WATER RELATIONS

of the
WHOLE PLANT

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Transport in Vascular Plants

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 Physical forces drive the transport of materials in
plants over a range of distances.

• Transport in vascular plants occurs on three

scales:
– Transport of water and solutes by individual
cells, such as root hairs.
– Short-distance transport of substances from
cell to cell at the levels of tissues and organs.
– Long-distance transport within xylem and
phloem at the level of the whole plant.

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 CO2 O2

Light
H2O Sugar

O2
H2O
CO2
Minerals
 Solute Transport

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 Selective Permeability of Membranes: A Review

• The selective permeability of the plasma

membrane controls movement of solutes into and
out of the cell.
• Specific transport proteins enable plant cells to
maintain an internal environment different from
their surroundings.

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 The Central Role of Proton Pumps
in Solute Transport
• Proton pumps in plant cells create a hydrogen ion
gradient that is a form of potential energy that can
be harnessed to do work.
• They contribute to a voltage known as a
membrane potential.

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CYTOPLASM EXTRACELLULAR FLUID

ATP
Proton pump
generates mem-
brane potential
and gradient.
 • Plant cells use energy stored in the proton
gradient and membrane potential to drive the
transport of many different solutes.

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CYTOPLASM EXTRACELLULAR FLUID
Cations ( , for
example) are
driven into the cell
by the membrane
potential.

Transport protein
Membrane potential and cation uptake
 • In the mechanism called cotransport, a transport
protein couples the passage of one solute to the
passage of another.

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 Cell accumulates
anions ( ,
for example) by
coupling their
transport to; the
inward diffusion
of through a
cotransporter.

Cotransport of anions
 • The “coattail” effect of cotransport is also
responsible for the uptake of the sugar sucrose by
plant cells

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 Plant cells can
also accumulate
a neutral solute,
such as sucrose
( ), by
cotransporting
down the
steep proton
gradient.

Cotransport of a neutral solute

 Water Transport

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 The Central Role of Water Potential
in Water Movement

• To survive, plants must balance water uptake and

loss.
• Osmosis determines the net uptake or water loss
by a cell.
-- affected by solute concentration and pressure.
-- regulated by the semi-permeable membrane.
-- governed by water potential gradient.

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• Water potential is a measurement that combines
the effects of solute concentration and pressure.
• Water potential determines the direction of
movement of water.
• Water flows from regions of higher water potential
(i.e., less negative) to regions of lower water
potential (i.e., more negative).

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 How Solutes and Pressure Affect Water Potential

• Both pressure and solute concentration affect

water potential.
• The solute potential of a solution is proportional to
the number of dissolved molecules.
• Pressure potential is the physical pressure on a
solution.

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 Quantitative Analysis of Water Potential

• The addition of solutes reduces water potential.

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 Addition of
solutes
0.1 M
solution

Pure
water
H2O

P = 0
S = –0.23
 = 0 MPa  = –0.23 MPa
 • Physical pressure increases water potential.

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 Applying
physical
pressure

H2O
H2O
P = 0.23 P = 0.30
S = –0.23 S = –0.23
 = 0 MPa  = 0 MPa
 = 0 MPa  = 0.07 MPa
 • Negative pressure decreases water potential.

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 Negative
pressure

H2O

P = –0.30 P = 0
S = 0 S = –0.23
 = –0.30 MPa  = –0.23 MPa
 • Water potential affects uptake and loss of water by
plant cells.
• If a flaccid cell is placed in an environment with a
higher solute concentration, the cell will lose water
and become plasmolyzed.
• (The protoplasts of flaccid cells are in contact with
their walls but lack turgor pressure.)

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 Water Relations in Plant Cells

Initial flaccid cell:

P = 0
 S = –0.7
0.4 M sucrose solution:  P = –0.7 MPa Distilled water:
P = 0 P = 0
Plasmolyzed cell S = –0.9 S = 0 Turgid cell
at osmotic P = –0.9 MPa  P = 0 MPa at osmotic
equilibrium equilibrium
with its surroundings with its surroundings

P = 0 P = 0.7
S = –0.9  S = –0.7
P = –0.9 MPa  P = 0 MPa

 > environmental 
initial conditions: cellular  initial conditions: cellular  < environmental 
 • If the same flaccid cell is placed in a solution with
a lower solute concentration, the cell will gain
water and become turgid.

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• Turgor loss in plants causes wilting, which can be
reversed when the plant is watered.

