Whole Plant Water Relations

Chapter 2
 Learning objectives
Understand the driving force behind
transpiration.

Know how environmental factors influence

the rate of transpiration water loss.

Know the structure of water-conducting

tissues in plants.
 Learning objectives
Understand why a continuity of the water
column is essential to water transport and
plant water relations.

Understand how soil water is transported

into roots.
 Transpiration
Plants lose ~95% of the water to the
atmosphere through transpiration.

Transpiration is the loss of water from

plants as water vapor.

Water loss from other surfaces is reduced

by the presence of a waxes and the
cuticle.
 Transpiration
Water escapes primarily through the
stomata, but also through lenticels in the
stem.

Water loss through the stomata is regulated

by the guard cells.

The water transpired is principally water

vapor in the intercellular spaces
surrounding the leaf mesophyll cells.
 Transpiration
Figure 2.1
 Transpiration
Transpiration is a two-stage process.
Water evaporates from the mesophyll cell
walls into the substomatal air.
The water vapor diffuses through the
stomates into the atmosphere.

Other researchers have suggested that

transpiration involves peristomal
evaporation instead.
 Transpiration
Transpiration from leaves occurs via:
Stomatal transpiration, which is passage of
water through the stomates.
Cuticular transpiration, which is water loss
directly through the cuticular layer.

Cuticular transpiration is thought to be

only 5-10% of the total transpiration.
 Transpiration
The driving force for transpiration is a
difference in vapor pressure between the
substomatal and atmospheric water.

The concentration of water in the vapor

phase is the vapor density (g m-3).

The water concentration can be expressed

as the vapor pressure (e or kPA).
 Transpiration
When the gas phase is saturated with water, it
has reached the saturation vapor pressure.

The vapor pressure can be quantified with

Raoults law.
o
e = Xie
e is the vapor pressure
Xi is the mole fraction of water
eo is the saturation vapor pressure of the solvent
 Transpiration
Figure 2.2
 Transpiration
The rate of transpiration is affected by:
Humidity
Temperature
Wind speed

Transpiration (T) is governed by the

magnitude of the difference in vapor
pressure between the leaf and air.
T eleaf - eair
 Transpiration
Transpired water encounters resistance to
movement.
The movement through the intercellular
spaces is tortuous.
Movement through the stomata represents a
constriction.
The unstirred boundary layer impedes water
movement to the bulk air.
 Transpiration
The resistances (r) to movement can be
incorporated into the mathematical
expression for transpiration.
eleaf - eair
T
rleaf - rair
 Transpiration
Figure 2.3
 Transpiration
Relative humidity (RH) is the ratio of
actual water content to maximum water
content.

Precent relative humidity is the actual

vapor pressure relative to the saturation
vapor pressure.

Temperature increases the capacity of air to

hold water.
 Transpiration
Table 2.1
 Transpiration
The relative humidity of the air contributes to the
airs water potential ().

As humidity decreases, water potential becomes

more negative.

Since substomatal air is more humid, it has a

more positive water potential that bulk air, so
transpiration is the movement of water down its
water potential gradient.
 Transpiration
Table 2.2
 Transpiration
Temperature also affects the vapor
pressure gradient, increasing it as
temperature increases through the day.

This insures that transpiration occurs,

even as the relative humidity of the air
approaches 100%.
 Transpiration
Table 2.3
 Transpiration
Wind influences transpiration by affecting the
distance over which water must move.

Increasing wind velocity disrupts the

boundary layer and enhances transpiration
because the distance of water movement
decreases.

Wind movement also cools and dries the leaf,

which may further increase transpiration.
 Transpiration
Figure 2.4
 Water conducting tissues
The plant vascular tissues consist of the:
Xylem
Phloem

Xylem consists of three cell types.

Fibers are elongated cell with thickened cell
walls for structural support.
Parenchyma cells provide storage.
Tracheary elements, which consist of
tracheids and vessel elements, are the
conducting cells.
 Water conducting tissues
Figure 2.5
 Water conducting tissues
Tracheid and vessel element structure
provides for the efficient movement of
water through the xylem.

The tracheids have bordered pit pairs

with a central torus that provides a way of
sealing the tracheids if necessary.
 Water conducting tissues
Figure 2.6
 Water conducting tissues
Vessel elements are made up of vessel
members that have a perforation plate
at each end.

The perforation plate may have a

scalariform or reticulate structure.

Vessels are more complex than tracheids.

 Water conducting tissues
Figure 2.7
 Water conducting tissues
The movement of water through the xylem
is governed by Poiseuilles equation.
P r 4
Jv =
8

Jv is the flux rate.

The pressure gradient is P.
The solution viscosity is
 Sap ascent in the xylem
Atmospheric pressure can raise a column
of water only about 10 m, but xylem
transport must reach greater heights.

The mechanism of xylem transport

involves:
Root pressure
Capillarity
Cohesion theory
 Sap ascent in the xylem
Figure 2.8
 Sap ascent in the xylem
Root pressure is a positive pressure at the
root level that pushes water up the xylem.

