Properties of Water. Processes of Water Movement in Plants Short Distance Transport. Long Distance Transport. Water & Plant Cells: Chapter 3
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1 Water & Plant Cells: Chapter 3 Why is water important? 1. 1g organic material requires 500g water. 2. Roles of water in plants (97% is lost) A. Solvent - Fluid medium for cytoplasm B. Photosynthesis C. Turgor pressure D. Transpiration & Heat dissipation E. Transport dissolved minerals 3. Productivity Properties of Water 1. Hydrogen bonding 2. Excellent solvent shells of hydration 3. Specific heat = 4.187kJ kg - 1 (1c/g) 4. Latent heat of vaporization = 44kJ mol Cohesion, adhesion & capillarity 6. Surface tension 7. Tensile strength > -30MPa 1 MPa = 1 newton m -2 1 MPa = 1 Joule m -3 1 MPa = 9.9 atmospheres 1 MPa psi 30 MPa = 4351 lb in -2 1 Atmosphere = 14.7 lbs in -2 1 Atmosphere = 760mm Hg 1 Atmosphere = MPa 1 Atmosphere = Bar 1 Atmosphere = X 10 5 Pa Major Processes Related to Water Movement in Plants 1. Short distance transport = Molecular Diffusion 2. Long distance transport = Bulk Flow Processes of Water Movement in Plants Short Distance Transport 1. Molecular Diffusion 1. Fick s first law Short Distance Transport Long Distance Transport Js Ds = rate of transport = diffusion coefficient = how easily something moves in a medium c s = difference in concentration between two points = concentration gradient x = distance between two points/concentrations 2. Importance: Identifies factors governing short distance transport: t: 1. Concentration gradient 2. Distance 3. Medium through which something moves 1
2 Processes of Water Movement in Plants Short Distance Transport Influence of distance 1. Average time for a particle to diffuse L = distance traveled D = diffusion coefficient 2. Example: Glucose travels 50um in 2.5 seconds 1 meter in 32 years 3. Point: Diffusion rapid over short distances very slow over long distances Processes of Water Movement in Plants Long Distance Transport Bulk flow or Mass Flow 1. Molecules move in bulk driven by a pressure gradient. Volume flow rate = meters sec -1 r = radius of tube η= viscosity of the liquid ψp = pressure gradient x = distance between gradient points 2. Pressure driven bulk flow moves water long distances Chemical Potential of Water Pressure Units 1. Free energy per mole of water 2. Water potential = 3. Units of chemical/water potential Energy: J mol -1 Pressure: Pascal 4. The Pascal: 1Pa = lb in -2 1MPa = 1,000,000 Pa = 145 lb in -2 = 10 bars = 9.87 atmospheres 2
3 Water Potential and Its Components 1. Water Potential Equation ψ = ψ s + ψ p + ψ g ψ = water potential in MPa ψ s = solute or osmotic potential ψ p = pressure potential ψ g = gravity potential Solute or Osmotic Potential 1. Represents effect of solutes on water potential 2. Solutes always decrease water potential always reduce free energy of water 3. van t Hoff equation ψ s = -RTc s ψ s = solute potential R = gas constant (8.32 J mol -1 K -1 ) T = temperature (degrees K) c s = solute concentration (osmolal = moles solute per liter water) 4. Independent of solute identity Values of Osmotic Potential Pressure Potential 1. Hydrostatic Pressure of solution 2. Turgor pressure = Positive hydrostatic pressure 3. Tension = Negative hydrostatic pressure 4. ψ p = 0 in a standard state Flaccid Turgid Tension 3
4 1. Gravity causes water to move downward 2. ψg = ρw g h ρ w = density of water (1.0 kg/m 3 at 4C) h = height (meters) g = acceleration due to gravity (9.8 ms - 2 ) ρw g = 0.01 M Pa m -1 Gravity Potential For a 100 meter tree: ψg = 0.01 MPa m - 1 X100 = 1.0 MPa 1MPa = = 145 lb in-2 5. Here s s the point: In cells: ψg is negligible (no change in height) In tall trees (100 meters) = 1.0 MPa change in ψ w 1. Matric = surface which binds water 2. Can be strong or negligible 1. Neglibible hydrated tissues 2. Strong Dry surfaces (e.g. seeds) Examples: ψ m = Matric Potential Dry seed ψ 50 to -350 Hydrated seed ψ 0 MPa 350 MPa 3. Here s s the point: In hydrated tissue: ψ m is negligible In dry tissue: ψ m is very important 4. Application Soil matric potential is a critical variable in crop yield, runoff, evapotranspiration and irrigation scheduling. Numerical Examples of Water Potential & Its Components Numerical Examples of Water Potential and Its Components 4
5 Numerical Examples of Water Potential and Its Components Hoffler diagram: Water Potential and Change in Cell Volume Major ideas As cell water potential decreases first 5% 1. In a turgid cell, Water potential drop results in sharp turgor drop with little change in cell volume. 2. As cell volume decreases below 90%, osmotic potential decreases faster than turgor (Turgor = 0) with further decline in water potential Sensitivity of Various Physiological Processes to Dehydration END Chapter 3 Water and Plant Cells 5
