Mechanisms of life history evolution: the genetics and physiology of life history traits and trade-offs / edited by Thomas Flatt, Andreas Heyland

Incluye bibliografía: páginas 380-468 e índice: páginas 469-478

Preface.. List of contributors.. Part 1: Integrating mechanisms into life history evolution.. 1 Integrating mechanistic and evolutionary analysis of life history variation.. 1.1 Introduction.. 1.2 The life history framework.. 1.2.1 What is a life history?.. 1.2.2 Life history traits and fitness.. 1.2.3 Trade-offs and constraints.. 1.2.4 Empirical approaches in life history research.. 1.3 The study of causal mechanisms linking genotype to phenotype.. 1.4 How can mechanistic insights contribute to understanding life history evolution?.. 1.4.1 Why understanding mechanisms is important for answering evolutionary questions.. 1.4.2 The molecular identity and function of genes that affect life history.. 1.4.3 Are candidate life history genes ecologically and evolutionarily relevant?.. 1.4.4 How do trade-offs work?.. 1.5 Conclusions.. 2 Genomic insights into life history evolution.. 2.1 Introduction.. 2.2 Genomic analysis of trade-offs.. 2.2.1 Case Study 1: A transgenic analysis of the cost of resistance in Arabidopsis thaliana.. 2.2.2 Case Study 2: A QTL analysis of the cost of resistance to parasite infection in Tribolium.. 2.2.3 Case Study 3: A microarray analysis implicating a single gene in the cost of resistance to DDT in Drosophila melanogaster.. 2.2.4 Case Study 4: A microarray analysis of antagonistic pleiotropy and gene expression in Drosophila melanogaster.. 2.3 To what extent is the phenotype determined by different molecular/developmental mechanisms?.. 2.3.1 Comparisons among species.. 2.3.2 Comparisons among natural populations of the same species.. 2.3.3 Arti' cial selection experiments.. 2.3.4 A proposed experiment and predictions.. 2.4 Summary.. 2.5 Acknowledgments.. Part 2: Growth, development, and maturation.. 3 Emerging patterns in the regulation and evolution of marine invertebrate settlement and metamorphosis.. 3.1 Background.. 3.2 Introduction to marine invertebrate life histories

3.3 Regulation of larval development and the evolution of feeding modes in echinoids: Energy allocation trade-offs during larval development.. 3.3.1 Hormonal regulation of juvenile development.. 3.3.2 Hormonal signaling and the evolution of alternative life history modes.. 3.4 Mechanisms underlying larval settlement and the evolution of alternative settlement strategies: Signal detection and modulation during settlement.. 3.4.1 The sensory system: Cues, receptors, and signal transduction mechanisms.. 3.4.2 The competence system.. 3.5 Settlement strategies: Evolution of sensory structures and signaling networks.. 3.6 Future directions.. 3.7 Summary.. 3.8 Acknowledgments.. 4 Evolution and the regulation of growth and body size.. 4.1 Introduction.. 4.2 The regulation of body size in insects.. 4.2.1 The regulation of critical size.. 4.2.2 The regulation of TGP.. 4.3 The regulation of growth rate.. 4.4 Environmental variation in body size: The functional interaction between critical size, TGP, and growth rate in insect size regulation.. 4.5 Trade-offs between body size and other traits.. 4.6 The evolution of body size.. 4.6.1 Evolutionary trends.. 4.6.2 Artificial selection.. 4.6.3 The developmental mechanisms underlying the evolution of body size.. 4.6.4 The relationship between evolutionary and environmental variation in body size.. 4.6.5 Can we predict which size-regulatory mechanisms are the target for selection on body size?.. 4.7 Summary.. 4.8 Acknowledgments.. 5 The genetic and endocrine basis for the evolution of metamorphosis in insects.. 5.1 Introduction.. 5.2 Endocrine regulation of metamorphosis.. 5.3 Comparative endocrinology across insect life history strategies.. 5.4 Endocrine titers and cuticle progression during embryonic development.. 5.5 Comparison of hemi- and holometabolous endocrine events during postembryonic development.. 5.6 The "status-quo" action of juvenile hormone and its signal transduction

