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Early Life Developmental Effects Workshop 9th-12th September 2015

St Michael’s Hotel, Falmouth

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Table of contents Workshop outline 3 Information for speakers 4 Programme 5 Abstracts 7 Session 1: Theories of early-life effects 7 Session 2: Mechanisms of information transfer 8 Session 3: Individual effects 10 Session 4: Implications and future perspectives 12 Map of Falmouth and local information 13 List of participants 14 Workshop sponsors 16

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Workshop outline Early life developmental effects: unifying evolutionary and biomedical perspectives

Summary

Variation in early life conditions can trigger developmental switches that lead to predictable individual differences in adult behaviour and physiology. Despite evidence for such early life effects being widespread both in humans and throughout the animal kingdom, the evolutionary causes and consequences of this developmental plasticity remain unclear.

Possible hypotheses explaining this developmental plasticity include the suggestion that it reflects ‘predictive adaptive responses’ to information provided by the maternal or early life social environment which maximise an offspring’s phenotypic match to future environments. However, this hypothesis has been contested on the grounds that long-range forecasts of the adult environment are inherently unreliable. Alternative hypotheses suggest that developmental plasticity during early life may simply reflect variation in constraints and conditions during development, non-adaptive instability, or the outcome of parent-offspring conflict over developmental trajectories.

This cross-disciplinary, NERC-funded workshop aims to bring together researchers of early life effects from the fields of both evolutionary ecology and biomedicine to synthesise and advance current knowledge of how information is used during development, the mechanisms involved, and how early life effects evolved. With a main focus on mammalian systems, the workshop will include three themed talk and discussion sessions and a further perspectives session where we will discuss the broader implications of early life effects.

Workshop Sessions and Talk Topics

1. Theories of early-life effects Plenary Speaker: John McNamara FRS, University of Bristol To establish a conceptual framework our first session will focus on theoretical approaches to questions that are at the cutting edge of current understanding of evolution and development. What sources of information are used to shape development, and why? Can humans and other mammals evolve ‘predictive adaptive responses’ to improve their phenotypic match to future environments? How are within-family conflicts over offspring developmental trajectories resolved? Questions such as these can be addressed using evolutionary game theory, quantitative genetics, and economic models of ‘skill formation’. Our hope in this session is to find common principles that emerge from different theoretical perspectives; to discuss the plausibility and testability of models, and identify new avenues for empirical research.

2. Mechanisms of information transfer Plenary Speaker: Ton Groothuis, University of Groningen, Netherlands This session will focus on the mechanisms by which the early life environment can impinge on the health, behaviour, and fitness of individuals via genetic, epigenetic, physiological and social effects. Talks will examine the role of resource availability and maternal state in determining offspring phenotype, and whether maternal endocrinology and epigenetic effects have evolved to buffer or mediate these effects. They will also discuss how pre- and postnatal parental care, early social environment, and stress physiology can have profound impacts on the fitness and adult health of offspring. A major goal of this session is to evaluate whether the mechanisms of developmental switches are best understood as the consequence of constraints or as adaptive responses to cues and signals from the maternal or ecological environment.

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3. Individual effects Plenary Speaker: Rebecca Sear, London School of Hygiene and Tropical Medicine In this session we will explore the individual responses to and consequences of the physiological and epigenetic changes discussed in the previous session. We will discuss how early life stressors and investment affect an individual’s ageing, cognition, behaviour, health, and reproductive decisions and whether these developmental responses are adaptive. We will examine why there is variation in an individual’s response to early life stressors, which early life effects are important and if there are critical windows of vulnerability or remediation. We also hope to discuss whether these responses can be primed, moderated or reversed and if so how can we predict them?

4. Implications and future perspectives This final session will focus on the broader implications of early life effects, and identify the most promising frontiers for future research. We will explore how improved conceptual understanding of early life effects can inform fields such as demography, life history evolution and conservation, what major outstanding questions need to be answered, and whether the aims and insights of evolutionary and biomedical research can be reconciled and synthesised.

