WATER BALANCE

OF PLANTS

KRISPINUS K. PUKAN
Life in earth’s atmosphere presents a formidable challenge to
land plants. On the one hand, the atmosphere is the source
of CO2, which is needed for photosynthesis. Plants therefore
need ready to access to the atmosphere. On the other hand,
the atmosphere is relatively dry and can dehydrate the
plant.
To meet the contradictory demands of maximizing CO2
uptake while limiting water loss, plant have evolved
adaptations to control water loos from the leaves, and to
replace the water lost to the atmosphere
Transpirational water loss from the leaf is driven by a
gradient of water vapor concentration
Long distance transport in the xylem is driven by pressure
gradient, as is water movement in the soil
Water transport through cell layers such as the root cortex is
complex, but it responds to water potential gradient a cross
the tissue
Troughout this journey water transport is passive, in
the sense that the free energy of water decreases as
it moves
WATER IN THE SOIL
The water content and the rate of water movement in soil
depends to a large extent on soil type and soil structure
Sandy soil have a relatively low surface area per gram of soil
and have large spaces or channels between particles
Clay soils have much greater surface areas and smaller
channels between particles
With the aid of organic substances such as humus, clay
particles may aggregate into “crumbs” allowing large
channels to form that help improve soil aeration and
infiltration of water
When soil is heavely watered by rain or by irrigation, the
water percolates downward by gravity through the spaces
between soil particles, partly displacing and in some cases
traping, air in the channels
Because water is pulled into the spaces between soil particles
by capillarity, the smaller channels becomes filled first.
Depending on the amount of water available, water in the soil
may exist as a film adhering to the surface of soil particles,
it may fill the smaller but not the larger channels, or it may
fill all of the spaces between particles.
WATER ABSORBTION BY ROOT
Intimate contact between the surface of the root and the soil
is esential for effective water absorbtion by the root
This contact provides the surface area nedeed for water
uptake and is maximazed by the growth of the root and of
root hairs into the soil
Root hairs that greatly increase the surface area of the root,
thus providing greater capacity for absorbtion of ions and
water from the soil
Water enters the root most readly near the root tip.
Mature regions of the root are less permeable to
water, because they have developed an outer layer of
protective tissue, called exodermis or hypodermis,
wich contains hydrophobic materials in its walls
WATER MOVES IN THE ROOT VIA APOPLAST, SYMPLAST AND TRANSMEMBRANE
PATHWAYS

The apoplast is the continous system of cell walls,

intercellular air spaces, and the lumens of nonliving cells
(e.g xylem conduits and fibers). In this pathway water
moves through cell walls and extracellular spaces
without crossing any membrane as it travels across the
root cortex

The symplast consist of the entire network of cell sytoplasm

interconected by plasmodesmata. In this pathway, water
travels the root cortex via palsmodesmata
The transmembrane pathway is the route by which waters
enters a cell on one side, exits the cell on the other side
enters the next in the series, and so on. In this pathway,
water crossis the plasma membrane of each cell in its
path twice. Transport across the tonoplast may also be
involved
The Casparian strip break the continuity of the apoplast
pathway, forcing water and solutes to pass through the
plasma membrane in order to cross the endodermis
Water uptake decreases when root are subjected to low
temperature or anaerobic condition or treated with
respiratory inhibitors
Decreased rates of respiration, in respons to low
temperature or anaerobic conditions, can lead to
increase in intracellular pH. This increase in
cytoplasmic pH alter the conductance of aquaporins
in root cells, resulting in roots that are markedly less
permeable to water.
SOLUTE ACCUMULATION IN THE XYLEM CAN GENERATE “ROOT PRESSURE ”