5.7 The broad gene and specification of the pupal stage.. 5.7.1 Molecular aspects of Broad action.. 5.7.2 A broad-based view of the pronymph hypothesis.. 5.7.3 The appearance of broad in the arthropods.. 5.8 Summary.. 5.9 Acknowledgments.. 6 Thyroidal regulation of life history transitions in fish.. 6.1 Introduction.. 6.2 Fish ontogeny and life history transitions.. 6.3 Overview of the hypothalamic-pituitary-thyroid axis.. 6.4 The hypothalamic-pituitary axis.. 6.5 Thyroid tissue and hormone synthesis.. 6.5.1 Serum thyroid hormone distributor proteins, cellular uptake, and cytosolic transport.. 6.5.2 Thyroid hormone deiodinases.. 6.5.3 Thyroid hormone nuclear receptors.. 6.6 Thyroidal regulation of fish ontogeny and life history transitions.. 6.6.1 Embryogenesis and embryo to larval transitions.. 6.6.2 Larval to juvenile transitions.. 6.6.3 First or "true" metamorphoses in ray-finned fish.. 6.6.4 First or "true" metamorphosis in lampreys (Agnatha.. 6.6.5 Smoltification: A juvenile transition in salmonids.. 6.7 Summary.. 6.8 Acknowledgments.. 7 Hormone regulation and the evolution of frog metamorphic diversity.. 7.1 Introduction.. 7.2 Ecological context of metamorphic life history evolution.. 7.2.1 Escape from the growth versus development trade-off.. 7.3 Key concepts in the endocrinology of metamorphosis.. 7.3.1 Overview of the endocrinology of metamorphosis.. 7.3.2 Tissue sensitivity and tissue-specific responses to thyroid hormones.. 7.3.3 Tissue developmental asynchrony.. 7.4 Endocrine basis of amphibian life history evolution.. 7.4.1 Larval period duration.. 7.4.2 Size at metamorphosis.. 7.4.3 Direct development.. 7.4.4 Neoteny.. 7.5 Molecular mechanisms of peripheral control: Potential evolutionary targets underlying diversity in larval period diversity.. 7.5.1 Thyroid hormone transporters.. 7.5.2 Thyroid hormone metabolizing enzymes.. 7.5.3 Cytosolic thyroid hormone binding proteins

7.5.4 Thyroid hormone receptors.. 7.5.5 Modulation of thyroid hormone responsiveness by corticosterone and prolactin.. 7.6 Conclusions.. 7.7 Summary.. Part 3: Reproduction.. 8 Asexual reproduction in Cnidaria: Comparative developmental processes and candidate mechanisms.. 8.1 Introduction.. 8.2 Diversification of clonal reproduction in cnidarians.. 8.2.1 Diversity of clonal reproduction modes in cnidarian polyps.. 8.2.2 The role of developmental modularity in life history diversification.. 8.2.3 Evidence of modularity in cnidarian developmental programs.. 8.2.4 Elucidating the genetic architecture of cnidarian modules.. 8.3 Trade-offs and environmental signaling in asexual reproduction.. 8.4 Trade-offs between methods of asexual reproduction.. 8.5 Environmental signals and reception in cnidarian asexual reproduction.. 8.6 Looking ahead: Combining signaling with developmental mechanisms.. 8.7 Summary.. 8.8 Acknowledgments.. 9 The genetics and evolution of flowering time variation in plants: Identifying genes that control a key life history transition.. 9.1 Introduction.. 9.2 The natural and laboratory history of Arabidopsis.. 9.3 The molecular genetics of flowering time.. 9.3.1 Getting at the mechanistic basis: Genes controlling flowering time variation and what they do.. 9.3.2 CRY2.. 9.3.3 PHYC.. 9.3.4 FRI.. 9.3.5 FLC.. 9.4 Epistatic effects among FRI and FLC.. 9.5 Pleiotropic effects of genes controlling flowering time variation.. 9.6 Comparative functional genomics: The genetics of flowering time in other species.. 9.7 Synthesis and prospectus.. 9.8 Summary.. 9.9 Acknowledgments.. 10 Mechanisms of nutrient-dependent reproduction in dipteran insects.. 10.1 Introduction.. 10.2 Larval nutrition and reproduction.. 10.2.1 Ovary size.. 10.2.2 Meal size.. 10.2.3 The effects of mate size.. 10.2.4 Larval nutrition and teneral reserves.. 10.3 Adult-acquired resources.. 10.3.1 Hunger.. 10.3.2 Finding nutrition.. 10.3.3 Oogenesis and ovulation