Organisers: The workshop is funded by a NERC award to Michael Cant and Andrew Young of the University of Exeter, Cornwall, and is organised by Harry Marshall, Jennifer Sanderson, Emma Vitikainen, Michelle Hares, Faye Thompson, Emma Inzani, Andy Young and Mike Cant. This team is currently funded by grants from NERC and the ERC to study early life effects, social and cognitive development in wild mammals. Our research integrates >20 years continuous field study on wild banded mongooses in Uganda with investigation of telomere dynamics, oxidative stress, stress endocrinology, disease biology, behaviour and life history evolution. Full information on our project is given on our website www.bandedmongoose.org

Information for speakers: There will be both a Windows PC and Mac laptop loaded with Microsoft Office software available for speakers to upload their talks to. It would be ideal if speakers could load their presentations on to one of these laptops the day before they are scheduled to give a talk. The laptops will be available during the welcome drinks on the Wednesday evening and after talks and discussions finish on the Thursday.

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Programme Wednesday 9th September Afternoon Residential participants arrive at St Michael’s Hotel* 17:30 Welcome drinks reception in the Locker room, St Michael’s Hotel 19:00 Dinner in St Michael’s restaurant *N.B Please note that participants may arrive at St. Michael’s Hotel from 1pm onwards on 9th September, although please note luggage and room check will commence at 3pm. Thursday 10th September

09:15 Session 1: Theories of early-life effects Chair: Mike Cant

09:15 Welcoming words Mike Cant 09:20 The evolution of transgenerational integration of information

in heterogeneous environments John McNamara

10:00 Short break

10:15 Do older mothers know best? Nongenetic effects in age-structured populations Bram Kuijper

10:30 The evolution of sensitive windows in a model of incremental development Willem Frankenhuis

10:45 The adaptive use of information during growth and long-term effects of early-life experiences Sinead English

11:00 Coffee break

11:30 Theory discussion in small groups 12:00 Discussion summing up 12:30 Lunch at St Michael’s restaurant

13:30 Session 2: Mechanisms of information transfer Chair: Michelle Hares

13:30 On the interpretation of (hormone mediated) maternal effects Ton Groothuis 14:10 Mechanisms of information transfer: maternal endocrinology Mandy Drake 14:25 Epigenetic pathways to mental illness Jonathan Mill 14:40 Profiling vertebrate epigenome’s dynamic response to a

fluctuating early life environment, and their effects on populations’ phenotype Ursula Paredes

14:55 Coffee Break

15:25 The snowshoe hare 10-year population cycle: impact of predator-induced maternal effects Sophia Lavergne

15:40 Transgenerational impacts of oxidative damage explain life history variation in a wild mammal Emma Vitikainen

15:55 Early-life IGF-1 predicts life history traits in wild spotted hyenas Nora Lewin

16:10 Effects of early adversity on ageing and cognition in the European starling Daniel Nettle

16:25 Short Break

16:40 Mechanisms discussion in small groups 17:10 Discussion summing up 19:15 Dinner in St Michael’s restaurant

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Friday 11th September

09:15 Session 3: Individual effects Chair: Harry Marshall

09:15 What can humans tell us about the consequences of early life stress for life history strategies? Illustrated with data on parental absence Rebecca Sear

09:55 Mechanisms of information transfer: prematurity in an evolutionary context Thomas Williams

10:10 Evidence for metabolic programming by maternal nutrition in humans Caroline Fall

10:25 Coffee break Chair: Emma Vitikainen

10:55 Early ecological environmental influences on later-life behaviour and life history in banded mongooses Harry Marshall

11:10 Developmental plasticity to prepare for life in a complex world Barbara Taborsky 11:25 Assessing links between early-life disease exposure and human

life-history traits Adam Hayward 11:40 Early life developmental effects: unifying evolutionary and

biomedical perspectives Jonathan Wells

11:55 Short break

12:10 Individual effects discussion in small groups 12:40 Discussion summing up 13:10 Lunch at St Michael’s Restaurant

14:10 Session 4: Implications and future perspectives Chair: Andy Young

14:10 Early in life experience and heredity Étienne Danchin 14:40 Early life programming - current knowledge and future

potential Susan Ozanne

15:10 Coffee break

15:40 What is this thing called developmental programming? Tobias Uller 16:10 Evolutionary insights for global health Mark Hanson

16:40 Short break

17:00 Implications and perspectives discussion in small groups 17:30 Discussion summing up 19:15 Dinner in Falmouth at The Stable Saturday 12th September 10:00 Workshop summing up 10:45 Final workshop admin 11:00 Participants depart

