Evolutionary physiology topics 1. Patterns 2. Processes Evolutionary physiology topics 1. Patterns • How and why of particular transitions How and why did endothermic vertebrates evolve from ectothermic ancestors? Endothermy versus ectothermy scala naturae Endothermy versus ectothermy Endothermy versus ectothermy Advantages of endothermy: • Independent of environment • Stenothermy • Aerobic metabolism Endothermy versus ectothermy Advantages of ectothermy: • Low energetic requirements metabolism (Wg-1day-1) 2 1.8 1.6 1.4 mammals 1.2 Passerine birds 1 reptiles 0.8 0.6 0.4 0.2 0 0.1g 10g 1kg 100kg 1000kg Endothermy versus ectothermy Advantages of ectothermy: • Low energy requirements Low food habitats Endothermy versus ectothermy Advantages of ectothermy: • Low energy requirements Low food habitats Fluctuating food habitats Flinders Island Mt Chappell Island Cape Barren Island Chappell Island tiger snake (Notechis ater serventyi) Short-tailed shearwater (Puffinus tenuirostris) Gila monster (Heloderma suspectum) Western banded gecko (Coleonyx variegatus) Endothermy versus ectothermy Advantages of ectothermy: • Low energy requirements Low food habitats Fluctuating food habitats Small body dimensions surface/volume 1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0 0 2 4 6 body length 8 10 mammals: >20 g lizards: 8% spp. < 1 g 80% spp. < 20 g salamanders: 20% spp. < 1 g 90% spp. < 20 g Dwarf chameleon Dwarf gecko Monte Iberia Eleuth Kitti’s hog-nosed bat Etruscan shrew L: 29-33 mm W: 1.5-2.5 g FR: 4xW/day HR: 835 b/min RR: 661 p/min Endothermy versus ectothermy Advantages of ectothermy: • Low energy requirements Low food habitats Fluctuating food habitats Small body dimensions Elongate body forms surface/volume 10 diameter 5 height 0 0 10 20 30 height/diameter 40 50 wood rat (Neotoma sp.) weasel (Mustela nivalis) energy loss: x2 Afrocaecilia taitana Desmognathus ochrophaeus Bipes bipes Anguis fragilis Opheodrys aestivus Endothermy versus ectothermy Advantages of ectothermy: • Low energy requirements Low food habitats Fluctuating food habitats Small body dimensions Elongate body forms Low water habitats Sauromalus obesus Scaphiopus couchii Endothermy versus ectothermy Advantages of ectothermy: • Low energy requirements Low food habitats Fluctuating food habitats Small body dimensions Elongate body forms Low water habitats Low oxygen habitats Iguana iguana Amblyrhynchus cristatus Neoseps reynoldsi Scincus mitranus Dilong paradoxus Xu et al. 2004. Nature 431: 680-684. Synapsida (mammal-like reptiles) Dimetrodon (Pelycosauria) Moschops (Therapsida) Endothermy in Mammalia: 1. RM x5 2. Tb > Ta, 28°C < Tb < 40°C 3. DTcore < 1-2°C 4. Maero x5 • Thermoregulation first physiological version Synapsida evolve from small ectotherms decrease in size increase in size (30-100 kg) Tb constant, physiological benefits become inertial homeotherms evolve insulation increased metabolism improved insulation McNab 1978. Am. Nat. 112: 1-21. • Thermoregulation first brain version Synapsida evolve from small ectotherms evolve larger, more complex brains increase in size (30-100 kg) Tb constant, physiological benefits Hulbert 1980. become inertial homeotherms evolve insulation • Thermoregulation first ecological version Synapsida evolve from small ectotherms evolve nocturnal habits increase in size (30-100 kg) Tb constant, physiological advantages become inertial homeotherms evolve insulation Crompton et al. 1978. Nature 272: 333-336. • Aerobic capacity first sustained ativity version small change in basal metabolic rate minimal effect on thermoregulatory capacity large effect on maximal aerobic metabolic rate Ruben 1995 Ann. Rev. Physiol. 