10.4 The evolutionary genetics of reproduction: Future prospects.. 10.5 Summary.. 10.6 Acknowledgments.. 11 Mechanisms underlying reproductive trade-offs: Costs of reproduction.. 11.1 Introduction.. 11.2 Key life history traits and costs of reproduction.. 11.3 Intrinsic costs of reproduction: Trade-offs between reproductive activity and survival or future reproductive rate.. 11.3.1 Physiological costs of reproduction.. 11.3.2 Evolutionary costs of reproduction.. 11.3.3 Mechanisms underlying reproductive costs.. 11.3.4 Nutrients, nutrient sensing, and costs of reproduction between reproductive rate and lifespan.. 11.3.5 The presence of a germ line and costs of reproduction between reproductive rate and lifespan.. 11.4 Reproductive hormones as mediators of trade-offs between reproductive rate and lifespan.. 11.5 Male seminal fluid proteins as mediators of trade-offs between reproduction and lifespan in females.. 11.6 The immune system as a mediator of costs between current reproductive rate and survival.. 11.7 Damage as a mediator of trade-offs between current reproductive rate and survival.. 11.8 Resource allocation: Allocation versus adaptive signaling.. 11.9 Costs of reproduction in a fitness-based framework.. 11.10 New directions.. 11.10.1 Mechanistic data are incomplete.. 11.10.2 The evolution and conditional economics of reproductive costs.. 11.10.3 Integration of life history data from social species.. 11.11 Summary.. 11.12 Acknowledgments.. 12 Patterns and processes of human life history evolution.. 12.1 The evolution of human life histories.. 12.1.1 Ecological dominance: Lowered mortality, better food and tools, and increased sociality.. 12.1.2 Human cognitive evolution.. 12.1.3 Prolonged development.. 12.1.4 High fertility, biparental and alloparental care.. 12.2 Proximate mechanisms of human life history patterns.. 12.2.1 Reproductive development.. 12.2.2 Ovarian and testicular functions.. 12.2.3 Reproductive behaviors

12.2.4 Reproductive senescence.. 12.3 Summary.. Part 4: Lifespan, aging, and somatic maintenance.. 13 Parallels in understanding the endocrine control of lifespan with the firebug Pyrrhocoris apterus and the fruit fly Drosophila melanogaster.. 13.1 Introduction.. 13.2 Reproductive diapause.. 13.3 Reproduction and its trade-offs.. 13.4 Endocrine regulation.. 13.5 Conclusion.. 13.6 Summary.. 13.7 Acknowledgments.. 14 The genetics of dietary modulation of lifespan.. 14.1 Introduction.. 14.2 Calorie restriction as a modulator of life history traits.. 14.2.1 Is lifespan extension due to calorie restriction universal?.. 14.2.2 The difficulty of defining what constitutes calorie restriction.. 14.3 The evolution of dietary restriction and its lifespan-extending effect.. 14.4 Dietary restriction in lower organisms.. 14.4.1 C. elegans.. 14.4.2 D. melanogaster.. 14.5 Dietary restriction in higher organisms.. 14.5.1 Rodents.. 14.5.2 Primates.. 14.6 Concluding remarks.. 14.7 Summary.. 14.8 Acknowledgments.. 15 Molecular stress pathways and the evolution of life histories in reptiles.. 15.1 Reptiles possess remarkable variation and plasticity in life history.. 15.2 The molecular stress networks: What is known in reptiles?.. 15.2.1 Metabolic pathways.. 15.2.2 Molecular mechanisms to regulate the production of reactive oxygen species.. 15.2.3 Molecular mechanisms to neutralize reactive oxygen species.. 15.2.4 Tolerance and resistance to reactive oxygen species.. 15.2.5 Molecular pathways for repair.. 15.2.6 Insulin/insulin-like growth factor signaling pathway.. 15.3 Environmental stress and evolving molecular pathways: Evidence in reptiles.. 15.3.1 Temperature (heat stress.. 15.3.2 Hibernation: Supercooling, freeze tolerance, and anoxia tolerance.. 15.3.3 Dietary stress: Availability and type of food.. 15.3.4 Type of food.. 15.4 Perspective.. 15.5 Summary.. 15.6 Acknowledgments.. 16 Mechanisms of aging in human populations.. 16.1 Introduction