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Abstracts Thursday 10th September Session 1: Theories of early-life effects

1. Plenary: The evolution of transgenerational integration of information in heterogeneous environments John McNamara It is often useful to think of developmental systems as integrating available sources of information about current conditions to produce mature organisms. Such processes are under genetic control, and subject to Darwinian selection, so we expect them to evolve to produce mature organisms that fit well with current ecological (including social) conditions. Genes and inherited physiology provide cues, as does the state of the environment during development. I present a general framework for linking genetic and phenotypic variation from an informational perspective, illustrating this perspective by specific models.

2. Do older mothers know best? Nongenetic effects in age-structured populations Bram Kuijper & Rufus Johnstone Parental information transmitted via nongenetic effects can inform offspring about the environment they will face, allowing them to develop an appropriate response. Current predictions on the evolution of these nongenetic effects assume, however, that parents facing a variable environment are all identical in state. This raises the question whether parents of different age or condition should all respond similarly to enviromental heterogeneity. To assess this, we model the evolution of nongenetic effects in an age-structured population. Interestingly, we find substantial differences between young and old mothers in the probability of nongenetic inheritance, with younger mothers being much less likely to copy their phenotype to offspring than older mothers. The reason for these age differences is that an older parent's phenotype is more likely to be adapted, as maladapted parents are less likely to reach old age. Overall, our model predicts that the magnitude and directionality of nongenetic effects should vary drastically over the lifespan of an individual.

3. The evolution of sensitive windows in a model of incremental development

Willem Frankenhuis

Sensitive windows are widespread in nature. Despite a recent focus on neural-physiological explanation, few formal models have examined the evolutionary selection pressures that result in developmental mechanisms that produce sensitive windows. Here, we present such a model. We model development as a specialization process during which individuals incrementally adapt to local environmental conditions, while receiving a constant stream of cost-free, imperfect cues to the environmental state. We compute optimal developmental programs across a range of ecological conditions and use these programs to simulate developmental trajectories and obtain distributions of mature phenotypes. This talk will focus on our main results.

4. The adaptive use of information during growth and long-term effects of early-life experiences

Sinead English Why do experiences in early life have long-term effects when individuals have the opportunity to adjust their phenotype? Here, I describe a model of growth and maturation in an uncertain world to explain how the coupled dynamics of information use and phenotypic change can result in such effects. Our model shows that, even when plasticity is unconstrained, early experiences can have long-term impacts on the timing of maturation. Key insights that may be useful for empiricists investigating early-life effects are, first, the importance of considering the later offspring environment; and, second, experiences should be manipulated across multiple stages of development.

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Session 2: Mechanisms of information transfer

5. Plenary: On the interpretation of (hormone mediated) maternal effects Ton G. G. Groothuis

Parents can provide resources and signals to their offspring already in the prenatal stage, containing information about the environment is which the offspring will develop. I will focus on hormone mediated maternal effects and discuss the possibility that its information transfer is less constrained than previously thought, that not only the role of the mother but also that of the embryo should be taken into account, and that the adaptive value of maternal effects can only be evaluated by taking into account that postnatal information can interact in several ways with information acquired during the prenatal phase, explaining the many discrepancies in the literature.

6. Mechanisms of information transfer: maternal endocrinology

Mandy Drake

Many epidemiological and animal studies have now demonstrated a clear link between the environment experienced in early life and an increased risk of cardiometabolic and neurodevelopmental disorders. In this talk I will discuss how maternal endocrine effects may affect the in utero environment, leading to programmed effects in the offspring. These effects include maternal stress, resulting in fetal glucocorticoid overexposure and maternal overnutrition/obesity with consequent insulin resistance/ hyperglycaemia. Such effects may explain the transmission of programmed effects across generations through the maternal line.