6 Physical Characteristics of Soil Relative Size of soil particles Chapter 4 Water Balance of Plants Mechanisms & driving forces operating on water transport in plants Field Capacity Wet Clay soil 40% Sand 3% Components of soil water potential Soil Hydraulic Conductivity Soil hydraulic conductivity how easily water moves through soil ψsoil = ψs + ψp ψp = about -2MPa in dry clay soil. Permanent Wilting Point = ψsoil > ψplant 6
7 Role of Water Evaporation in Water Movement Ψ P 2T = r T = surface tension of water = 7.28 X 10-8 MPa m r = radius of curvature Water Uptake by the Root Water Movement: Soil to Root Water Absorption: Root Root Hairs increase surface area. Epidermis Cortex Concave menisci form at soil-water interface.. creating very negative matric potentials. More water removed More acute menisci More negative ψsoil. Aquaporins regulate water movement through endodermis. 7
8 Role of Aquaporins in water transport Root Pressure and Guttation How does water move up a tree? 1. Diffusion 2. Capillarity 3. It s pushed up 4. It s pulled up Processes of Water Movement in Plants Diffusion is a very slow process 1. Average time for a particle to diffuse L = distance traveled D = diffusion coefficient 2. Example: Glucose travels 50um in 2.5 seconds 1 meter in 32 years 3. Point: Diffusion rapid over short distances very slow over long distances 8
9 Capillarity as a Mechanism of Long Distance Water Transport Could capillarity generate the force needed to draw water up a tree? To calculate the height liquid can be raised by capillarity: h = 2Tcosø dgr h = height of capillary (meters) T = surface tension (@20C, T for water = Nm-1) cos ø = lifting component of meniscus angle (approx = 1) d = density of liquid (998kgm-3) g = gravitational constant (9.80 ms-2) r = radius of capillary (in meters) Capillarity as a Mechanism of Long Distance Water Transport Capillary diam (µm) height of water column (m) Point: Height to which a column of water can be lifted by a capillary is inversely related to the diameter of the capillary. Xylem and Water Transport Circular Bordered Pits 9
10 Water Movement Through Xylem Requires less pressure than movement through cells From Pouiseuille s equation: Rate = 4mm s -1 Tube radius = 40um Tissue Xylem Layer of cells Pressure gradient (MPa m -1 ) X 10 8 Pressure Needed to Move Water Up a Tree Pressure gradient = 0.02 MPa m -1 Tree height = 100 meters (a Redwood tree) Total Pressure Needed = 2.0 MPa Weight of Water added Pressure to overcome 100 meters X 0.01MPa m -1 = 1MPa Total Pressure Needed = 3MPa Fig 4.16 Soil Soil-Plant-Atmosphere Continuum Soil-Plant Plant-Atmosphere Continuum 10
11 Relative Humidity and Absolute Water Concentrations cwv RH % = x100 cwv( sat ) Relative Humidity and Water Potential RT Ψ w = ln( RH ) Vw Ψ w = water potential (kpa) R = gas constant = kpa L mol -1 K -1 T = temperature ( o K) V w = partial molal volume of water = Liters / mol RH = relative humidity (0-1.0) Note: MPa = kpa/1000 RH = Relative Humidity (%) Cwv = water concentration (moles m -3 ) Cwv(sat) = saturation water concentration (moles m -3 ) Diurnal course of Relative Humidity over a deciduous forest Water Movement from Leaf To Atmosphere 1. Water Vapor diffuses from Leaf to Air = Transpiration 2. Diffusion influenced by 2 factors 1. Water Vapor Concentration Gradient 2. Diffusional Resistance 11
12 Stomatal aperture and Boundary Layer Resistance at leaf surface Role of Water Evaporation in Water Up a Tree Water Evaporation in a leaf generates Negative Pressure in the Xylem Fig Soil-Plant- Atmosphere Continuum Experimental Demonstration of Water Under Tension 1. Dye uptake in cut stems 2. Change in stem diameter (Fritts 1958 Beech Trees) 3. Direct measurement Pressure bomb 12
13 Measuring Water Potential in Plants with a Pressure Bomb Guard Cells Control Short Term Regulation of Water Loss 1. Turgor increase in guard cell solute concentration pores open 2.Turgor decrease in guard cell solute concentration pores close Physical Challenges of Cohesion Tracheary Elements are connected by Pits Physical Challenges of Cohesion-Tension 1. Break strength of water is 30MPa << Pressure needed to raise water. 2. Water in a metastable state -3MPa - 30 Atmospheres 3. Cavitation or embolism A. Controlled by Redissolving trapped gas Secondary growth replaces xylem 13
14 Physical Challenges of Cohesion-Tension Pit pairs regulate embolism Trachary element cavitation in trees infected with wilt diseases Transpirational Water Loss Plant Water transpired Potato 25 gallons Winter wheat 25 gallons Tomato 34 gallons Corn 54 gallons Species vary in xylem hydraulic conductivity drought tolerance Silver Maple Tree 58 gallons hr -1 per kg corn grain per kg corn plant 600 kg (about 158 gallons) 225 kg (about 60 gallons) 14
15 Water Transport in Mangroves Mangroves in Esmeraldas- Ecuador, Majagual Region. Florida Mangroves Water Transport in Mangroves END Water Balance of Plants 15
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