16.2 Mechanisms of aging.. 16.2.1 Insulin/IGF-1 signaling.. 16.2.2 Lipid metabolism.. 16.2.3 Antioxidant enzymes.. 16.2.4 Macromolecule repair mechanisms.. 16.2.5 Cellular responses to damage.. 16.3 Convergence of longevity signals.. 16.3.1 Dietary restriction.. 16.4 Integration of genetic pathways and the environment.. 16.5 Summary.. 16.6 Acknowledgments.. Part 5: Life history plasticity.. 17 Mechanisms underlying feeding-structure plasticity in echinoderm larvae.. 17.1 Introduction.. 17.2 Plasticity of feeding structures.. 17.3 Evidence for adaptive plasticity.. 17.4 Developmental regulation.. 17.5 Mechanisms of perception.. 17.6 Mechanisms of morphological response.. 17.7 Integrative response.. 17.8 Future directions.. 17.9 Summary.. 17.10 Acknowledgments.. 18 Evolution and mechanisms of insect reproductive diapause: A plastic and pleiotropic life history syndrome.. 18.1 Introduction.. 18.2 Advances and methods.. 18.2.1 Clines.. 18.2.2 Temporal variation and seasonality.. 18.3 Diapause as a model system for life history evolution.. 18.4 Identifying genes for seasonality.. 18.4.1 Dormancy in D. melanogaster.. 18.4.2 Genes for diapause in Drosophila.. 18.5 Pathway and genomic analyses.. 18.5.1 Diapause and insulin signaling.. 18.5.2 Expression analyses.. 18.6 Summary.. 18.7 Acknowledgments.. 19 Seasonal polyphenisms and environmentally induced plasticity in the Lepidoptera: The coordinated evolution of many traits on multiple levels.. 19.1 Introduction.. 19.2 Frameworks for dissecting the evolution of polyphenisms.. 19.3 Case studies on the adaptive nature of seasonal polyphenisms.. 19.4 Environmental cues and the physiological regulation of plasticity.. 19.5 Genetics of the evolution of the seasonal polyphenism in wing pattern.. 19.6 Life history evolution in polyphenic butterflies.. 19.7 Perspectives: Suites of adaptive traits in combination with an ability to acclimate.. 19.8 Summary.. 19.9 Acknowledgments

20 Honey bee life history plasticity: Development, behavior, and aging.. 20.1 Introduction.. 20.2 Development.. 20.2.1 Female caste morphology: Physiology, function, and reproduction.. 20.2.2 An integrative molecular model for caste development: Differential nutrition during larval development triggers caste differentiation.. 20.3 Behavioral maturation and specialization.. 20.3.1 Specialization of foraging behavior.. 20.3.2 Central nervous system changes during behavioral maturation.. 20.3.3 Metabolic changes during behavioral maturation.. 20.4 Worker aging.. 20.4.1 Plasticity of aging.. 20.4.2 Oxidative stress.. 20.4.3 Metabolic patterns of senescence.. 20.4.4 Cognitive senescence.. 20.4.5 Impact of nutrition sensing pathways on lifespan.. 20.5 Concluding remarks.. 20.6 Summary.. 20.7 Acknowledgments.. Part 6: Life history integration and trade-offs.. 21 Molecular mechanisms of life history trade-offs and the evolution of multicellular complexity in volvocalean green algae.. 21.1 Introduction.. 21.2 The volvocalean green algal group.. 21.2.1 Overview.. 21.2.2 Life history trade-offs and the evolution of multicellularity in volvocalean algae.. 21.3 Mechanisms of life history trade-offs and the evolution of multicellularity in volvocalean algae.. 21.3.1 Overview.. 21.3.2 Acclimation and life history trade-offs in Chlamydomonas.. 21.3.3 The genetic basis for cell differentiation in Volvox carteri.. 21.4 Co-opting mechanisms underlying environmentally induced life history trade-offs for cell differentiation.. 21.5 Conclusion.. 21.6 Summary.. 21.7 Acknowledgments.. 22 Molecular basis of life history regulation in C . elegans and other organisms.. 22.1 Introduction.. 22.2 C. elegans life history.. 22.3 Genetics of dauer formation.. 22.4 Dauer pheromone.. 22.5 Neurosensory signaling and processing.. 22.6 Sensory signal transduction and longevity.. 22.6.1 Insulin/IGF-1 signal transduction.. 22.6.2 TGF-beta signaling

22.6.3 Biogenic amine signaling.. 22.6.4 Neuropeptide-Y-like signaling.. 22.6.5 Steroid hormone signaling.. 22.7 Developmental timing and life history specification.. 22.8 Reproduction and longevity.. 22.9 Dietary restriction.. 22.10 Diapause in other nematode strains.. 22.11 D. melanogaster reproductive diapause.. 22.12 Torpor/hibernation of mammals.. 22.13 Prospectus.. 22.14 Summary.. 22.15 Acknowledgments.. 23 The costs of immunity and the evolution of immunological defense mechanisms.. 23.1 Introduction.. 23.2 Innate immune defense in Drosophila.. 23.3 Trade-offs between reproduction and immunity.. 23.4 Deployment costs, tolerance, and the evolution of immune regulation.. 23.5 Multiple-fronts costs of immunity.. 23.6 Future directions.. 23.7 Summary.. 24 Intermediary metabolism and the biochemical-molecular basis of life history variation and trade-offs in two insect models.. 24.1 Introduction.. 24.2 Gryllus firmus : Biochemical and molecular studies of trade-offs in lipid metabolism and life histories.. 24.2.1 Background on life history variation in Gryllus and methodological perspective.. 24.2.2 Lipid reserves: The physiological context of biochemical studies of life history trade-offs.. 24.2.3 Morph-specific differences in flux through pathways of lipid biosynthesis and oxidation.. 24.2.4 Enzymatic basis of flux trade-offs: Digging deeper into the functional hierarchy of life history trade-offs.. 24.2.5 Enzymological and molecular causes of differences in enzyme activities between morphs.. 24.2.6 Amino acid metabolism and life history trade-offs in Gryllus.. 24.3 Drosophila melanogaster.. 24.3.1 Laboratory selection on life history.. 24.3.2 Clinal variation in intermediary metabolism and life history traits in the field.. 24.4 Other studies and issues relevant to Drosophila.. 24.4.1 Additional biochemical and molecular studies.. 24.4.2 Influence of changes in allocation versus nutrient input on life history evolution