7. Epigenetic pathways to mental illness

Jonathan Mills

Current approaches to understanding the etiology of mental illness has primarily focussed on uncovering a genetic contribution to disease-onset. Despite the success in identifying novel susceptibility genes for disorders including schizophrenia, bipolar disorder and autism using genome-wide association study approaches, these loci account for only a small proportion of attributable risk and the mechanism behind their action remains unknown. Sequencing the genome was only the first step in our quest to understand how genes are expressed and regulated. Sitting above the DNA sequence is a second layer of information (the"epigenome") that regulates several genomic functions, including when and where genes are expressed. There is growing recognition that epigenetic mechanisms are important in the etiology of complex disease, acting at the interface between the genome and the environment. Recent technological advances mean that it is now feasible to study epigenetic marks such as DNA methylation at base-pair resolution across the genome. In this talk I will present data from our group showing how dynamic epigenetic processes are involved in neuropsychiatric phenotypes, and can be influenced by environmental, genetic and stochastic factors. I will describe results from ongoing studies of disease-associated epigenetic changes using post-mortem brain material, discordant monozygotic twins and clinical samples.

8. Profiling vertebrate epigenome’s dynamic response to a fluctuating early life environment, and their

effects on populations’ phenotype Ursula Paredes

I will discuss my recent work, studying the methylome of vertebrates affected by fluctuating environments in early life. I will examine the effect of differences in levels of pollution and nest habitat quality (birds) and incubation temperature (lizards) on the methylome, and how such epigenetic variation correlates with population’s phenotype. I will also discuss how social status of female baboons affect their epigenome, and that of their daughters and mediate the social stress reactivity of their offspring. Finally, I will discuss how female epigenome contribute to individuals’ response fluctuations in food availability and drought.

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9. The snowshoe hare 10-year population cycle: impact of predator-induced maternal effects Sophia Lavergne

Snowshoe hares are a keystone herbivore species whose predictable 8-10 year population cycle dominates terrestrial community dynamics in Canada’s boreal forest. The cycle is top-down regulated by predation, which is key to all but the 2-5 year low phase. The lag in population recovery that characterizes the low has been attributed to lingering chronic stress effects from the intense predation risk hares experience during the decline, where annual survival can drop to 0.5%. The fear and psychological stress of predator exposure have been associated with many long-term effects such as altered dendritic morphology and glucocorticoid levels, and anxiety-like behaviours. If experienced during sensitive developmental periods (pregnancy and lactation), stress can have long term impacts on offspring brain organization and trait expression. Previous work on hares has shown that females subjected to artificially increased predation risk during pregnancy have reduced reproductive fitness and their offspring inherit an altered stress axis. My research uses experimental manipulations and natural variation of predator risk in wild snowshoe hares to assess the impact of predator-induced stress on gene expression and epigenetic patterns in the brain, to see if these patterns can be induced experimentally by manipulating the perceived risk of predation, and to test whether prenatal stress has potentially adaptive downstream impacts on offspring exploratory and antipredator behaviour. My talk will discuss the challenges and the benefits of using a non-model species to study maternal effects and consider future research directions into the role of epigenetic programming as the mechanism underlying stress inheritance in hares.

10. Transgenerational impacts of oxidative damage explain life history variation in a wild mammal Emma Vitikainen Life history theory assumes a trade-off of resources between reproduction and survival, yet the mechanism remains debated. The oxidative stress theory of aging (OSTA) suggests that energy expensive activities, such as reproduction, generate reactive oxygen species (ROS) which damage tissues and reduce survival. In the banded mongoose, oxidative damage predicted individual and sex differences in survival, but contrary to the theory, females had reduced oxidative damage during pregnancy. We show that this can be explained if mothers are selected to shield their offspring from oxidative harm during pregnancy. Our model incorporating transgenerational effects can explain life history allocation across the lifespan.

11. Early-life IGF-1 predicts life history traits in wild spotted hyenas

Nora Lewin The early nutritional environment can have profound effects on later life phenotypes and thus may figure importantly in the timing of life history traits. One proposed mechanism mediating this relationship is the anabolic hormone, insulin-like growth factor-1 (IGF-1). Although much laboratory research has elucidated the role of IGF-1 in the mediation of life history traits, no study to date has comprehensively explored this in any free-ranging mammal. Here, we investigated variation in and consequences of juvenile IGF-1 concentrations using longitudinal data on free-ranging spotted hyenas (Crocuta crocuta). We found significant effects of maternal social rank, local prey density, maternal parity, and intra-litter dominance on juvenile plasma IGF-1 concentrations in males, but not females. Relative size of juveniles of both sexes was strongly correlated with their plasma IGF-1 concentrations. Among females, higher concentrations of juvenile IGF-1 predicted younger age at first parturition, and increased survival to sexual maturity, but at a cost of decreased longevity after sexual maturity. To our knowledge, this is the first study to explore the effects of early-life IGF-1 on later-life traits in a free-ranging mammal, and greatly enhances our understanding of developmental plasticity and endocrine-mediated effects on life history patterns.