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 Aquaporin Proteins and Water Transport

• Aquaporins are transport proteins in the cell

membrane that allow the passage of water.
• Aquaporins do not affect water potential, but
rather the rate at which water diffuses down its
water potential gradient.
- regulated by phosphorylation of these proteins induced
by changes in second messengers (e.g., Ca 2+).

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 Three Major Compartments of Vacuolated Plant
Cells

• Transport is also regulated by the compartmental

structure of plant cells.
• The plasma membrane directly controls the traffic
of molecules into and out of the protoplast
• The plasma membrane is a barrier between two
major compartments, the cell wall and the
cytosol.

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 • The third major compartment in most mature plant
cells is the vacuole, a large organelle that
occupies as much as 90% or more of the
protoplast’s volume.
• The vacuolar membrane regulates transport
between the cytosol and the vacuole.

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 • In most plant tissues, the cell walls and cytosol
are continuous from cell to cell.
• The cytoplasmic continuum is called the
symplast.
• The apoplast is the continuum of cell walls and
extracellular spaces.

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 Cell wall Key
Cytosol
Symplast
Vacuole
Apoplast

Plasmodesma Vacuolar membrane

(tonoplast)
Plasma membrane
Cell compartments
 Functions of the Symplast and Apoplast in Transport

• Water and minerals can travel through a plant by

three routes:
– Transmembrane route: out of one cell, across
a cell wall, and into another cell.
– Symplastic route: via the continuum of
cytosol
– Apoplastic route: via the the cell walls and
extracellular spaces

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 Key

Symplast
Apoplast
Transmembrane route
Apoplast

Symplast

Symplastic route
Apoplastic route
Transport routes between cells
 Bulk Flow in Long-Distance Transport

• In bulk flow, movement of fluid in the xylem and

phloem is driven by pressure differences at
opposite ends of the xylem vessels and sieve
tubes.

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 Absorption of Water
by
Root Systems

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 Roots absorb water and minerals from the soil

• Water and mineral salts from the soil enter the

plant through the epidermis of roots and ultimately
flow to the shoot system.

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 Casparian strip

Endodermal cell
Pathway along
apoplast

Pathway
through
symplast

Casparian strip

Plasma
membrane
Apoplastic
route

Vessels
(xylem)
Symplastic Root
route hair

Epidermis Endodermis Vascular cylinder

Cortex
 The Roles of Root Hairs, Mycorrhizae, and
Cortical Cells

• Much of the absorption of water and minerals

occurs near root tips, where the epidermis is
permeable to water and root hairs are located.
• Root hairs account for much of the surface area of
roots.

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 • Most plants form mutually beneficial relationships
with fungi, which facilitate absorption of water and
minerals from the soil.
• Roots and fungi form mycorrhizae, symbiotic
structures consisting of plant roots united with
fungal hyphae.

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 2.5 mm
Mycorrhizae
 • After soil solution enters the roots, the extensive
surface area of cortical cell membranes
enhances uptake of water and selected minerals.

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 The Endodermis: A Selective Sentry

• The endodermis is the innermost layer of cells in

the root cortex.
• It surrounds the vascular cylinder and is the last
checkpoint for selective passage of minerals from
the cortex into the vascular tissue.

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 • Water can cross the cortex via the symplast or
apoplast.
• The waxy Casparian strip of the endodermal wall
blocks apoplastic transfer of minerals from the
cortex to the vascular cylinder

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 WATER TRANSLOCATION:
Transpiration and Ascent
of the Xylem Sap

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 Ascent of Xylem Sap

• Water and minerals ascend from roots to shoots

through the xylem.
• Plants lose an enormous amount of water through
transpiration, the loss of water vapor from leaves
and other aerial parts of the plant.
• The transpired water must be replaced by water
transported up from the roots.

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 Bulk Flow in Long-Distance Transport

• movement of fluid in the xylem and phloem is driven

by pressure differences at opposite ends of the xylem
vessels and sieve tubes.

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 Factors Affecting the Ascent of Xylem Sap

• Xylem sap rises to heights of more than 100 m in

the tallest plants.
• Is the sap pushed upward from the roots, or is it
pulled upward by the leaves?

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 Pushing Xylem Sap: Root Pressure

• At night, when transpiration is very low, root cells

continue pumping mineral ions into the xylem of
the vascular cylinder, lowering the water potential.
• Water flows in from the root cortex, generating
root pressure.

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 Guttation

• Root pressure sometimes results in guttation, the

exudation of water droplets on tips of grass blades or
the leaf margins of some small, herbaceous eudicots.