24.4.3 Quantitative-genetic variation in enzyme activities and fitness.. 24.5 Summary.. 24.6 Acknowledgments.. 25 Epistatic social and endocrine networks and the evolution of life history trade-offs and plasticity.. 25.1 Introduction.. 25.2 Endocrine networks and life history trade-offs.. 25.3 An example of a gated switch in developmental life history trade-offs.. 25.4 Social networks and life history trade-offs.. 25.5 Social dimensions to trade-offs: Endocrine mediation and fitness consequences.. 25.6 Corticosterone, egg size, and the trade-off between aspects of offspring quality in a lizard.. 25.7 Juvenile hormone, vitellogenin, and reproductive trade-offs in eusocial honeybees.. 25.8 Testosterone, growth hormone, social dominance, and smolting in Atlantic salmon.. 25.9 Gibberellins, auxin, ethylene, and reproductive allocation in monoecious plants.. 25.10 Conclusions and future directions.. 25.11 Summary.. 26 Hormonally-regulated trade-offs: Evolutionary variability and phenotypic plasticity in testosterone signaling pathways.. 26.1 Introduction.. 26.2 Testosterone and trade-offs.. 26.3 Conservation versus variation in testosterone-regulated traits on an interspecific level.. 26.4 Signal production pathway.. 26.5 Signal transduction pathway.. 26.6 Plasticity of testosterone-regulated trade-offs on an individual level.. 26.7 Costs.. 26.8 Phenotypic plasticity and reaction norms.. 26.9 Development.. 26.10 Future directions of evolutionary endocrinology.. 26.11 Summary.. 26.12 Acknowledgments.. Part 7: Concluding remarks.. 27 Does impressive progress on understanding mechanisms advance life history theory?.. 27.1 Introduction.. 27.2 How research on mechanisms is changing views on life history evolution.. 27.2.1 The nature of trade-offs: Signals, allocation, or both?.. 27.2.2 Ancient, conserved, broadly shared mechanisms?.. 27.2.3 Decoupling functions by duplicating modules.. 27.3 Is work on mechanisms changing theory?

27.3.1 Are these data forcing the theory to change?.. 27.3.2 Are we identifying general features of intermediate structure in the genotype-phenotype map?.. 27.3.3 Are the empirical problems with the theory being addressed?.. 27.4 Conclusion.. 27.5 Summary.. 28 What mechanistic insights can or cannot contribute to life history evolution: An exchange between Stearns, Heyland, and Flatt.. 28.1 Why mechanisms are important for life history theory: A response by Flatt and Heyland.. 28.2 Reply by Stearns: Mechanisms do not yet force the theory to change.. References.. Index

Life history theory seeks to explain the evolution of the major features of life cycles by analyzing the ecological factors that shape age-specific schedules of growth, reproduction, and survival and by investigating the trade-offs that constrain the evolution of these traits. Although life history theory has made enormous progress in explaining the diversity of life history strategies among species, it traditionally ignores the underlying proximate mechanisms. This novel book argues that many fundamental problems in life history evolution, including the nature of trade-offs, can only be fully resolved if we begin to integrate information on developmental, physiological, and genetic mechanisms into the classical life history framework. Each chapter is written by an established or up-and-coming leader in their respective field; they not only represent the state of the art but also offer fresh perspectives for future research. The text is divided into 7 sections that cover basic concepts (Part 1), the mechanisms that affect different parts of the life cycle (growth, development, and maturation; reproduction; and aging and somatic maintenance) (Parts 2-4), life history plasticity (Part 5), life history integration and trade-offs (Part 6), and concludes with a synthesis chapter written by a prominent leader in the field and an editorial postscript (Part 7). eng