Through the life history theory perspective, this paper seeks to demonstrate how early adversity shapes the development of effortful control in ways that aim to best match the individual to the proximal environment toward ultimate goals despite trade-offs related to social, academic, and later health outcomes. Investigation linking early life harshness (i.e., cues of extrinsic morbidity-mortality; Ellis et al., 2009) and unpredictability (i.e., stochastic changes in environmental conditions; Ellis et al., 2009) to the development of self-regulation could facilitate a more nuanced understanding of early environmental effects on development. The current study investigates early environmental harshness and unpredictability as unique predictors for a self-regulation construct, effortful control. It was hypothesized that early life harshness and unpredictability would uniquely and negatively predict effortful control among preschoolers. While there was no evidence that cues of unpredictability predicted effortful control, cues of harshness, specifically neighborhood harshness, did statistically significantly predict effortful control in the direction expected. This appears to be the first study to explicitly investigate effortful control development in early childhood within the harshness and unpredictability framework.

Abstract This study investigated the ecophysiology of one of the world's northernmost Atlantic salmon (Salmo salar L.) stocks in the River Teno. The juvenile years of salmon of same genetic background were studied in the wild and in the hatchery conditions. In addition, the maternal size effect on reproduction was studied in wild females. Benefit of body size was not only quantitative but also qualitative in reproduction success of the wild female salmon in the River Teno. Total number of eggs and energy content of eggs were higher in big females than in smaller ones. In the hatchery, under natural day length and water temperature conditions, the growth rate, liver glycogen content and condition factor of the parr was higher than in the wild. The liver glycogen content of the hatchery-reared parr increased throughout the growing season and decreased during winter, whereas that of the wild parr was the lowest in summer, and stayed relative stable from September to May. The observed differences in annual fluctuation in liver glycogen content may reflect the differences in carbohydrate content of feed and in behaviour between the hatchery and wild. Overall, the hatchery-reared juveniles maturated and smoltificated 1–2 years earlier than the wild fish. The mean age of wild precocious males was 3 years and that of wild smolts 4 years. However, there was considerable variation in the age of precocious males (1–6 yr) and smolts (2–8 yr) in the wild. The maturing and smolting juvenile age groups were restricted to two (1–2 yr and 2–3 yr, respectively) in the hatchery. In June, the hypo-osmoregulatory ability of hatchery smolts was developed parallel to the wild smolts. Some differences in physiological parameters between different smolt groups could be observed in the wild and between hatchery-reared and wild smolts indicating that completing of smolting process varies to some extent under different conditions. However, the hatchery-reared smolts showed higher levels of fin damage and body energy stores than the wild smolts. As the hatchery practices should aim at controlling the quantity and quality of the juvenile salmon in production, the environmental conditions governing the physiological development of the juvenile fish should be taken into account. Hatchery practices should be planned so that the seasonal timing of smolting would follow the wild fish as close as possible.

Abstract Females may either mate with one (monandry) or several (polyandry) males during a single breeding season. The polyandrous mating system has prompted numerous studies since the recognition of a widespread occurrence of mixed paternity among animals. Consequently, the benefits of polyandry have become well-established, and the female role in sexual selection upgraded. Females may gain both material and genetic benefits from multiple mating. Hence, the occurrence of polyandry is understandable, whereas monandry remains an evolutionary puzzle especially among species with male nutrient provisioning. I studied both the life history variation among female mating tactics and the adaptive significance of monandry in varying environmental conditions in the green-veined white butterfly [Pieris napi, (L. 1758)], which is a predominantly polyandrous species with nuptial feeding. I used a combination of explicit laboratory experiments and field studies. My results show that monandry and degrees of polyandry are distinct strategies with life history differences reaching beyond mating frequencies. Polyandry corresponded with a higher lifetime fecundity than monandry in P. napi. Polyandry was, however, associated with relatively low fecundity during the early days of reproduction. Thus, monandry is beneficial if time for reproduction is limited severely enough or other female traits or behaviours associated with polyandry are traded off against longevity. Due to temporal variation in reproductive rate among mating tactics, offspring of polyandrous females have less time to complete development. Accordingly, polyandrous females developed at a faster rate as larvae than monandrous ones under optimal conditions. Despite growth rate variation, monandrous females were more likely to contribute to additional summer generation in conditions that allow production of only a partial second generation, and thus monandry is favoured under these conditions. Genetic variation in female mating tactics will not only prevail if environmental conditions do not allow all individuals to contribute evenly to the directly breeding generation in bivoltine populations, but also if even the production of a single generation per year is time-limited. A general conclusion would be that seasonality and unpredictability of fitness in the wild drives the evolution of optimal female mating tactics and promotes the maintenance genetic variation in mating frequencies, regardless of the direct benefits of nuptial feeding. Even if a high degree of polyandry would be the most profitable mating tactic in an average year, strong annual variation in weather conditions and the duration of summer may create possibilities for a temporally fluctuating selection that promotes a co-existence of different mating tactics because variance of fitness is likely increase with an increasing mating frequency.