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 Pulling Xylem Sap:
The Transpiration-Cohesion Tension Mechanism

• Water is pulled upward by negative pressure in

the xylem.

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 Transpirational Pull

• Water vapor in the airspaces of a leaf diffuses

down its water potential gradient and exits the leaf
via stomata.
• Transpiration produces negative pressure
(tension) in the leaf, which exerts a pulling force
on water in the xylem, pulling water into the leaf.

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Water Movement out of the Leaf

Y = –0.15 MPa Y = –10.00 MPa

Cell wall
Air-water
Airspace interface

Low rate of High rate of

Cuticle transpiration transpiration
Upper
epidermis

Cytoplasm
Evaporation
Mesophyll Airspace
Air
space Cell wall

Lower Evaporation
epidermis Water film Vacuole

Cuticle Stoma
CO2 O2 CO2 O2
Xylem
 Water Movement thru Leaf Xylem:
Governed by Water Potential
• Water flows from the xylem of the leaves, where water
potential is least negative, toward air, where water
potential is most negative.

 Y gradients :
tracheids > sheath
sheath > apoplast
sheath > spongy mesophyll
spongy mesophyll > apoplast
apoplast > stoma
(outside air  -10 MPa)

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 Cohesion and Adhesion in the Ascent of Xylem Sap

• The transpirational pull on xylem sap is

transmitted all the way from the leaves to the root
tips and even into the soil solution.
• Transpirational pull is facilitated by cohesion and
adhesion.

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 Ascent of Xylem Sap by Bulk Flow:
A Take-Home Lesson

• The movement of xylem sap against gravity is

maintained by the transpiration-cohesion-
tension mechanism.

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 Xylem
sap
Outside air  Mesophyll
= –100.0 MPa cells
Stoma
Leaf  (air spaces) Water
= –7.0 MPa molecule
Transpiration
Leaf  (cell walls) Atmosphere
= –1.0 MPa

Xylem Adhesion Cell

cells wall

Water potential gradient

Trunk xylem 
= –0.8 Mpa Cohesion,
Cohesion and by
adhesion in hydrogen
the xylem bonding

Water
molecule
Root
Root xylem  hair
= –0.6 MPa Soil
particle
Soil 
= –0.3 MPa Water
Water uptake
from soil
 Ascent of Xylem Sap: Summary

• H-bonding forms an unbroken chain of water molecules

extending from leaves all the way to the soil.
• Y gradient: the force that drives the ascent of xylem sap.
• For the bulk flow over long distance, the Y gradient is due
mainly to pressure potential gradient (Yp).
• Transpiration results in the Yp at the leaf end of the xylem
being lower than the Yp at the root end.

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Stomata

20 µm
 Stomata: Major Pathways for Water Loss

• About 90% of the water a plant loses escapes

through stomata.
• Each stoma is flanked by a pair of guard cells,
which control the diameter of the stoma by
changing shape.

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 Stomata help regulate the rate of transpiration

• Leaves generally have broad surface areas and

high surface-to-volume ratios.
• These characteristics increase photosynthesis and
increase water loss through stomata.

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 Effects of Transpiration on Wilting and Leaf
Temperature

• Plants lose a large amount of water by

transpiration.
• If the lost water is not replaced by absorption
through the roots, the plant will lose water and wilt.

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 • Transpiration also results in evaporative cooling,
which can lower the temperature of a leaf and
prevent denaturation of various enzymes involved
in photosynthesis and other metabolic processes.

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 Cells turgid/Stoma open Cells flaccid/Stoma closed

Radially oriented
cellulose microfibrils

Cell
wall

Vacuole
Guard cell

Changes in guard cell shape and stomatal opening and closing

(surface view)
 • Changes in turgor pressure that open and close
stomata result primarily from the reversible uptake
and loss of potassium ions by the guard cells.

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 Cells turgid/Stoma open Cells flaccid/Stoma closed

H 2O H 2O H 2O
H 2O

H 2O
K+ H 2O

H 2O
H 2O H 2O H 2O

Role of potassium in stomatal opening and closing

 Xerophyte Adaptations That Reduce Transpiration

• Xerophytes are plants adapted to arid climates.

• They have leaf modifications that reduce the rate
of transpiration.
• Their stomata are concentrated on the lower leaf
surface, often in depressions that provide shelter
from dry wind.

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Cuticle Upper epidermal tissue

Lower epidermal Trichomes Stomata

tissue (“hairs”) 100 µm