57: 69-95. • Aerobic capacity first parental care version small change in basal metabolic rate minieme verandering in thermoregulatie-capaciteit large effect on maximal aerobic metabolic rate necessary for locomotor costs related to parental care Koteja 2000 Proc. R. Soc. Lond. 267: 479-484 Evolutionary physiology topics 1. Patterns • How and why of particular transitions • Testing a-priori-hypotheses plastic responses are adaptive Dicerandra linearifolia • leaf length • leaf thickness • density of stomata Winn A.A. 1999. J. Evol. Biol. 12: 306-313. 0.165 35 96 94 25 20 15 92 Density of stomata (mm-2) 0.160 Leaf thickness (mm) Leaf length (mm) 30 0.155 0.150 0.145 10 90 88 86 84 82 80 78 5 0.140 winter summer 76 winter summer winter summer Winn A.A. 1999. J. Evol. Biol. 12: 306-313. -0.01 0.60 0.55 0.50 0.45 0.40 0.35 0.30 Selection gradiënt for stomata density 0.08 Selection gradiënt for leave thickness Selection gradiënt for leave length 0.65 0.06 0.04 0.02 0.00 -0.02 -0.04 winter summer -0.02 -0.03 -0.04 -0.05 -0.06 winter summer winter summer Winn A.A. 1999. J. Evol. Biol. 12: 306-313. Beneficial acclimation hypothesis Beneficial acclimation hypothesis Colder is better Hotter is better Beneficial acclimation hypothesis Deleterious acclimation hypothesis Beneficial acclimation hypothesis Escherichia coli Leroi et al. 1994.Proc. Natl. Acad. Sci. USA 91: 1917-1921. Beneficial acclimation hypothesis > 32° 32° 41.5° 32° 41.5° Escherichia coli 37° acclimation > 41.5° competition Leroi et al. 1994.Proc. Natl. Acad. Sci. USA 91: 1917-1921. Beneficial acclimation hypothesis Bicyclus anynana Geister T.L. & Fischer 2007. Behav. Ecol. 18: 658-664. Beneficial acclimation hypothesis 20° 27° development larvae 20° 20,20° 27,27° 27° 20,27° 27,20° 27° 20,20° 27,27° 20° acclimation 20,27° 27,20° Beneficial acclimation hypothesis Oribatida Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Marion Island, Prince Edward Islands Beneficial acclimation hypothesis 15° Locomotor tests -5° up to 35° Halozetes marinus Halozetes marionensis 10° Halozetes belgicae Halozetes fulvus 5° Podacarus auberti 0° acclimation 7 days Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis Halozetes marinus deleterious acclimation Halozetes marionensis Halozetes belgicae Halozetes fulvus Podacarus auberti Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis 15°C 10°C 5°C 0°C Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis Halozetes marinus deleterious acclimation Halozetes marionensis beneficial acclimation Halozetes belgicae Halozetes fulvus Podacarus auberti Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis 15°C 10°C 5°C 0°C Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis Halozetes marinus deleterious acclimation Halozetes marionensis beneficial acclimation colder is better Halozetes belgicae Halozetes fulvus Podacarus auberti Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis 15°C 10°C 5°C 0°C Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis Halozetes marinus deleterious acclimation Halozetes marionensis beneficial acclimation colder is better Halozetes belgicae Halozetes fulvus Podacarus auberti geen plasticiteit geen plasticiteit Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Beneficial acclimation hypothesis 15°C 10°C 5°C 0°C Deere J.A. & Chown S.L. 2006. Am. Nat. 168: 630-644. Evolutionary physiology topics 1. Patterns • How and why of particular transitions • Testing a-priori-hypotheses plastic responses are adaptive phenotypic plasticity ~ environmental variability Rana temporaria Lind & Johansson 2006. J. Evol. Biol. 20: 1288-1297 • • • • • • 14 small islands 10 clutches < 20-50 eggs depth pools variability drying / island lab: 4 tadpoles / container 2 regimes: Constant & Drying • developmental time ~ regime (D<C) • developmental time ~ island • phenotypic plasticity ~ variability island Lind & Johansson 2006. J. Evol. Biol. 20: 1288-1297 constant developmental time island 1 (homo) 28 plasticity=11 17 drying island 2 (hetero) 28 plasticity=18 10 • devolopmental time ~ regime (D<C) • developmental time ~ island • phenotypic plasticity ~ variability island Lind & Johansson 2006. J. Evol. Biol. 20: 1288-1297 Evolutionary physiology