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LECTURE 4 CH 2+3 PHYSICAL ENVIRONMENT: CONSTRAINTS AND SOLUTIONS Background Physical laws set limits on life. Life exists out of equilibrium with physical world; uses energy to counteract. Life tolerates extreme conditions. Adaptations evolved in response to physical environment. Shelford’s ‘Law of Tolerance’ Individual’s physical responses show optimum and limits of tolerance. Tolerance limits affect individual’s performance and survival/reproduction. Distribution of species controlled by tolerance limits. More than 1 factor may limit distribution. Tolerance is related to functioning of enzymes. Physical factors organisms face 1) Water Why is it the ideal medium for life? How do constraints of water account for adaptation of form? What are thermal properties of water? Buoyancy and viscosity Limits of water determine distribution of deserts (coupled with high T in hot deserts) Adaptations of animals to aridity (e.g. kangaroo rat) Vary body temperature Concentrated urine and dry feces Nocturnal activity; use burrows in day Food provides metabolic water Adaptations of plants to aridity How do plants obtain water? Soil – plant – atmosphere continuum Soil - roots via osmosis Movement upward in plant aided by cohesion and adhesion Transpiration pull from atmosphere to leaf to xylem water column Water gradient (in to out) is very steep CO2 gradient (out to in) is very shallow >>>>Water loss relative to CO2 gain Structural adaptations: hairs, sunken stomates Photosynthetic adaptations C3: Problem = Rubisco has higher affinity for O2 than CO2; Especially at high temp and low H2O; less CO2 fixed C4: Solution = Spatial separation of CO2 uptake from Rubisco; keep O2 low CAM: Problem = High water loss with stomates open in day to take in CO2 Solution = Temporal separation of CO2 uptake (night; high humidity) and CO2 fixation by Rubisco (day; low humidity); save water When and where did C3 vs. C4 evolved relative to CO2 conc.? 2) Dissolved solutes and nutrients Water as powerful solvent; contains solutes Solutes reflect bedrock chemistry Plants obtain mineral nutrients from soil water Root:shoot ratio > in low nutrients Ionic conc. in soils affects plant distributions Evolution of tolerance to heavy metals 3) Dissolved solutes: Salt and water balance High or low conc. of ions in water osmotic strain Hyperosmotic: gain water, increase pressure in cell; lose solutes Hypoosmotic: lose water, cells collapse; gain solutes Fish in fresh water: retain water and excrete ions Fish in salt water: retain ions and excrete excess water Animals excrete excess nitrogen in urine Plants in salt (e.g. mangroves) actively excrete salt.