Coping with Environmental Variation: Temperature and Water Photo of differential drought stress tolerance in 2 genotypes of Arabidopsis thaliana from http://www.riken.jp/en/research/rikenresearch/highlights/7320/ The Ecological Niche Key niche dimensions include: Energy availability Availability of other resources Physical conditions (e.g., pH) Figure from Bruno et al. (2003) Trends in Ecology & Evolution Generalists & Specialists Two species with the same niche breadths along both niche axes Water Availability (resource & physical condition) Species 1 Species 2 Temperature (C) Generalists & Specialists Species 1 has a narrower niche breadth along both niche axes Water Availability (resource & physical condition) Species 1 (Specialist) Species 2 (Generalist) Temperature (C) Generalists & Specialists Individuals with much narrower niche breadths than their respective populations Water Availability (resource & physical condition) Species 1 Specialist? Species 2 Generalist? Individual Temperature (C) Stress Tolerance Range of environmental tolerances helps determine potential geographic distribution (fundamental niche) Photo of saguaro in Sonoran Desert and range map from Wikimedia Commons Tolerance & Avoidance of Stress To reduce exposure to, or consequences of, environmental stress: physiological changes, e.g., drought deciduousness Photo of dry season Tabebuia aurea from Wikimedia Commons Tolerance & Avoidance of Stress To reduce exposure to, or consequences of, environmental stress: behavioral/physiological changes, e.g., dormancy (highly reduced metabolic activity), torpor (reduced met. & temp. in animals), hibernation (extended torpor); winter sleep / denning (extended sleep with slight reduction in temp.) Hibernating northern bat in Norway from Wikimedia Commons Tolerance & Avoidance of Stress To reduce exposure to, or consequences of, environmental stress: behavior, e.g., migration Migration routes for some champion avian migrants from Wikimedia Commons Acclimatization Usually reversible physiological, morphological or behavioral adjustment by individuals to reduce stress (under lab conditions = acclimation) Photo of Mt. Everest base camp from https://studentadventures.co.uk/adventures/everest_base_camp_trek Adaptation Ecotypes are populations adapted to local conditions Image re Clausen, Keck & Heisey’s (1948) classic study from http://www2.nau.edu/~gaud/bio300w/ecotype.htm Temperature Enzymes have physical optima Isozymes’ optima differ Denature at high temp. Lower activity limit -5 C Image from https://wikispaces.psu.edu/pages/viewpage.action?pageId=112526688&navigatingVersions=true Temperature Influences the properties of biological membranes Image of cell membrane from Wikimedia Commons Thermal Energy Balance (Plants) Energy input vs. energy output determines an object’s heat energy change (and internal temp.) Hplant = SR + IRin – IRout Hconv Hcond – Het Cain, Bowman & Hacker (2014), Fig. 4.8 Thermal Energy Balance Boundary layer lowers convective heat loss Cain, Bowman & Hacker (2014), Fig. 4.11 Thermal Energy Balance (Animals) Ectotherm – regulates temp. through energy exchange with environ. Hectotherm = SR + IRin – IRout Hconv Hcond Endotherm – relies primarily on internal heat generation Hendotherm = SR + IRin – IRout Hconv Hcond + Hmet Photos of basking snake and weasel from Wikimedia Commons Thermal Energy Balance (Animals) Cain, Bowman & Hacker (2014), Fig. 4.16 A Properties of Water Maximum density at 3.98 C High heat capacity – ratio of change in heat energy to change in temperature Photo of water in 3 physical phases from Wikimedia Commons Properties of Water Universal solvent for biologically important solutes “Water is the medium in which all biochemical reactions necessary for physiological function occur” Quote from Cain, Bowman & Hacker (2014), pg. 98; photo of bacteria grown from Lake Whillans (beneath Antarctic ice sheet) from http://www.nature.com/news/lakes-under-the-ice-antarctica-s-secret-garden-1.15729 Water and Biology Organismal water content for normal physiology 60% to 90% of body mass