Whilst individual differences in circulating hormone levels can influence life history traits throughout an animal’s lifetime, this remains a poorly understood area of research, particularly for wild systems where sufficient sets of individual-based data are rare. This thesis aimed to address this dearth of information by identifying key drivers of hormone variation, as well as exploring potential fitness consequences within a single system of wild red deer (Cervus elaphus) on the Isle of Rum National Nature Reserve in Scotland. It focussed on both androgen (e.g. testosterone) and glucocorticoid (e.g. cortisol) levels, and examined among-individual variation in these two hormone groups from samples collected using both traditional (blood: chapters 3 & 4) and non-invasive (faecal: chapters 5 & 6) methods. Results showed both intrinsic and extrinsic factors to influence an individual’s hormone levels. In general, current or recent environment explained the greatest variation, with both hormone groups exhibiting strong temporal trends at multiple scales. Concentrations changed substantially across an individual’s lifetime as they aged (chapters 5 & 6), and calves born in different years differed in their neonatal testosterone levels (chapter 3). Hormone levels also varied across the year, showing clear seasonal cycles which peaked during key reproductive events: the calving season in females (chapter 6) and the rut in males (chapter 5). An individual’s current life history state was also important, particularly a female’s reproductive state (chapter 6). Whilst there was some evidence of maternal effects on neonatal hormone levels (chapter 3) these were not extensive, and maternal hormone concentrations did not appear to influence those in their new-born calves (chapter 6). There was, however, evidence of neonatal circulating testosterone levels being heritable, and despite overall differences between the sexes the underlying genetic architecture of this trait did not differ between male and female calves (chapter 4). Associations were also found between an individual’s hormone levels and their fitness, although these consequences were only apparent in short-term fitness measures or proxies such as reproductive behaviour (e.g. male reproductive effort in chapter 5). Effects were also not ubiquitous within the population. Whilst a calf’s circulating testosterone levels indicated their probability of surviving their first year of life, these effects were only apparent in firstborn males, a group which is particularly vulnerable to mortality (chapter 3). In general, this thesis suggests that the fitness consequences identified by broad-scale hormone manipulation studies can still be found when looking at subtle individual-level differences. The limited evidence of persistent hormone phenotypes (indicated by the lack of among individual variance for most measures, chapter 5 & 6) does, however, emphasise the importance of repeatedly sampling individuals before drawing extensive conclusions about fitness consequences.

The purpose of this dissertation is to test how growth, reproduction, and immune functioning interact in two populations of adult women residing in vastly different socio-economic and ecological environments, the Kalunga quilombo in Brazil, and the United States of America. The presence of life history trade-offs was tested to determine how the different envirnonments, and socio-ecological contexts of the populations were creating differential risks for health and reproductive outcomes, and life history trade-offs. I hypothesized that the Kalunga people, living in very difficult and harsh conditions, would experience greater amounts of, and more severe, life history trade-offs than the U.S. women. I also hypothesized that in both populations, syndemic interactions between health outcomes, and the larger macro-social conditions, would influence stature, immune functioning, and reproduction. Additionally, these syndemic interactions would perpetuate poor health and reproductive outcomes within the entire Kalunga population, and portions of the U.S. population. I collaborated with the University of Brasilia to collect data adult women from in the Kalunga quilombo in Goiás State, Brazil (n=38) via semi-structured interviews in 2016. In addition, these data were compiled with reproductive, demographic, anthropometric, biological, and socio-economic data collected from adult Kalunga women in 2015 by the University of Brasilia’s Department of Genetics and Morphology. Demographic, anthropometric, biological, and socio-economic data recorded in the National Health and Nutrition Examination Survey (NHANES) 2005-2006 from adult U.S. women was also analyzed. Life history traits were measured through stature (cm), Immunoglobulin E (IgE; kU/l), and reproductive variables (fertility, and reproductive ages) for both populations. My results demonstrated that life history trade-offs impact diverse populations. I found evidence to support life history trade-offs occurred between stature and fertility, and stature and reproductive ages, in both populations. However, my results also indicated that the socio-economic, and ecological, conditions of both populations heavily influenced stature and reproduction in the women. Life history trade-offs were not observable between immune functioning and fertility in the Kalunga population, and not readily evident in the U.S. sample. The positive associations between IgE and fertility in the U.S. women indicate that while life history trade-offs may be suggested, other biocultural variables including: low income, ethnicity, education, and body size, are stronger influences on immune functioning, and fertility, in the U.S. women. Though the Kalunga women had a history of infectious and parasitic diseases, and lived in conditions of poor sanitation and poverty, they had multiple ways of mitigating the stresses of their everyday life. The shared socio-ecological conditions of their community decreased the deprivations they experienced, reduced the disabilities they felt from their health statuses, and was reflected in relative homogeneity of their lived experience. In contrast, the U.S. population was dictated by disparities. Poor health and reproductive outcomes were concentrated disproportionately in low-income, less educated, and/or minority ethnicity status, women. The findings of this dissertation have important implications for applied anthropology and the study of life history theory. My results demonstrate that Western lifestyles must be considered as instigators of life history trade-offs, and life history scholarship in humans must also focus on populations living in these conditions. Additionally, the secular changes resulting in lowered ages of menarche in U.S. women could lead to future poor health outcomes. As the Kalunga transitions into a more Western lifestyle, they are at risk for disparities within their population. It is imperative for applied anthropologists to be engaged in addressing the immune, nutritional, and psychosocial stressors within populations that perpetuate disparities, and instigate and/or exacerbate life history trade-offs.

