Endocrine Axis: HP-prolactin/growth Negative and positive Endocrine regulation: 1. Larval growth high: +ve PrL, -ve TRH inhibited Metabolic effect Growth hormones Endocrine Axis Thyroid-Growth +ve +ve Prl -ve 3. Prl 2.Growth drops slows T3 -ve Endocrine T3 Cascade: Tissue specific Aquatic larval phase Monoidonase TSH Endocrine Organ: PrRH 4.TRH inhibition released LEGEND Hypothalmus (PrRH or TRH) Pituitary (Prl or TSH) +ve Endocrine Axis HP-thyroid T4 α and β nuclear receptors (retinoid X) Thyroid Prolactin Releasing Hormone Prolactin T3 induces Metamorphosis Tissue specific effects: cell death (gills, tail fin) pigmentation (melanin) lung development skull remodeling hyoid remodeling Prl Thyroxine Releasing Hormone TRH Thyroxine Stimulating Hormone TSH Thyroxine gene transcription: reorganize larval form PrRH T3, T4 Hierarchical Structure of the Endocrine System: 1. Endocrine Organ Terrestrial adult phase Adult 2. Endocrine Cascade 3. Positive-Negative +ve Feedback Loops are made up of 3 -ve endocrine axes a) EVOLUTION of LIFE CYCLE STATES of the SALAMANDER FAMILIES Yolky Eggs Aquatic Larvae M M* Desmognathus M Metamorphosis P Paedomorphosis D Direct Development Terrestrial Adult Aneides Terrestrial Adult Ensatina D Plethodon Eurycea Plethodontidae (M, P, D) P Paedomorph Pseudotriton Bolitoglossa Thorius D Terrestrial Adult Batrachoseps Amphiumidae (P) P Adult Ambystoma M Axolotl Adult Ambystoma Aquatic Larvae Paedomorphic Adult Axolotl P M Salamandridae (M, P) Hynobiidae (M) Anura (Frogs) Larva Dicamptodontidae (M, P) Adult Larvae Cryptobranchidae (P) Sirenidae (P) P OUTGROUPS Eurycea Ambystomatidae (M, P) Proteidae (P) P M Rhyacotritonidae (M) Aneides Hyrdomantes Ensatina Anura (Frogs) Gymnophiona (Caecelians) Desmognathus Hemidactylium Bolitoglossa Bolitoglossa Option 2 Free Tongue Adult Miniaturisation Larva Adult M* M M D Option 1 Free Tongue Embryo D D Larva M Adult Larva M Adult Embryo Atavism Hypothetical Ancestral State Thorius Adult Option 2 Free Tongue b) TONGUE and HYOID EVOLUTION in the PLETHODONTID SALAMANDERS a. Ancestral Complex Metamorphosis b Collapse of adult trade-offs in paedomorphs LARVAL GROWTH Metabolic effect Growth hormone WATER DRIVE PrRH Prl Prl PrRH Hypothalmus Pituitary TRH -ve TSH Metabolic effect PROLACTIN Prl LARVAL GROWTH WATER DRIVE Metabolic effect PROLACTIN Metabolic effect PROLACTIN T3 -ve MUTATION Thyroid Adrenal Aquatic larval phase Metamorphosis size timetludA larval survival Thyroid LEGEND Gonad Reproductive Effort Adrenal Prolactin PrRH Releasing Hormone T3 Prolactin Prl Thyroxine T3, T4 Hierarchical Structure of the Endocrine System: Migratory distance 1. Endocrine Organ 2. Endocrine Cascade T3 disabled Aquatic adult phase Migration Life history trade-offs collapse d. Complex metamorphosis with an alternative paedomorph strategy LARVAL GROWTH Metabolic effect Growth hormone Behavioral effect PROLACTIN WATER DRIVE Prl Prl -ve TSH TSH PrRH PrRH Hypothalmus Pituitary TRH Prl Metabolic effect PROLACTIN T3 -ve Thyroid Thyroid Adrenal size Gonad Reproductive Effort Adrenal Aquatic larval phase Metamorphosis size No T3 Terrestrial adult phase Adult survival Maternal Care Trade-offs associated with adult reproduction such as offspring quality and care are elaborated Gonad Reproductive Effort T3 quantity quality time Life history trade-offs associated with aquatic larvae collapse