topics 1. Patterns • How and why of particular transitions • Testing a-priori-hypotheses plastic responses are adaptive phenotypic plasticity ~ environmental variability a jack-of-all-trades is a master of none sprint speed sprint speed ‘specialist’ 40 35 30 25 20 ‘generalist’ 15 10 5 0 6 8 10 14 18 22 26 lichaamstemperatuur 30 34 38 rank 12 10 Laudakia stellio 8 6 4 2 0 18 22 26 30 34 38 42 lichaamstemperatuur Huey R.B. & Hertz P.E. 1984. Evolution 38:441-444. rank 5 4 3 Amoeba 2 1 0 10 15 20 25 lichaamstemperatuur 30 38 Huey R.B. & Hertz P.E. 1984. Evolution 38:441-444. Escherichia coli Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. Escherichia coli 7.8 7.0 6.3 5.3 2000 generations non-active Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. Escherichia coli 7.8 7.0 6.3 5.3 2000 generations non-active C > P in constant and fluctuating environments Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. Escherichia coli 7.8 7.0 6.3 5.3 2000 generations non-active R > P in some fluctuating and constant environments Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. Escherichia coli 7.8 7.0 6.3 5.3 2000 generations non-active B > P in fluctuating environments, but not in 7.8 Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. Escherichia coli 7.8 7.0 6.3 5.3 2000 generations non-active A > P in constant, not in fluctuating environments Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. Escherichia coli (1) adaptation to cycling pH, randomly changing pH and constante pH follows different patterns (2) in variable environments generalists evolve, in constant environments specialists evolve; (3) in variable environments the ‘cycling’ lines have a higher fitness than the ‘random changes’ lines; (4) an acclimation benefit (BAH) was not always detected. Hughes et al. 2007. Physiol. Biochem. Zool. 80: 406-421. • 18 Lygosominae • sprinting, jumping, clinging, climbing Goodman et al. 2007. Evol. Ecol. Res. 9: 527-546. Evolutionary physiology topics 1. Patterns • How and why of particular transitions • Testing a-priori-hypotheses plastic responses are adaptive phenotypic plasticity ~ environmental variability a jack-of-all-traits is a master of none symmorphosis: design satisfies need Evolutionary physiology topics Evolutionary physiology topics king pin one half rule Weibel et al. 1991. Proc.Natl. Acad. Sci. USA 88: 10357-10361 mitochondria in muscle cells capillary design (volume, surface) hematocrite V02max heart stroke volume surface pulmonary vesicles diffusion capacity membrane Evolutionary physiology topics 1. Patterns 2. Processes • natural selection performance gradient genetic variation ? fitness gradient design variation performance fitness ? variation ? variation quantitative genetics physiology morphology biochemistry kinematics biomechanics ecology behavioral biology juvenile survival Zootoca vivipara limited food supply abundant food supply initial endurance LeGalliard et al. 2004. Nature 432: 502-505. Evolutionary physiology topics 1. Patterns 2. Processes • natural selection • sexual selection • intrasexual selection (male-male combat) • intersexual selection (female choice) Neriene litigiosa deCarvalho et al. 2004. Anim. Behav. 68: 473-482. Phase 3 Locomotion Joint male energy use (EmW) 1200 X 11.5 1200 800 Phase 2 X 7.4 600 Phase 1 X 3.5 400 200 0 0 1 2 3 4 5 6 7 8 Time (min) Neriene litigiosa deCarvalho et al. 2004. Anim. Behav. 68: 473-482. Necora puber Thorpe et al. 1995. Anim. Behav. 50: 1657-1666 Uca lactea Matsuma & Murai 1995. Anim. Behav. 69: 569-577 anaerobic respiration Agkistrodon contortix Schuett & Grober 2000 Physiol & Behav 71: 335-341. Agkistrodon contortix Schuett & Grober 2000 Physiol & Behav 71: 335-341. Anolis sagrei Evolutionary physiology implications Evolutionary physiology implications Evolutionary physiology implications Evolutionary physiology implications http://www.sfecologie.org/blog/2011/09/30/evolrescueonline-topic-1/