Water and Xylem Transport

Water transport in the xylem
In most plants, the xylem

constitutes the longest part of the
whole plant water transport pathway
For a 1 m tall, more than 99.5% of
the water transport pathway will
occur within the xylem.
Compared to the root, xylem water
transport is a fairly simple low
resistance transport pathway.
We will be considering now:
The role of negative pressures
in xylem transport
And the importance of xylem
transport safety under negative
pressure
Negative or positive pressure driven

water flow?
Pressure gradients needed to move water through the xylem could come
about in two ways:
Positive pressures- hypothetically positive pressure could drive water
to the tops of trees, but
root pressures are generally less than 0.1 MPa (lifts water only 1
m above the ground) and pressures of at least 2 MPa are required
to supply some of the tallest trees on the planet
Root pressure is energetically expensive...
Also, evaporation will easily collapse positive pressure
gradients...
Negative pressures - Solar power! Instead plants use the evaporative
power of solar energy to pull water through the vascular system.
So instead, water at the tops of trees develops a large tension to pull
water through the xylem
Cohesion-tension theory of sap ascent
The well-supported theory of sap

ascent is that water moves up
plants under tension
Requires that the cohesive
properties of water sustain large
tensions in the xylem water column
Despite the attractiveness of the
theory, still continues to generate
controversy; mainly along whether
tension exists in the xylem.
Pressure probe studies failing to
find high tensions
Later refinements
Xylem water transport under tension is

risky
The large tensions that develop in the
xylem of trees can create some
problems
First, tension results in an INWARD
pull on the tracheary cell system. The
development of lignified walls is
necessary to allow resistance to
implosion from this force...
Metastable state of water under

tension; air entry problems
A second problem is that water is METASTABLE and very

sensitive to slight changes in gas-content
Recall that pure degassed water is very strong, but with gas
added, the water column can become increasingly easily broken
As tensions increase, there is an increased tendency for air to be
pulled in from microscopic pores in the xylem wall that contain air
(from respiring living cells or just close to lenticels etc.)
This is called air-seeding.
Another way that air can be introduced into the xylem is when the
sap freezes (bubbles form) and if this occurs when the xylem is
under tension, then after thawing the bubbles will expand under
tension
Embolism or cavitation
This phenomenon of bubble formation is called cavitation or

embolism.
Analogous to a vapor lock in a fuel line or an air bubble in a blood
vessel
Breaks like this in the water column are not all that unusual
However, these breaks in xylem water continuity, if not repaired,
end up blocking xylem water transport in the plant
Cavitation blocks xylem water

transport
Note that water can detour,

but as the number of blocked
vessels or tracheids
increases, transpirational
surfaces of the plant are
supplied with less and less
water.
Plants attempt to minimize the

consequences of xylem cavitation
Gas bubbles can be eliminated from the xylem

At night by root pressure, by dissolving gas back into solution
of the xylem
Recent data suggests that bubbles may be collapsed even
under tension; but mechanism is not known.
Or, plants may grow out of the effects of cavitation; add new
sapwood to replace older embolized vessels; we see this in ringporous plants where the early wood vessels become blocked and
are not refilled year to year
Root pressure and in vivo embolism refilling
From Holbrook et
al. (2001) Plant
Physiology
Water evaporation in the leaf generates

a negative pressure on the xylem
IMPORTANT: The tensions

need to pull water through the
xylem are the result of
evaporation of water from
leaves.
This tension develops at the
surface of the cell walls in the
leaf that are in contact with the
air.
The analogous to the situation
in soil
As evaporation increases and leaf

water content declines, the motive
force for water transport increases
Note how the curvature changes
Curved water surfaces at the

cellulose microfibrils in the leaf
cells become smaller and smaller.
The p can be estimated as: p = -2
T/r
where T is surface tension of
water (7.28 x 10-8 MPa m) and r
is the radius of the curvature at
an air water interface
Water movement from the leaf to the

atmosphere
After water has

evaporated from the
leaf, diffusion is the
primary means of
water movement out
of the leaf.
The rate of diffusion
out the leaf is
controlled by three
interacting pathways
cuticular
stomatal
boundary layer
Diffusion and the water vapor

concentration gradient
Because movement of water

out of the leaf is based on
diffusion, the gradient in water
vapor concentration from
inside the leaf intercellular air
spaces to outside controls the
rate of diffusion.
The atmosphere gradient is
strongly dependent on
temperature
Diffusional water loss regulated

resistors along the pathway
Cuticular
direct diffusion through
the cuticle, generally
very small (~ 5% of
total flux)
Stomatal and boundary
layer
These two processes
interact, stomata
generally control 95%
of the water loss by
changes in their
aperture; but the
boundary layer can
become important
Stomatal control and

non-control of leaf
water loss rate
Measurements of leaf water loss rates
When can boundary layer resistance

override stomatal resistance to leaf
water loss?
Still air condition

Plants with big leaves
Plants that have stomata covered or embedded (sunken deep) in
the leaf
generally thought to be drought-avoid structures
But remember waxy stomatal plugs in Drimys?
Stomatal function
Guard cells function as

multisensory hydraulic
valves
respond to light,
temperature,
humidity, intercellular
CO2, and water
potential of the leaf
An increase in guard
cell turgor causes
stomatal pores to
open.
Cellulose microfibril orientation and guard

cell bending
A distinctive feature of guard

cells are their cell wall
structure
Portions of the wall are
thickened
Also, the alignment of
cellulose microfibrils, which
recall determine cell shape,
are oriented radially and this
plays an important role in their
movement
cellulose microfibrils are
oriented like steel-belts in a
radial tire
Overview of the soil-plant-atmospheric

continuum
OK, a brief word about phloem transport: the

pressure flow hypothesis
In lab 8

Water and Xylem Transport

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Water transport in the xylem

In most plants, the xylem

Negative or positive pressure driven

Cohesion-tension theory of sap ascent

The well-supported theory of sap

Xylem water transport under tension is

Metastable state of water under

A second problem is that water is METASTABLE and very

This phenomenon of bubble formation is called cavitation or

Cavitation blocks xylem water

Note that water can detour,

Plants attempt to minimize the

Gas bubbles can be eliminated from the xylem

Root pressure and in vivo embolism refilling

Water evaporation in the leaf generates

IMPORTANT: The tensions

As evaporation increases and leaf

Note how the curvature changes

Curved water surfaces at the

Water movement from the leaf to the

After water has

Diffusion and the water vapor

Because movement of water

Diffusional water loss regulated

Stomatal control and

Measurements of leaf water loss rates

When can boundary layer resistance

Still air condition

Guard cells function as

Cellulose microfibril orientation and guard

A distinctive feature of guard

Overview of the soil-plant-atmospheric

OK, a brief word about phloem transport: the

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