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12. Effects of early adversity on ageing and cognition in the European starling Daniel Nettle

I will present some of our recent research on the European starling. We have shown that early-life competitive disadvantage is associated with accelerated cellular ageing as measured by telomere attrition, and that telomere attrition is related to cognitive traits in adulthood. The current theoretical literature on developmental plasticity tends to focus on two types of explanation, predictive adaptive responses and maladaptive constraints. There is however an important third category which better characterises the starling case, and perhaps others: adaptive behavioural accommodation of the individual to its own somatic state, which is in turn the product of its developmental history.

Friday 11th September Session 3: Individual effects

13. Plenary: What can humans tell us about the consequences of early life stress for life history strategies? Illustrated with data on parental absence Rebecca Sears

There has been a recent flurry of literature in the human evolutionary sciences suggesting that early life stress can kickstart a ‘fast’ life history. Most evidence for this relationship comes from data correlating early life stress with earlier first births in women. Other measures of ‘life history strategy’ are rarely investigated. This talk will consider how the wide availability of data from a range of human populations can be harnessed to answer the question of whether early stress does speed up life history strategy. Particular emphasis will be placed on the loss of a parent(s) as an indicator of stress.

14. Mechanisms of information transfer: prematurity in an evolutionary context

Thomas Williams

Prematurity is a significant public health problem, and is currently the leading cause of infant mortality worldwide. Compared to other primates, humans have a wider range of gestations at birth at which survival is viable. This raises the possibility that giving birth prematurely may be an evolutionarily adaptive approach for pregnant individuals faced with adverse environmental conditions. Epidemiological studies have consistently demonstrated a link between lower socioeconomic status and an increased risk of giving birth prematurely. Supporting the hypothesis that this may be due in part to adverse environmental conditions, rates of premature delivery fall in populations showing upward socio-economic mobility, and increase in populations experiencing downward mobility. Our group has examined changes in a cohort of infants born prematurely and at term. In addition to differences in body composition between the two groups, we found changes in cortisol responses to a stressor at three months of age, and in percentage methylation at one of the differentially methylated regions controlling expression of the key growth factor IGF2 at birth. This suggests that prematurity is associated with a distinct phenotype, similar to that of infants born at a low birth weight. As with low birth weight, it is not clear whether the conditioning changes associated with prematurity are mainly a marker of adverse in utero environments (or cycles of intergenerational deprivation), or truly an adaptive response that has evolved since evolutionary divergence from other non-human primates. I will examine the data available from animal, archaeological and human studies to outline possible approaches to this question.

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15. Evidence for metabolic programming by maternal nutrition in humans Caroline Fall I will present what is known about the relationship of maternal nutritional status during pregnancy to offspring phenotype in humans, drawing upon the work of others as well as on my own work in India, where maternal undernutrition and gestational diabetes (an example of fetal ‘overnutrition’) are common problems. We have shown associations between maternal undernutrition and gestational diabetes and offspring insulin resistance, and have called these effects ‘nutrient-mediated teratogenesis’ and ‘fuel-mediated teratogenesis’ respectively, showing our bias towards ‘constraint’ rather than ‘adaptation’. Although data are limited, I will focus on the results of nutritional interventions in pregnancy, both experimental and ‘catastrophes of history’ such as famine. I will also summarise what is known about effects of maternal nutrition on the offspring epigenome in humans.

16. Early ecological environmental influences on later-life behaviour and life history in banded

mongooses Harry Marshall The ecological environment an individual experiences in early-life is expected to have profound effects on its behaviour and life history. Despite this, evidence for such effects in wild animal systems is scarce. I will present results from analyses of the Banded Mongoose Research Project’s long-term data set showing that individuals born in more variable periods invested more in reproductive behaviours as an adult, lived longer and had a greater lifetime reproductive success. These results highlight the importance of considering the variability, as well as the average, of early-life conditions and suggest that more variable early-life conditions may provide individuals with the ability to cope better with environmental challenges in later life.

