Water movement through the soil-plant continuum · Water movement through the soil-plant continuum...
Transcript of Water movement through the soil-plant continuum · Water movement through the soil-plant continuum...
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CO2!
Plant Ecology in a Changing World Jim Ehleringer, University of Utah
http://plantecology.net
Water movement through the soil-plant continuum
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Part 1 Critical features essential
to plant invasion of land
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Land plants evolved from aquatic algae!
The transition to land created numerous benefits and challenges!
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Plants evolved a cuticle and stomata to regulate gas exchange,!which prevented leaves from drying out!
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Part 2 Water moves along gradients of decreasing water potential
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Plants tend to lose water when taking in CO2 !
~1 : 400!
http://biobook.nerinxhs.org/bb/special_topics/microscopy/plantanatomy/syringaleafhplabeled.htm!
Ohm’s Law analogy!!A = gCO2(ca – ci)!!E = gH2O(ei – ea)!!gCO2 = gH2O/1.6!
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Water moves through soils and plants:!• as a liquid!• by bulk flow !• along gradients of decreasing water potential!
Taiz & Zeiger 2010!
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Water along gradients of deceasing total water potential!
pure, free water at ground level and atmospheric pressure !
Plants generally have negative Ψ values.!The unit is MPa (or historically bars) !
Ψtotal = Ψpressure + Ψosmotic + Ψmatric + Ψgravitational!
Ψtotal = 0 !
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cell wall!
cell membrane!(semi-permeable)!
Ψtotal = Ψpressure + Ψosmotic + Ψmatric + Ψgravitational!
ΨT = -1! ΨT = 0!
non-equilibrium!starting condition!
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Ψtotal = Ψpressure + Ψosmotic + Ψmatric + Ψgravitational!
ΨT = 0! ΨT = 0!
equilibrium!condition!
ΨP = +1!Ψo = -1!
water diffuses inward!
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higher!
lower!
Ψtotal = Ψpressure + Ψosmotic + Ψmatric + Ψgravitational!
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higher!
lower!
(1 MPa = 145 psi)!
density of water!
gravity!
height!
Ψtotal = Ψpressure + Ψosmotic + Ψmatric + Ψgravitational!
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Matric potential is capillary forces (most relevant for water in soil)!
narrower capillaries = stronger forces!
adhesion + cohesion!
Ψtotal = Ψpressure + Ψosmotic + Ψmatric + Ψgravitational!
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Matric potential is capillary forces!
narrower capillaries = stronger forces!
matric! osmotic! gravitational!total!
surface tension!
contact angle!
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Water is held in capillaries between soil particles !
Nobel 2009!
smaller particles house smaller capillaries!
water evaporates from larger capillaries first, leaving more tightly held water in smaller capillaries!
water potential decreases!
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Soil type influences matric potential !
-2 -1.5 -1.0 -0.5 0!
(MPa)!
Campbell & Norman 1998!
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Part 3 Water moves from soil to
leaves under tension (negative pressure)
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How do plants pull water from the soil?!
http://biobook.nerinxhs.org/bb/special_topics/microscopy/plantanatomy/syringaleafhplabeled.htm!
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How do plants pull water from the soil?!
http://biobook.nerinxhs.org/bb/special_topics/microscopy/plantanatomy/syringaleafhplabeled.htm!
cell wall!
cellulose !microfibril!
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How do plants pull water from the soil?!
10 nm!
http://biobook.nerinxhs.org/bb/special_topics/microscopy/plantanatomy/syringaleafhplabeled.htm!
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How do plants pull water from the soil?!
10 nm!
-29 MPa!
Water transport is driven solely by evaporation from cell walls!
http://biobook.nerinxhs.org/bb/special_topics/microscopy/plantanatomy/syringaleafhplabeled.htm!
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A range of minimum total water potentials in chaparral species!
Jacobsen et al 2007 Ecol Mono!
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A range of minimum water potentials in chaparral species!
Jacobsen et al 2007 Ecol Mono!
metastable! stable!
Xylem water is commonly!beyond its boiling point!
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Part 4 Plants have evolved features
to reduce cavitation
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Conduits cavitate when air enters from embolized conduits!
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Two inter-conduit check valve types!
homogenous torus-margo
Pittermann et al 2006 Science!
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Vulnerability curves quantify resistance to xylem cavitation!
Sperry 2000 AgForMet!
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Nobel 2009!
Measuring with a pressure chamber!
How do we typically measure total water potential?!
Water is under tension (negative pressure). In a pressure chamber,!positive pressure is added until water appears at the cut surface.!
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At equilibriun and!without transpiration:!
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without transpiration:!
with transpiration:!
sapflow!
hydraulic!conductance!
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without transpiration:!
with transpiration:!
sapflow!
hydraulic!conductance!
e! e!
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Gas exchange with the environment and internal water transport are coupled!
e! e!
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Hydraulic conductance depends on the path of water flow!
Taiz & Zeiger 2010!
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When plants lack a conducting system, plants are small and water movement!from source to the environment through plants is slow.!
http://commons.wikimedia.org/wiki/File:Sphagnum_-_moss_-_Moos_01.jpg!
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The evolution of xylem greatly increased hydraulic conductance (k)!
conduit diameter!conduit
number!
viscosity! length!
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An increased hydraulic conductance capacity and higher water transport rates support increased growth and larger plants.!
Smith & Sperry 2014 PCE!
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Part 5 Stomates control water loss
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Plants can control water loss by changing stomatal aperture!
Franks & Farquhar 2010 Plant Phys!
increase!turgor!
e! e!
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Changing stomatal aperture is an active process!
ATP!
ADP!
K+!
H+!
H+!
K+!
H2O!
H2O!
stimulus!
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Franks & Farquhar 2010 Plant Phys!
-closed-!
-open-!
K+ transfer between guard (G) and subsidiary (S) cells!allow for increased stomatal movement!
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Changing stomatal aperture is an active process!
abscisic acid!(ABA) !
ATP!
ADP!
K+!
H+!
H+!
K+!
H2O!
H2O!
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Franks & Farquhar 2010 Plant Phys!
Huperzia prolifera!(lycopod)!
Tradescantia virginiana (angiosperm)!
Subsidiary cells (S) allow guard cells (G) to overcome the mechanical advantage of the epidermis (E)!
-closed-!
-open-!
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Buckley, Mott, Farquhar 2003 PCE!
Stomata respond to a variety of environmental conditions!
Atmospheric humidity!
Sunlight!Atmospheric!
carbon dioxide!
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Not all graphics and images are original. We acknowledge materials from these sites:!http://www.goldiesroom.org/Note%20Packets/05%20Nutrition/04%20Nutrition--Lesson%204.htm!http://www.slideshare.net/eziennker/seed-plants-characteristics!https://c1.staticflickr.com/9/8007/7131296595_1c276b0261_b.jpg!http://prometheuswiki.publish.csiro.au/tiki-index.php?page=Making+leaf+surface+imprints!http://www.microscopy-uk.org.uk/micropolitan/botany/schlumbergera_stomata.jpg!!!!