Bio205H5: lecture 2 physiological ecology tolerance intolerance Q: How does the ability of an organism to cope with its abiotic environment influence its distribution, abundance and interactions with other species? stress intolerance ecophysiology = physiological responses of individual organisms to abiotic factors organism response stress low factor high 2 phenotypic plasticity: responses to abiotic factors: • genotypic (selection → adaptation) • phenotypic (plasticity; reaction norms) 3 4 reaction norms • reaction norm = range of phenotypes produced by a genotype under different environmental conditions (phenotypic plasticity) • ideally: look at genetically identical individuals over range of conditions 5 6 1 reaction norms: 7 comparing reaction norms: 8 genetic x environment interaction: Genotype 2 Genotype 1 e.g. height phenotype: phenotype Genotype 1 Genotype 2 environment 9 time scales of responses to abiotic factors: environment 10 developmental responses: 1. Evolutionary - over generations (∆ in allele frequencies) - genotypic (adaptations) 2. Developmental - within lifespan of organism - abiotic factors can irreversibly influence development of an individual • responses to environmental conditions during ontogeny (development) e.g. temperature - dependent sex determination (TSD) most lizards, turtles, crocodiles 11 12 2 e.g. reproductive maturation in whitefooted mice: day length: • affects testicular development in juveniles • some breed in year of birth; some don’t 13 - 3. Biochemical / physiological timescale measured in units much less than the lifespan of an individual • acute responses (e.g. transient increase in metabolic rate with temp) • gradual responses (e.g. seasonal changes in metabolic rate with temp) acclimation response 15 How do organisms cope with abiotic factors? • 14 results from exposure to new conditions may be regular or unpredictable changes seasonal; daily; etc. reversible e.g. Arctic fox: summer phase winter phase 16 1. Avoiders • in space: alter microhabitat alter macrohabitat (migrates) 3 main strategies: 1. avoidance 2. conformity 3. regulation • in time: e.g. diapause; torpor 17 18 3 conformers: 2. Conformers: undergo changes of internal state similar to external environment 3. Regulators: maintain internal environment regardless of external conditions (homeostasis) 19 20 mixed strategies: regulators 21 22 temperature relations • temperature has profound effect on organisms • climate broadly defines species distribution: e.g. trees in N. Am. 23 24 4 • conduction: movement of heat between solids in contact Q: How does heat transfer between organisms and environment? • convection: movement of heat between solids and fluid (gas or liquid) Heat (energy) transferred by: 1. conduction 2. convection 3. radiation • radiation: transfer of heat through electromagnetic waves (infrared) 25 heat gain and heat loss 26 evaporation solar radiation H s = H m ± H cd ± H cv ± H r − H e radiation from vegetation convection radiation from animal s = stored heat m = metabolic heat cd = conduction cv = convection r = radiation e = evaporation (1-way transfer) reflected solar radiation 27 conductance from ground radiation from ground conductance to ground 28 effects of temperature psychrophiles -50°C thermophiles 100°C • most organisms: much narrower range! 29 30 5 extreme temperatures: • many species have broad geographic distributions extreme low temp: freezing of body fluids - ice crystals damage cells - osmotic movement of water → ECF precipitates enzymes • many live in seasonal environments • how do they cope with temp extremes and temp variation? extreme high temp: denaturation of enzymes, DNA - brain damage, middle ear (equilibrium) no vertebrate can tolerate ≥ 50°C 31 32 temperature affects physiological processes • reaction rate: R2 = R1 ( Q10 ) Q10 = T2 − T1 10 increase in reaction rate for every 10°C increase in temperature often ~ 2 - 3 33 34 metabolic rate and temperature temp effects on plants: • • effect on photosynthetic rate • metabolic rate → physiological “performance” enzymes function best in narrow range of temp How to ensure that physiological performance is optimal over a range of temperatures? 35 36 6 adaptations to thermal regime recall: 3 strategies 1. avoid • don’t function under temp extremes poikilotherm = thermal conformer - body temp ≈ ambient temp - body temp varies 2. conform • body temp ≈ ambient; function maintained homeotherm = thermal regulator - body temp ≈ constant (usually > ambient) 3. regulate • maintain body at optimal temp 37 contrast: variability (constancy) of body temp 38 however..... ectotherm = body heated externally - e.g. basking etc. ectothermic homeotherms = body temp constant, but dictated by env’t endotherm = body heated internally e.g. deep sea fish contrast: source of body heat endothermic poikilotherms = body heated internally, but varies usually : poikilotherm = ectotherm homeotherm = endotherm e.g. naked mole-rat 39 Ectotherms: don’t regulate temp internally: 40 avoid activity in extreme temps • dormancy burrow under frost line, underwater • freeze tolerance e.g. wood frogs: can’t dig; hibernate in leaf litter; actively control freezing: proteins nucleate ice crystals in ECF 1. avoid extreme temperature • in space: migration e.g. monarch butterfly 41 42 7 oxygen consumption Consider: fish at 5°C; acute change to 25°C acute response: increase in metabolic rate with increasing temp 100 10 Oxygen Consumption 2. conform: maintain function at “sub-optimal” temps fish allowed to acclimate to 25°C 100 full compensation 10 0 10 20 30 40 Temperature °C 0 10 20 30 40 temperature °C 43 44 • molecular/cellular changes can permit physiological function at varying temperatures compensation involves molecular changes • expression of isozymes (alternate forms of enzymes) • e.g. goldfish: alter expression of heavy and light myosin chains in muscle at low temp • changes in abundance of mitochondria 45 46 3. regulate: • ectotherms do not regulate temperature internally • manipulate heat exchange with environment • compensation allows ectotherms to avoid effects of temp on metabolic rate - not sluggish in polar regions - not hyperactive in tropical regions in ectotherms, metabolic rate can be independent of ambient temp (not true of endotherms!) basking to warm 47 burrowing to cool can maintain optimal temperatures through behavioural adaptations 48 8 plants are poikilothermic ectotherms • usually do not produce significant amounts of metabolic heat • 1. 2. 3. most terrestrial env’t fluctuate in temp avoid conform regulate 49 50 conform: • molecular adaptations that permit photosynthesis at temp extremes plants can avoid: • grow in microhabitats that are sheltered from prevailing conditions e.g. clover: arctic saxifrage produces cyanide in response to cold temp moist slope has trees, dry slope doesn’t protects from freezing 51 52 regulate: • morphology & behaviour regulate heat exchange with environment • alter Hr = heat from radiation • alter Hcv = heat from convection • alter Hcd = heat from conduction 53 54 9 • some plants use metabolic heat to warm flowers • e.g. skunk cabbage: large root stores starch • moved to inflorescence • burned at high rate; generates heat • regulates temp 55 56 next lecture: • temperature and water relations continued • introduction to life-histories (Chapter 12) 57 10
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