17. Developmental plasticity to prepare for life in a complex world

Barbara Taborsky

The early environment often shapes phenotypes for a lifetime. The functional significance of such long-term developmental effects is still little understood. A key problem hampering our understanding is that experiments usually target only a single environmental variable and/or a single ontogenetic period, while in nature development is influenced by multiple environmental factors and during a succession of ontogenetic stages. Here I present results from cooperatively breeding cichlids suggesting that (i) maternal effects on behaviour and brain architecture can be fully reversed by direct, postnatal experience of offspring, and (ii) organisms require information on multiple environmental factors in order to express adequate plastic responses.

18. Assessing links between early-life disease exposure and human life-history traits

Adam Hayward Why has human life expectancy increased over the last 150 years? A leading hypothesis, the ‘cohort morbidity phenotype’, proposes that limited exposure to childhood infections has reduced chronic inflammation, immunosenescence and risk of death from associated diseases. Tests of this hypothesis have typically used all-cause mortality rates to estimate disease exposure in early life, but such associations could arise from factors other than early-life disease exposure. Additionally, the impact of early-life disease exposure on reproduction remains unknown, and thus previous work ignores a major component of fitness, through which life-history responses to early environmental stressors are subject to selection. We used data from 19th century Finnish church records to show that early-life exposure to disease, measured as the prevalence of deaths from infectious disease during the first five years of a child’s life, was associated with reductions in later-life survival and fertility. Early-life disease especially increased probability of death from infections in older individuals, while women experienced stronger effects of early disease than men. These results support the hypothesis that diminished early-life disease exposure may have played a role in increasing life expectancies and suggest a previously unexplored link with reproductive success.

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19. Early life developmental effects: unifying evolutionary and biomedical perspectives Jonathan Wells

In the last two decades, epidemiological studies have provided compelling evidence that the risk of many adult diseases is strongly shaped by growth patterns in early life. These findings have much in common with those of evolutionary ecology, which has demonstrated the contribution of developmental plasticity to reproductive fitness and longevity. The pivotal connection between these perspectives lies in the metabolism of hormones such as insulin, leptin and cortisol, which on the one hand orchestrate life history flexibility in the face of ecological stresses, and on the other hand give rise to the metabolic syndrome when perturbed in modern environments.

Session 4: Implications and future perspectives

20. Early in life experience and heredity Étienne Danchin

During the last decade the claim that we should rethink heredity occurred as a consequence of or in parallel with the realization of the importance of nongenetic inheritance. I will illustrate a major characteristic of nongenetic inheritance, which is that its influence occurs mostly very early in life in most animals, an even within the womb in mammals. Furthermore, in some cases we have strong evidence that events that occurred a long time before fertilization significantly influence the development of several subsequent generations, thus clearly illustrating the fact that the phenotype not only recapitulates the experiences of the concerned individual but also that of its recent and less recent ancestors.

21. Early life programming - current knowledge and future potential

Susan E Ozanne Over twenty years ago epidemiological studies revealed an association between patterns of early growth and risk of type 2 diabetes and cardiovascular disease. This led to the developmental origins of health and disease hypothesis– the concept that the environment experienced during critical periods of development has a permanent impact on our long-term health. Animal models have been invaluable in identifying underlying mechanisms. Three mechanisms have emerged: (i) permanent structural changes (ii) accelerated cellular ageing and (iii) epigenetic programming. Further understanding of these mechanisms will give the potential to develop markers of disease risk and help design of rational intervention strategies.

22. What is this thing called developmental programming?

Tobias Uller

The evidence for long-term effects of early life experiences has given rise to the concept of ‘developmental programming’. The program metaphor invokes notions of determinism and adaptation – both key components of evolutionary theory. But what are the mechanisms of programming and how are these related to models of phenotypic evolution? Here I show how patterns of programming can arise from a number of different developmental processes, and discuss how these fit with contemporary perspectives on adaptive plasticity in the evolutionary sciences.

23. Evolutionary insights for global health

Mark Hanson

The simplest view of the epidemic of non-communicable diseases is that their origins lie in our evolved genotype plus unhealthy lifestyle. From this point of view classical evolutionary theory explains much of the risk. But, thankfully, it isn't that simple. From a global health perspective, attention is shifting to the epigenetic and other processes underlying plasticity during development - and later - and this offers new insights into biomarkers and interventions.