Climatic variation is an important driver of avian life history and population dynamics. Climate change models predict increased variability for many regions and to predict the effects on species, we need to examine how their life history characteristics influence their response to climate. I studied how environmental conditions influenced the ecology of white-tailed (Lagopus leucura) and rock ptarmigan (L. mutus) in tundra habitats of the southern Yukon Territory. Although sympatric in the study area, breeding territories were generally segregated, with white-tailed ptarmigan selecting steep, rocky slopes at higher elevations and rock ptarmigan preferring lower elevation sedge meadows. For both species, cold spring temperatures delayed the onset of breeding, resulting in smaller clutch sizes and fewer hatched young per female. However, delayed breeding led to a stronger reduction in these rates for rock ptarmigan, suggesting a lower resilience to extend reproductive effort in colder years. White-tailed ptarmigan were also more likely to re-nest following failure and had higher daily nest survival, both of which contributed to greater annual productivity compared to rock ptarmigan. Annual adult survival showed the opposite pattern to productivity as rock ptarmigan survival was 24 percent higher than white-tailed ptarmigan. This finding suggested a reproduction-survival trade-off exists for the two species, which may be driven by differing susceptibility to environmental factors in the region. Life history theory predicts that if the likelihood of future breeding opportunities is low, individuals should increase current reproductive effort, which may explain why white-tailed ptarmigan have longer breeding seasons and higher reproductive effort under unfavourable climatic conditions. Population models showed that growth rates (λ)were approximately stable for rock ptarmigan (λ=1.01), but declining for white-tailed ptarmigan (λ=0.96). Simulations showed that warmer spring temperatures over the next few decades would elevate λ by ~0.05 for both species, but the extent of increase in λ may be reduced with more variable spring conditions. Population growth will also depend on how changing winter conditions influence survival for each species. Model simulations suggest that if juvenile and adult survival are positively correlated, rock ptarmigan would be more resilient to severe years that simultaneously depress reproduction and survival. / Forestry, Faculty of / Graduate

京都大学 / 新制・課程博士 / 博士(理学) / 甲第23040号 / 理博第4717号 / 新制||理||1676(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 渡辺 勝敏, 教授 曽田 貞滋, 教授 沼田 英治 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Early exposure to hypoxia is related to a variety of physiological and metabolic changes that have lasting effects on organisms’ physiology and life history. We measured the effects of maternal and embryonic mild, intermittent hypoxia on the life history of four clones of microcrustacean Daphnia magna, an emerging model organism for the studies of senescence and longevity. Daphnia individuals were produced parthenogenically, maintained in individual vials, and fed standard algal concentration daily. The cohort consisted of 189 individuals. We measured body size at first reproduction, fecundity (including late-life fecundity peak), offspring sex ratio, and longevity. We found no effect of maternal and embryonic hypoxia on body size and longevity; however, there was a slight but statistically significant increase in age-specific mortality in the early hypoxia treatment cohort. Daphnia from the hypoxia group showed higher early fecundity which disappeared by the age of 100 days. A late-life spike in fecundity was observed at the age of 100 days when hypoxia group individuals showed significantly lower fecundity. There was little evidence of a trade-off between early- and late-life fecundity. Finally, early hypoxia affected mid-life male production in one of the four clones, and we discuss possible physiological changes triggered by maternal and embryonic exposure to hypoxia.

No description available.

Predation is known to have a significant effect on life history, eliciting predictable responses. Physical constraints of body shape and size may also limit life history divergence. There may be a trade-off between adapting to predation, and limits placed by constraints that decrease life history divergence. We test this idea in the Costa Rican livebearing fish Alfaro cultratus. This species has a keeled ventral surface and does not develop a distended abdomen when pregnant like other livebearers. We describe the life history of A. cultratus in 20 different populations across predator and non-predator environments. We found significantly lower reproductive allotment in predator environments relative to non-predator environments, but no significant difference in female or male size at maturity, number of offspring, or size of offspring. We found that A. cultratus exhibit isometric patterns of allocation for clutch dry mass in relation to female dry mass in predator and non-predator environments. We suggest that body shape constraints in this species limit the life history divergence we typically see in predator and non-predator environments in other species.