Water uptake, water transport and transpiration

1. Water uptake, water transport and transpiration

2. Topics 1. General features of water flow through plants 2. Adhesion and cohesion in the water column 3. Water potential and how water moves through the plant 4. The energy budget of foliage 5. Measuring water potential 6. Stomatal control of leaf water potential 7. Consequences to the plant of changes in water potential

3. 1. General features 1. General features Functions: cooling CO2/O2 exchange nutrient flow translocation physiological processes + Water flows through the plant with rate largely determined by physics Plant structure and control systems minimize loss and ‘shortage’ in tissues Translocation is not unidirectional

4. 2. Cohesion and adhesion in the transpiration stream Fig. 32.3

5. Hydrogen Bonds and Cohesion H H O H O H H H O O H H H O H Water molecules have a weak negative charge at the oxygen atom and weak positive charge at the hydrogen atoms. The positive and negative regions are attracted to the oppositely-charged regions of nearby molecules. The force of attraction, dotted line, is called a hydrogen bond. Each water molecule is hydrogen bonded to four others. The hydrogen bond has ~ 5% of the strength of a covalent bond. However, when many hydrogen bonds form, the resulting union can be sufficiently strong as to be quite stable. Adhesion is the tendency of molecules of different kinds to stick together. Water sticks to the cellulose molecules in the walls of the xylem, counteracting the force of gravity. http://www.ultranet.com/~jkimball/BiologyPages/H/HydrogenBonds.html

6. Descriptive movie clip

7. 3. Water potential and how water moves through the plant At 22oC (72F) and 50% Relative Humidity yair = 100 MPa negative Water potential indicates how strongly water is held in a substance. It is measured by the amount of energy required to force water out of it. Think of squeezing a sponge or cloth. Water potential , referred to as y (psi), is measured in megapascals, Mpa, (SI, SystÈme Internationale) units. For pure water at standard temperature and pressure (STP) y = 0 Mpa. Typically yleaf=-1 to - 4MPa Water potentials of connected tissues defines rate of water flows through a plant. ysoil= 0.01 to - 0.1 MPa

8. 4. The energy budget of foliage Some radiation is reflected and some energy is re-radiated Radiation input In addition to radiation input leaf temperature can also be affected by wind speed and humidity because these conditions affect rate of cooling If Tleaf > Tair then the leafwarms the air Evaporative cooling depends upon latent heat of evaporation Only 1-3% of radiation is used in photosynthesis

9. 3.0 Wind speed influences transpiration 2.5 2.0 1.5 Transpiration flux, g H2O/cm2 leaf surface/second X10-7 1.0 0.5 Stomatal aperture, m The boundary layer around a leaf is thick in still air, and constitutes a major resistance to the fluxof H2O from the leaf. A slight increase in wind speed will reduce the boundary layer, and increase transpiration. Further increase in wind speed may reduce transpiration, especially for sunlit leaves, because wind speed will cool the leaf directly  http://forest.wisc.edu/forestry415/lecture6/windspd.htm

10. Laboratory measurement of transpiration A laboratory potometer 1. Fill the potometer by submerging it – make sure there are no air bubbles in the system. 2. Recut the branch stem under water and, keeping the cut end and the potometer under water, put the cut end into the plastic tubing.

11. 5. Measuring water potential The pressure bomb! Compressed air

12. Field measurements of  Forest laboratory in south west Scotland Measurement every hour for 7 days

13. Diurnal pattern of shoot water potential Midnight Midday 500 400 Transpiration 300 Mg/sec/tree 200 100 Shoot water 0  potential -1 MPa -2 30 Jul 31 Jul 1 Aug 2 Aug 3 Aug 4 Aug 5 Aug 6 Aug During daylight water loss from foliage exceeds water gain from soil so shoot water potential decreases.On sunny days  reaches –2 Mpa

14. Stomatal control clip

15. 6. Stomatal control of leaf water potential

16. … about osmosis?

17. Review of osmosis Diffusion of water across a selectively permeable membrane from a hypotonic to a hypertonic solution Hyper - above Hypo - below Water crosses the membrane until the solute concentrations are equal on both sides

18. Control of stomatal opening and closing Guard cells actively take up K causing water to enter by osmosis. The guard cell’s walls are unevenly thickened causing the cells to bow as they becomes turgid Fig. 32.4

19. Factors that can affect stomatal aperture How could you analyze them experimentally? Low leaf water potential High temperature Low CO2 concentration in the air spaces of the leaf causes a plant to open its stomata Circadian rhythm: a cycle of opening during daylight and closing during the dark.

20. Leaf hairs increase boundary layer resistance Trichomes or hairs cells grow out of the surface of  the epidermis. These may be uni-or multicellular depending on species. Both uni-and multicellular hairs may be branched. Some leaves have glandular hairs with an enlarged cell or group of cells at the end of  a stalk. Curatella americana from Cerrado (Brazil). Leaf surface showing stomata. Note the silicified hairs in the shape of a star

21. Consequences of changes in water potential Cavitation and tree ecology: The water column in wood can break when the tension on it exceeds the forces of adhesion and cohesion Contraction of tissues: At low  tissues that have not developed strong thickening can contract Cessation of physiological processes: Different physiological processes may cease at different 

22. Cavitation and tree ecology: Under very low water potentials the water column in xylem can break and cells become filled with air. This is called cavitation Air is in solution in water, but as water potential decreases it may come out of solution unless the forces of cohesion and adhesion are strong enough to overcome that. Cavitation becaused by a sustained period of dry conditions, i.e., a summer drought. It can also occur when xylem water freezes. Air has very low solubility in ice, so bubbles form; when the water thaws, the bubbles will coalesce and cause cavitation. Conifers have smaller conducting cells than angiosperm trees. Conifers only have tracheids while angiosperm trees have vessels. So, conifers may grow less quickly, but they may also cavitate less and this may help them survive in more stressed environments

23. Contraction of tissues: Measuring the extension rate of a conifer shoot

24. Shoot extension Midnight Shoot extension 4 Mm/h 2 0 -2 8 Total radiation MJm2 6 4 2 24 June 25 June 26 June 27 June

25. Cessation of physiological processes: Cell growth and wall synthesis are very sensitive and may stop at -0.5 MPa Photosynthesis, respiration and sugar accumulation are less sensitive. They may be affected between -1 and -2 MPa

26. Sections you need to have read 32.1 32.2 32.3 32.4 Courses that deal with this topic Botany 371/372 Plant physiology laboratory