Figure 4.10 Water pathway through the leaf Figure 4.12 Electron micrographs of stomata; (B) Stomatal complexes of Carex Figure 4.13 Electron micrograph showing a pair of guard cells from the dicot tobacco 4.16 Radial alignment of the cellulose microfibrils in guard and epidermal cells 1 Figure 4.10 Water pathway through the leaf Water movement into and out of guard cells is controlled by ion transport processes which causes changes to cell s and p Pumping K+ into guard cells increases p - stomatal opening Pumping K+ out of guard cells decreases p - stomatal closing Ohm’s Law Analogy for Transpiration Transpiration – Water Loss from Leaves Current (amps) Transpiration is a flux of water vapour from a leaf Potential Difference (volts) Resistance (ohms) Diffusion of water vapour out of leaf is controlled by: (i) a driving force – which is the gradient in water vapour concentration (or partial pressure) between the leaf and the air (ii) resistance to water loss which is controlled primarily by stomatal opening Transpiration Rate = Driving Force / Resistance Conductance = 1/Resistance Transpiration Rate = Driving Force x Conductance Abbreviations used in Transpiration and Conductance Calculations Symbol Definition Usual Units E gw P ei ea mmol m-2 s-1 mmol m-2 s-1 kPa kPa kPa Transpiration rate Conductance to water vapour Atmospheric pressure Intercellular water vapour partial pressure Ambient water vapour partial pressure Ohm’s Law Analogy for Transpiration E gw Further subscripts can be applied to the symbol for conductance (gw) to indicate which section of the diffusion pathway is being referenced. For example, if you are referring to conductance over the total pathway from the intercellular air spaces to the atmosphere, the total conductance term is indicated as: gw-total If you are referring only to stomatal conductance: gw-stomata If you are referring only to boundary layer (BL) conductance: gw-BL (ei ea ) P Transpiration (E) = conductance (gw) x driving force (VPD) 2 Atmosphere Epidermis Internal Air Space Leaf Cells Xylem Liquid Water ea ei Humidity Concepts 1. 2. 3. 4. Saturation Vapour Pressure Relative Humidity Actual or Absolute Humidity Vapour Pressure Difference (VPD) Water Vapor Water Vapor Relative Humidity << 100 % Liquid Water Relative Humidity 100 % 4.12 Concentration of water vapor in saturated air as a function of air temperature Humidity Concepts 1. Saturation Vapour Pressure The absolute amount of water vapour the air can hold when saturated varies as an exponential function of air temperature Saturation Vapour Pressure (kPa) = 0.611 e ((17.502 T)/(T + 240.97)) Saturation Vapour Pressure (kPa) where e is the base of the natural logarithm and T is air temperature in °C Saturation Vapour Pressure Varies with Temperature 14 12 10 8 6 4 2 0 0 10 20 30 40 Air Temperature (°C) 50 Humidity Concepts Humidity Concepts 2. Relative Humidity 3. Absolute Humidity The ratio of the absolute amount of water vapour in the air divided by how much the air could hold if it was saturated Relative Humidity (RH) = Actual or Absolute Humidity (kPa) Saturation Vapour Pressure (kPa) RH may be expressed as a proportion (as done here) or as a percentage The absolute amount of water vapour in the air It is calculated from measurements of air temperature and air RH Air temperature is used to determine the saturation vapour pressure Absolute Humidity (kPa) = Saturation Vapour Pressure (kPa) x RH 3 Transpiration – Water Loss from Leaves Humidity Concepts 4. Vapour Pressure Difference (VPD) The difference in absolute vapour pressure between the leaf intercellular air spaces (ei) and the well-mixed air outside of a leaf (ea) VPD (kPa) = ei (kPa) – ea (kPa) • Solar radiation energy input causes the water-to-vapour phase change • Driving force is a vapor pressure gradient (VPD) – Depends on air temperature and the water vapor content in the bulk well-mixed air and also on leaf temperature • Stomata are usually the major resistance – Stomatal conductance varies and is dependent on: • • • • Soil moisture Vapor pressure of air Light intensity CO2 concentration of air Stomatal Conductance (g) Conductance (gw) is calculated by re-arranging the Ohm’s Law analogy equation E gw gw (ei ea ) P EP (ei ea ) Humidity Response Light Response PPFD = Photosynthetically active Photon Flux Density Soil Moisture Response • Boundary layers can also be a resistance to water loss from leaves A boundary layer develops as air flows across a leaf Boundary layer conductance depends on • Leaf size and leaf shape • Wind speed This boundary layer retards the transfer of CO2 and H2O from the leaf to the surrounding air 4 Figure 4.11 Dependence of transpiration flux on the stomatal aperture of zebra plant Plant Response to Water Stress • water shortage - drought • water excess – flooding Textbook Reading: Chapters 3 & 4 All Chapter 26: pp. 756-760 pp. 765-772 Plant Response to Water Stress: Drought MPa -0.03 -0.2 -0.81 -1.60 Soil Water Potential Figure 26.4 Effects of water stress on photosynthesis and leaf expansion of sunflower Figure 26.1 Dependence of leaf expansion on leaf turgor 5 Osmotic Adjustment Figure 26.10 Solute adjustments during osmotic stress • it is possible to decrease w by decreasing s - this allows high p which is required for growth • this can be actively done in living cells by accumulating compatible solutes in the cytoplasm • compatible solutes: sugars, sugar alcohols, proline w=-0.9 MPa • ions can be compartmented into the vacuole to prevent the charged molecules from interfering with enzyme activity in the cell cytoplasm Pressure potential (p) Figure 26.11 Four groups of molecules frequently serve as compatible solutes Drought Induced Reduction in Stomatal Conductance: Mechanisms and Signals Initial ideas about water stress-induced reduction in stomatal conductance assumed that reductions in leaf water potential and pressure potential (p or turgor) were the mechanism that controlled changes in guard cell turgor and reduced stomatal opening. This is what occurs in primitive (non-seed plants). However, in more advanced seed plants (conifers and angiosperms) stomatal closure is part of a complex signaling system involving the plant hormone, Abscisic Acid (ABA). 6 7
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