quantity quality Adult survival aquatic adult survival MATERNAL CARE HP Metamorphosis tludA time Terrestrial adult phase Life History Trade-off c. Adult Trade-offs Associated with Direct Development Metabolic effect Growth hormone Gonad Reproductive Effort T3 larval survival 3. Positive-Negative +ve Feedback Loops -ve Life history trade-offs associated with complex life cycles LARVAL GROWTH Aquatic larval phase size Terrestrial quantity quality adult phase Adult survival larval survival HP larval survival timetludA Terrestrial quantity quality adult phase Adult survival Aquatic adult phase aquatic adult survival Migratory distance Life history trade-offs associated with complex life cycles ROAD MAP FOR ENDOCRINE EPISTASIS: Physiological causes of life history plasticity and trade-offs and alternative pathways in larval spadefoot toads 2. Biotic Inputs (e.g., crowding) 1. Metabolic Inputs (e.g., growth hormone and prolactin) PrRH Hypothalamus- Prl +ve 3. Abiotic Inputs (e.g., water level) TRH Prl TSH Pituitary CRH ACTH TSH +ve -ve -ve T3 B +ve T3 -ve T3 Monoidonase Tissue specificity T4 α and β nuclear receptors (retinoid X) Thyroid B gene transcription Eat Fairy Shrimp Exogenous T 3 I nd uc ti Life history plasticity on e ivor Carnivore LEGEND keratenized beak Hypothalamus-Pituitary Hormones: jaw muscles larval survival gut length cannibalism vs. social Slow development Fast Thyrotropin Releasing Hormone Corticotropin Releasing Hormone Prolactin Releasing Hormone Thyroid Stimulating Hormone Adrenocorticotropic hormone Prolactin Steroid Hormones: Corticosterone B kin altruism TRH CRH PrRH TSH ACTH Prl Octopeptides: Thyroxines T3 T4 Hierarchical Structure of the Endocrine System: rate size Gonadal rn Ca INDUCED TRAITS: Adrenal of Alternative pathways for larval development: Omnivore B Stress 1. Endocrine Regulatory System/Organ(s) time 2. Endocrine Cascade juvenile survival larval survival and size and time to metamorphosis and survival costs of kin altruism LARVAL LIFE HISTORY TRADE-OFFS SOCIAL TRADE-OFFS 3. Positive-Negative Feedback Loops Life History Trade-off -ve +ve ROAD MAP FOR ENDOCRINE EPISTASIS: Life-history trade-offs and plasticity in reproduction of side-blotched lizards Biotic Inputs Social crowding by female morphs LEGEND Hypothalamus-Pituitary Hormones: Gonadotropin Releasing Hormone GnRH Thyrotropin Releasing Hormone TRH Corticotropin Releasing Hormone CRH Abiotic Inputs (e.g., photoperiod triggers reproduction) Thyroid Stimulating Hormone Follicle Stimulating Hormone Luteinising Hormone Adrenocorticotropic hormone HypothalamusPituitary FSH LH ACTH Life history plasticity Steroid Hormones: Corticosterone B Testosterone T Estrogen E GnRH CRH +ve +ve -ve B Stress -ve E T B Energy +ve Adrenal Hierarchical Structure of the Endocrine System: FSH LH 1. Endocrine Regulatory System/Organ(s) Gonadal Life history tradeoffs Liver cells E Immune function Cell-mediated Humoral FSH 3. Positive-Negative Feedback Loops Vitellogenin Reproductive Effort MHC 2. Endocrine Cascade Follicles FSH ERE B TSH FSH LH ACTH -ve +ve Life History Trade-off Yolk Yolk Clutch Size Egg Size Progeny Survival dispersal philopatry altruism Female Survival COSTS OF REPRODUCTION TRADE-OFF OFFSPRING QUANTITY VS. QUALITY TRADE-OFF SOCIAL TRADE-OFFS Density-dependent survival trade-offs r-strategist oo genotype K-strategists yy genotype bb genotype