The role of oxidative stress as an underlying mechanism in life-history trade-offs

A key aspect underpinning life-history theory is the existence of trade-offs. Trade-offs occur because resources are limited, meaning that individuals cannot invest in all traits simultaneously, leading to costs for traits such as growth and reproduction. Such costs may be the reason for the sub-maximal growth rates that are often observed in nature, though the fitness consequences of these costs would depend on the effects on lifetime reproductive success. Recently, much attention has been given to the physiological mechanism that might underlie these life-history trade-offs, with oxidative stress (OS) playing a key role. OS is characterised by a build-up of oxidative damage to tissues (e.g. protein, lipids and DNA) from attack by reactive species (RS). RS, the majority of which are by-products of metabolism, are usually neutralised by antioxidants, however OS occurs when there is an imbalance between the two. There are two main theories linking OS with growth and reproduction. The first is that traits like growth and reproduction, being metabolically demanding, lead to an increase in RS production. The second involves the diversion of resources away from self-maintenance processes (e.g. the redox system) when individuals are faced with enhanced growth or reproductive expenditure. Previous research investigating trade-offs involving growth or reproduction and self-maintenance has been equivocal. One reason for this could be that associations among redox biomarkers can vary greatly so that the biomarker selected for analysis can influence the conclusion reached about an individual’s oxidative status. Therefore the first aim of my thesis was to explore the strength and pattern of integration of five biomarkers of OS (three antioxidants, one damage and one general oxidation measure) in wild blue tit (Cyanistes caeruleus) adults and nestlings (Chapter 2). In doing so, I established that all five biomarkers should be included in future analyses, thus using this collection of biomarkers I explored my next aims; whether enhanced growth (Chapters 3 and 4) or reproductive effort (Chapter 5) can lead to increased OS levels, if these traits are traded off against self-maintenance. I accomplished these aims using both a meta-analytic and experimental approach, the latter involving manipulation of brood size in wild blue tits in order to experimentally alter growth rate of nestlings and provisioning rate (a proxy for reproductive expenditure) of adults. I also investigated the potential for redox integration to be used as an index of body condition (Chapter 2), allowing predictions about future fitness consequences of changes to oxidative state to be made. A growth – self-maintenance trade off was supported by my meta-analytic results (Chapter 4) which found OS to be a constraint on growth. However, when faced with experimentally enhanced growth, animals were typically not able to adjust this trade-off so that oxidative damage resulted. This might support the idea that energetically expensive growth causes resources to be diverted away from the redox system; however, antioxidants did not show an overall reduction in response to growth in the meta-analysis suggesting that oxidative costs of growth may result from increased RS production due to the greater metabolism needed for enhanced growth. My experimental data (Chapter 3) showed a similar pattern, with raised protein damage levels (protein carbonyls; PCs) in the fastest growing blue tit chicks in a brood, compared with their slower growing sibs. These within-brood differences in OS levels likely resulted from within-brood hierarchies and might have masked any between-brood differences, which were not observed here. Despite evidence for a growth – self-maintenance trade off, my experimental results on blue tits found no support for the hypothesis that self-maintenance is also traded off against reproduction, another energetically demanding trait. There was no link between experimentally altered reproductive expenditure and OS, nor was there a direct correlation between reproductive effort and OS (Chapter 5). However, there are various factors that likely influence whether oxidative costs are observed, including environmental conditions and whether such costs are transient. This emphasises the need for longitudinal studies following the same individuals over multiple years and across a wide range of habitats that differ in quality. This would allow investigation into how key life events interact; it might be that raised OS levels from rapid early growth have the potential to constrain reproduction or that high parental OS levels constrain offspring growth. Any oxidative costs resulting from these life-history trade-offs have the potential to impact on future fitness. Redox integration of certain biomarkers might prove to be a useful tool in making predictions about fitness, as I found in Chapter 2, as well as establishing how the redox system responds, as a whole, to changes to growth and reproduction. Finally, if the tissues measured can tolerate a given level of OS, then the level of oxidative damage might be irrelevant and not impact on future fitness at all.

A house, not a home? Examining the use of the private rented sector to resolve homelessness in Scotland

Changes to homelessness legislation in post-devolution Scotland have resulted in an expansion of rights for homeless households seeking formal assistance from local authorities. These changes have led to Scotland’s homelessness arrangements being considered among the most progressive in Europe. In recent years, however, the Scottish Government has increasingly promoted homelessness prevention and Housing Options approaches as a means by which homelessness might be avoided or resolved without recourse to statutory rights. As part of that, they have promoted greater use of the private rented sector (PRS) as a key housing option, with the potential to meet the needs of homeless households. The arguments made to support use of the PRS have much in common with arguments for privatisation in other areas of social policy, notably greater choice for the individual promoting better welfare outcomes, and competition among providers encouraging improvements in quality of service provision. Critics have argued that such benefits may not be realised and that, on the contrary, privatisation may lead to exclusion or act to worsen households’ outcomes. This thesis considers the extent to which the PRS has been utilised in Scotland to accommodate homeless households, and the consequences of this for their welfare. The thesis uses a combination of quantitative and qualitative methods. To examine trends in the use of the PRS, it presents quantitative analysis of the data on the operation of the statutory system and Housing Options arrangements, and of data from a survey of local authority homelessness strategy officers. To examine the consequences of this for homeless households, the thesis uses qualitative research involving face-to-face interviews with 35 homeless households across three local authority areas. This research considers the extent to which households’ experiences of homelessness, housing need and the PRS reflect the arguments presented in the literature, and how settled accommodation has impacted on households’ ability to participate fully in society. The research found an increasing but still limited role for the PRS in resolving statutory homelessness in Scotland, with indications that the PRS is being increasingly used as part of the Housing Options approach and as a means of resolving homelessness outside the statutory system. The PRS is being utilised to varying degrees across different local authority areas, and a variety of methods are being used to do so. While local authorities saw clear advantages to making greater use of the sector, a number of significant barriers including affordability, available stock and landlord preferences - made this difficult in practice. Research with previously homeless households in the PRS similarly found broadly positive experiences and views of the sector, particularly with regard to enabling households to access good quality accommodation in desirable areas of their choosing, with many households highlighting improvements relating to social inclusion and participation. Nevertheless, concerns around the security of tenure offered by the sector, repairs, service standards and unequal power relations between landlord and tenant persisted. As such, homeless households frequently expressed their decision to enter the sector in terms of a trade-off between choice and security.