern watermilfoil M. sibiricum did not. In contrast, only 2-4% mortality in N. integer was observed with intact and broken stems of M. spicatum in field experiments. The distribution of littoral mysids in different vegetations, however, suggests that N. integer avoids areas vegetated by M. spicatum.

Invertebrate predation and trophic cascades in a pelagic food web : The multiple roles of Chaoborus flavicans (Meigen) in a clay-turbid lake

In lake ecosystems, both fish and invertebrate predators have dramatic effects on their prey communities. Fish predation selects large cladocerans while invertebrate predators prefer prey of smaller size. Since invertebrate predators are the preferred food items for fish, their occurrence at high densities is often connected with the absence or low number of fish. It is generally believed that invertebrate predators can play a significant role only if the density of planktivorous fish is low. However, in eutrophic clay-turbid Lake Hiidenvesi (southern Finland), a dense population of predatory Chaoborus flavicans larvae coexists with an abundant fish population. The population covers the stratifying area of the lake and attains a maximum population density of 23000 ind. m-2. This thesis aims to clarify the effects of Chaoborus flavicans on the zooplankton community and the environmental factors facilitating the coexistence of fish and invertebrate predators. In the stratifying area of Lake Hiidenvesi, the seasonal succession of cladocerans was exceptional. The spring biomass peak of cladocerans was missing and the highest biomass occurred in midsummer. In early summer, the consumption rate by chaoborids clearly exceeded the production rate of cladocerans and each year the biomass peak of cladocerans coincided with the minimum chaoborid density. In contrast, consumption by fish was very low and each study year cladocerans attained maximum biomass simultaneously with the highest consumption by smelt (Osmerus eperlanus). The results indicated that Chaoborus flavicans was the main predator of cladocerans in the stratifying area of Lake Hiidenvesi. The clay turbidity strongly contributed to the coexistence of chaoborids and smelt at high densities. Turbidity exceeding 30 NTU combined with light intensity below 0.1 μE m-2 s-1provides an efficient daytime refuge for chaoborids, but turbidity alone is not an adequate refuge unless combined with low light intensity. In the non-stratifying shallow basins of Lake Hiidenvesi, light intensity exceeds this level during summer days at the bottom of the lake, preventing Chaoborus forming a dense population in the shallow parts of the lake. Chaoborus can be successful particularly in deep, clay-turbid lakes where they can remain high in the water column close to their epilimnetic prey. Suspended clay alters the trophic interactions by weakening the link between fish and Chaoborus, which in turn strengthens the effect of Chaoborus predation on crustacean zooplankton. Since food web management largely relies on manipulations of fish stocks and the cascading effects of such actions, the validity of the method in deep clay-turbid lakes may be questioned.

How Kings are Made, How Kingship Changes : A Study of Ritual and Ritual Change in Pre-Colonial Owamboland, Namibia

This study discusses the legitimacy basis of political power and its changes in historical African societies. It starts from Luc de Heusch s tenet that political power required a legitimacy basis of a spiritual kind, often formulated as sacred kingship. In ancient and pre-literate societies such kings were held to be responsible for the fertility of man, land and cattle. The king was a paradoxical figure, symbolising society, but standing above it, while simultaneously being its victim by being ritually killed at old age. This was also how Owambo sacred kings were conceived. De Heusch suggested that African kings derived their power over fertility from having been made sacred monsters in the rituals of installation. With the example of Owambo kingship, this study argues that the transgressive and monstrous aspect is only one of several dimension of a king s sacredness and brings out the nurturing and symbolically female aspect, identified but not analysed further by de Heusch. In the Owambo kingly installation a king-elect was made sacred, and part of it was that a link was ritually created to the early owners of the land. Their consent made it possible for the king to promote fertility and to appropriate power emblems needed for ruling. In the kingdom of Ondonga the early owners of the land were the spirits of early Bushman inhabitants and those of an early kingly clan, both neglected in public memory. The sacred dimension of kingship was further augmented when kings manipulated and appropriated rain rituals and initiation rituals, both of which were related to fertility. The study argues that even though there were aspects of the sacred monster in Owambo kingship, its manifestation was, in part, a distortion of the reciprocal aspect of kingship that was expressed in the homage paid to various ancestor spirits. A change in succession practices from ritual regicide to political assassination took place concomitant with the introduction of firearms, and this broke the sacrificial aspect of sacred kingship paving the way for a more predatory form of kingship while the sacred status of the king was retained.

Somali State Failure: Players, Incentives and Institutions

In Somalia the central government collapsed in 1991 and since then state failure became a widespread phenomenon and one of the greatest political and humanitarian problems facing the world in this century. Thus, the main objective of this research is to answer the following question: What went wrong? Most of the existing literature on the political economy of conflict starts from the assumption that state in Africa is predatory by nature. Unlike these studies, the present research, although it uses predation theory, starts from the social contract approach of state definition. Therefore, rather than contemplating actions and policies of the rulers alone, this approach allows us to deliberately bring the role of the society – as citizens – and other players into the analyses. In Chapter 1, after introducing the study, a simple principal-agent model will be developed to check the logical consistence of the argument and to make the identification of causal mechanism easier. I also identify three main actors in the process of state failure in Somalia: the Somali state, Somali society and the superpowers. In Chapter 2, so as to understand the incentives, preferences and constraints of each player in the state failure game, I in some depth analyse the evolution and structure of three central informal institutions: identity based patronage system of leadership, political tribalism, and the Cold War. These three institutions are considered as the rules of the game in the Somali state failure. Chapter 3 summarises the successive civilian governments’ achievements and failures (1960-69) concerning the main national goals, national unification and socio-economic development. Chapter 4 shows that the military regime, although it assumed power through extralegal means, served to some extent the developmental interest of the citizens in the first five years of its rule. Chapter 5 shows the process, and the factors involved, of the military regime’s self-transformation from being an agent for the developmental interests of the society to a predatory state that not only undermines the interests of the society but that also destroys the state itself. Chapter 6 addresses the process of disintegration of the post-colonial state of Somalia. The chapter shows how the regime’s merciless reactions to political ventures by power-seeking opposition leaders shattered the entire country and wrecked the state institutions. Chapter 7 concludes the study by summarising the main findings: due to the incentive structures generated by the informal institutions, the formal state institutions fell apart.