Salt balance is intimately tied to water balance Infra-red camera trap photo of bat at Peruvian mineral lick from http://news.mongabay.com/2008/0714-hance_bats_atbc.html Water Potential Flows along potential energy gradients (high to low) Gravitational potential – owing to gravity o = Osmotic potential – negative, owing to solutes p = Pressure (turgor) potential – positive, owing to pressure (negative if under tension) m = Matric potential – negative, owing to attractive forces on surfaces (e.g., large molecules, soil particles) Water potential = o + p + m Resistance – force that impedes movement of water (its reciprocal is conductance) Water Potential & Transpiration Daytime – decreasing water potential gradient from soil, through terrestrial plant, to atmosphere Cain, Bowman & Hacker (2014), Fig. 4.20 Allocation Tradeoff Root-shoot ratio Cain, Bowman & Hacker (2014), Fig. 4.22 Water & Salt Balance in Teleost Fishes seawater < teleost < freshwater Hypoosmotic Cain, Bowman & Hacker (2014), Fig. 4.24 Hyperosmotic Water & Salt Balance in Terrestrial Animals Exchange gases in dry environment (low water potential) Some adaptations to lower evaporative loss & water stress: High skin resistance Habitat selection (sufficient water to replace losses) Metabolic water High renal efficiency Image of camels in Chad from Wikimedia Commons Scaling Many scaling relationships can be expressed as power laws: Y = c Xs X is the independent variable – measured in units of a fundamental dimension; c is a constant of proportionality; and s is the exponent (or “power” of the function) The relationship is a straight line on a log-log plot: Log10(Y) = Log10(c) + s Log10(X) …and by rearranging, this is the form of the familiar equation for a straight line: y = mx + b Scaling Consider the scaling of squares & cubes as functions of the length of a side (the fundamental dimension) Area = Length2 Area Length2 Surface area = 6 * Length2 Surface area Length2 Volume = Length3 Volume Length3 Scaling 120 2 Y = X2 y = x 100 (accelerating function) Area 80 60 40 20 0 0 2 4 6 Length 8 10 12 Scaling 120 2 Y = X2 y = x 100 (accelerating function) Area 80 60 40 20 0 0 2 4 6 Length 8 10 12 Scaling 120 2 Y = X2 y = x 100 (accelerating function) Area 80 60 40 20 0 0 2 4 6 Length 8 10 12 Scaling 3.5 120 2 Y = X2 y = x (accelerating function) Area 80 Y = 2X 3 Log10(Area) 100 60 40 20 2.5 y = 2x 2 1.5 1 0.5 0 0 0 2 4 6 8 10 0 12 Length 0.2 0.4 0.6 0.8 Log10(Length) Etc… 1 1.2 Scaling 700 Log10(Surface Area) 3 2 Y = 6 * yX=26x Surface area 600 500 (accelerating function) 400 300 200 100 = 2x + 0.7782 Y = y2X + 0.778 2.5 2 1.5 1 0.5 0 0 0 2 4 6 8 10 0 12 Length 0.2 0.4 0.6 0.8 Log10(Length) Etc… 1 1.2 Scaling 3.5 1200 3 Y = X3 y = x (accelerating function) 800 600 400 2.5 2 1.5 1 200 0.5 0 0 0 2 4 Y = 3Xy = 3x 3 Log10(Volume) Volume 1000 6 8 10 0 12 Length 0.2 0.4 0.6 0.8 Log10(Length) Etc… 1 1.2 Scaling Consider the ways in which surface area & volume of a sphere scale with its radius Surface area = 4 r2 Surface area r2 Volume = 4/3 r3 Volume r3 Scaling Surface-to-volume ratio: Surface area r2 Volume r3 Surface area1/2 r Volume1/3 r Surface area1/2 Volume1/3 Surface area Volume2/3 Scaling Slope = 1 1400 Log10(Surface area) Y=4.83 * X 1200 Surface area 3.5 y = 4.8352x0.6667 0.667 1000 800 600 (decelerating function) 400 200 y = 0.6667x* +X0.6844 Y=0.667 + 0.68 3 2.5 2 1.5 1 0.5 0 0 0 1000 2000 3000 4000 0 5000 Volume 1 2 3 Log10(Volume) Etc… Volume increases proportionately faster than surface area 4 Scaling Slope = 1 1400 Log10(Surface area) Y=4.83 * X 1200 Surface area 3.5 y = 4.8352x0.6667 0.667 1000 800 600 (decelerating function) 400 200 y = 0.6667x* +X0.6844 Y=0.667 + 0.68 3 2.5 2 1.5 1 0.5 0 0 0 1000 2000 3000 4000 0 5000 Volume 1 2 3 Log10(Volume) Etc… This simple fact has myriad important implications for biology (e.g., heat exchange) 4 Scaling Assuming ecotothermy and environmental ceteris paribus, which one (dino or frog) warms up (or cools down) fastest? Paedophryne amauensis Image of dinosaurs and world’s smallest known vertebrate from Wikimedia Commons