Root pressure is a function of:

Root structure
Water uptake
Mineral uptake
 Sap ascent in the xylem
Root pressure is generated as follows:
Roots take up mineral ions from the soil.
The change in solute potential decreases
water potential and induces water uptake.
Water moves radially to the endodermis
surrounding the stele.
The endodermis has a Casparian band made
of suberin.
 Sap ascent in the xylem
Root pressure is generated as follows:
The Casparian band prevents the movement
of water back to the cortex
Therefore the continual influx of water causes
a positive pressure to develop which forces
water up the stem in the xylem.

Root pressure can be measured with a

manometer.
 Sap ascent in the xylem
Figure 2.10
 Sap ascent in the xylem
Figure 2.9
 Sap ascent in the xylem
Root pressure can only raise a column of
water a short distance so it is insufficient
to raise water to the necessary height.

Measurements have also shown that the

xylem sap is under negative pressure, or
tension, so root pressure cannot be the
driving force.
 Sap ascent in the xylem
Capillarity contributes to the rise of sap in
the xylem.

Capillarity occurs because water shows

adhesion, an attraction to solid surfaces.

The surface tension of water also

contributes to capillary rise.
 Sap ascent in the xylem
The capillary rise in the xylem can be calculated
using this equation:

1.49x10-5 m 2
h=
rm
h is the capillary rise
r is the radius (in meters)

For the radius of a typical xylem element, capillarity

would raise the water column <1 m.
 Sap ascent in the xylem
The current hypothesis describing sap
ascent is the cohesion theory.
Water evaporation from cell walls of
mesophyll cells in leaf is the driving force.
As the water evaporates, it creates a tension
in the minute contours of the wall.
Because water is cohesive, the tensile
strength that develops pulls the continuous
network of water up from the soil to leaves.
 Sap ascent in the xylem
Figure 2.11
 Sap ascent in the xylem
When water is under tension in the xylem,
it is in a metastable state and dissolved
gases are released as bubbles.

This formation of bubbles is called

cavitation.

These bubbles can cause an embolism, a

blockage of xylem vessels.
 Sap ascent in the xylem
Figure 2.12
 Sap ascent in the xylem
The structural nature of xylem elements
allows embolisms to be overcome.

The bordered pits in xylem elements

allows water to bypass emoblized vessels
to keep xylem flow moving.

As the xylem tension decreases during the

night, some embolisms will disappear.
 Sap ascent in the xylem
Embolisms may actually serve beneficial roles in
terms of preventing excess water loss during
stress.

The designed leakage hypothesis proposes

that plants use cavitation to protect themselves
when water potentials become too negative.

Root are also prone to cavitation, particularly

when first transplanted into new soil.
The soil-plant-atmosphere continuum
Water movement through plants relies on an
integrated connection of water from the soil,
through the roots, stem, and leaves, and into the
atmosphere.

This continuum must be maintained.

This means that plants are highly dependent

upon the environment and soil water.
The soil-plant-atmosphere continuum

Water is obtained principally from the soil.

Soil has three phases.

The solid surfaces
The soil solution phase
A gaseous phase

Soil also contains various organic

components and living organisms.
The soil-plant-atmosphere continuum

The size of the soil particles determines

the characteristics of the soil, including the
retention of water.
Soils have a porosity composed of large
pores and capillary pores, which together
account for 40-60% of the soil volume.
When the pore space is filled with water, the
soil is at field capacity.
The soil-plant-atmosphere continuum

Water in soils is subject to the same

forces as water in plants.
The curvature of soil particles create the
same adhesive force as cell wall water.
Water in the pore space shows cohesion and
tension, and can be drawn to roots.
Water moves in the soil primarily by pressure-
driven mass flow.
The soil-plant-atmosphere continuum

The water potential of the soil and the

roots determines water movement.
Root uptake of water can only occur if the
water potential of root cells is more negative
that the soil water potential.
The soil water percentage at which the root is
unable to obtain water corresponds to the
permanent wilting percentage.
The water content between field capacity and
permanent wilting percentage is available
water.
 Uptake of water by roots
The functions of roots include:
Anchoring the plant in the soil.
Storing carbohydrates and other molecules.
Synthesis of some hormones and secondary
metabolites.
Uptake and transport of water and minerals.

In order to provide the uptake and transport

needed, root systems need to be extensive in
length and surface area.
 Uptake of water by roots
Roots take up water primarily at the tips.

The uptake of water occurs along other

root sections as a function of age,
physiological condition, and water status.

Water is also taken up by the root along

the zone of maturation.
 Uptake of water by roots
Figure 2.13
 Uptake of water by roots
Root hairs are also primary sites for water
uptake.

Root hairs are outgrowths of the

membrane of individual epidermal cells.

The small diameter allows root hairs to

explore smaller pore spaces in the soil to
obtain water.
 Uptake of water by roots
Figure 2.14
 Radial movement of water in roots
Water moves across the diameter of the root to
the stele by two pathways.
The apoplast represents the continuity of the cell wall
space outside of the cells.
The symplast represents the connection of cellular
cytoplasm between cells provided by the
plasmodesmata.

Water movement radially also varies depending

upon whether the suberization of the
endodermis in that area of the root has fully
developed.
 Radial movement of water in roots
This radial movement of water creates a lag
between uptake and transpiration.

If transpiration exceeds root uptake of water,

there may be a midday closure of the stomates
while the balance is restored.

Water uptake is dependent upon physiological

processes in the root, such as respiration
Radial movement of water in roots
Figure 2.15