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Map of Falmouth and local information

Follow coastal path or Swanpool Hill road for Swanpool beach, nature reserve and cafe

St Michael’s Hotel wifi code: Falmouth

Follow coastal path or Pendennis rise for castle and viewpoint.

Taxis: Abacus: 01326 212141 Able: 01326 373007 Bizzy Bees: 01326 317898 Century: 01326 212000

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List of Participants

Stuart Bearhop University of Exeter, UK S.Bearhop@exeter.ac.uk Jon Blount University of Exeter, UK J.D.Blount@exeter.ac.uk Darren Croft University of Exeter, UK D.P.Croft@exeter.ac.uk

Michael Cant University of Exeter, UK M.A.Cant@exeter.ac.uk

Katie Cooke University of Exeter, UK K.Cooke@exeter.ac.uk

Feargus Cooney University of Exeter, UK fc251@exeter.ac.uk Étienne Danchin CNRS/Université Paul Sabatier, France etienne.danchin@univ-tlse3.fr Mandy Drake University of Edinburgh, UK mandy.drake@ed.ac.uk Sinead English University of Oxford, UK sineadenglish@cantab.net Caroline Fall University of Southampton, UK chdf@mrc.soton.ac.uk Willem Frankenhuis Radboud University Nijmegen, The Netherlands w.frankenhuis@psych.ru.nl Ton Groothuis University of Groningen, The Netherlands a.g.g.groothuis@rug.nl

Mark Hanson University of Southampton, UK m.hanson@soton.ac.uk Michelle Hares University of Exeter, UK M.C .Hares@exeter.ac.uk Adam Hayward University of Edinburgh, UK adam.hayward@ed.ac.uk Emma Inzani University of Exeter, UK eli203@exeter.ac.uk Bram Kuijper University College London, UK a.kuijper@ucl.ac.uk Sophia Lavergne University of Toronto, Canada sophia.lavergne@mail.utoronto.ca Nora Lewin Michigan State University, USA nora.lewin@gmail.com Harry Marshall University of Exeter, UK H.Marshall@exeter.ac.uk John McNamara University of Bristol, UK john.mcnamara@bristol.ac.uk Alex Mesoudi University of Exeter, UK A.Mesoudi@exeter.ac.uk Jonathan Mill University of Exeter/King’s College London, UK J.Mill@exeter.ac.uk Daniel Nettle University of Newcastle, UK daniel.nettle@ncl.ac.uk

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Susan Ozanne University of Cambridge, UK seo10@mole.bio.cam.ac.uk Ursula Paredes University of Oxford, UK ursula.paredes@zoo.ox.ac.uk Sarah Paul University of Exeter, UK S.C.Paul@exeter.ac.uk Nick Royle University of Exeter, UK N.J.Royle@exeter.ac.uk Andy Russell University of Exeter, UK A.Russell@exeter.ac.uk

Rebecca Sear London School of Hygiene and Tropical Medicine, UK rebecca.sear@lshtm.ac.uk Barbara Taborsky University of Bern, Switzerland barbara.taborsky@iee.unibe.ch

Faye Thompson University of Exeter, UK F.J.Thompson@exeter.ac.uk Tobias Uller University of Oxford, UK tobias.uller@zoo.ox.ac.uk Maddy Vierbuchen University of Bern, Switzerland susanna.vierbuchen@iee.unibe.ch Emma Vitikainen University of Exeter, UK E.I.K.Vitikainen@exeter.ac.uk Lindsay Walker University of Exeter, UK L.Walker@exeter.ac.uk Jonathan Wells University College London, UK jonathan.wells@ucl.ac.uk Thomas Williams University of Edinburgh, UK thomas.williams@cantab.net Andy Young University of Exeter, UK A.J.Young@exeter.ac.uk

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Workshop Sponsors

VWR Jencons Capital Equipment Specialist Mobile. 07827977705 Email. robin.lancashire@jencons.co.uk www.uk.vwr.com

Sartorius Lab Products and Services Adam Green Area Manager – South Wales & South West Mobile: 0044 (0) 7900 828991

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