When transpiration is low or absent, positive

hydrostatic pressure builds up in the xylem because
root continue to absorb ions from the soil and
transport them into the xylem
The building up solutes in the xylem sap leads to a
decrease in the xylem osmotic potentil (Ψs) and thus a
decrease in the xylem water potential (Ψw)
This lowering of the xylem Ψw provides a driving force
for water absorbtion, which in turn leads to a positive
hydrostatic preassure in the xylem
Root preassure in most likely to occure when
soil water potentials are high and
transpiration rates are low
When transpiration rates are high, water is
taken up into the leaves and lost to the
atmosphere so rapidly that a positive
pressure resulting from ion uptake never
develops in the xylem
Plants that develop root pressure frequently
produce liquid droplets on the edges of their
leaves,a phenomenon known as guttation
WATER TRANSPORT THROUGH THE XYLEM
The xylem consist of two types of tracheary elements are
tracheids and vessel elements
To conducting cells in the xylem have a specialized
anatomy that enables them to transport large quantities
of water with great efficiency
Vessel element are found in angiosperm, a small group of
gymnosperm, called Gnetales, and some ferns.
Tracheid are present in both angiosperm and
gymnosperm, as well as in ferns and other group of
vascular plants.
Water moves through the xylem by pressure-driven
bulk flow
Pressure-driven bulk flow of water is responsible for
long-distance transport of water in the xylem
If we consider bulk flow through a tube, the rate of
flow depends on the radius (r) of the tube, the
viscosity (η) of the liquid, and the pressure
gradient (ΔΨp/Δχ) that drives the flow. Volume
flow rate = [ see page 92]
WATER MOVEMENT THROUGH THE XYLEM REQUIRE A
SMALLER PRESSURE GRADIENT THAN MOVEMENT
THROUGH LIVING CELLS

The xylem provides a pathway of low resistivity

for water movement
Velocity of water transport in xylem is 4 mm
per second, r = 40 μm
Driving force needed to move water from cell to
cell, crossing the plasma membrane at 4 mm
per second is 2 x 108 Mpa m-1
WHAT PRESSURE DIFFERENCES IS NEEDED TO
LIFT WATER 100 METERS TO A TREETOP?

The pressure gradients needed to move water through

the xylem of very tall trees are on the order of 0.01
Mpa m-1, smaller than in our previous example.
If we multiplay this pressure gradient by the height of
the tree (0.01 x 100 m) we find the total pressure
difference needed to overcome the frictional
resistance to water movement to the stem is equal to
1 MPa
For a hight differnce of 100 m, the difference in Ψg is
approximately 1 Mpa higher at the top of the tree
than at the ground level.
So the other components of water potential must be 1
Mpa more negative at the top of the tree to counter
the effect of gravity.
To allow transpiration to occur , the prssure gradient
due to gravity must be added to that require to cause
water movement through the xylem
Thus we calculate that a pressure difference of roughly
2 Mpa from the base to the top branches, is needed to
carry water up the tallest tree.
THE COHESION-TENSION THEORY EXPLAIN WATER
TRANSPORT IN THE XYLEM
In theory , the pressure gradient needed to move water
through the xylem could result from the generation of
positive pressure at the base of the plant or negative
pressure at the top of the plant.
The water at the top of tree developes a large tension (a
negative hydrostatic pressure), as this tension pulls
water trhough the xylem
PLANT MINIMIZE THE CONSEQUENCES OF XYLEM
CAVITATION
At night , when transpiration is low, xylem Ψp increase,
and water vapor and gases may simply dissolve back
into the soulution of the xylem
Recent studies indicate that cavitation may be repaired
even when the water in the xylem is under tension
Finally, many plants have secondary growth in wich
new xylem form each year. The production of new
xylem conduits allows plants to replase looses in
water transport capasity due to cavitation
WATER MOVEMENT FROM THE LEAF TO THE
ATMOSPHERE
Leaf have a large hydraulic resistance
The driving force for transpiration is the difference in
water vapor cosentration
WATER LOSS IS ALSO REGULATE BY THE PATHWAY
RESISTANCES
The second important factor governing water loss from
the leaf is the diffusional resistance of the
transpiration pathway, which consist of two varying
components.
1. The resistance associated with diffusion through the
stomatal pore, the leaf stomatal resistance (rs)
2. The resistance due to the layer of unstirred air next
to the leaf surface through which water vapor must
diffuse to reach the turbulent air of the atmosphere.
This second resistance , rb, is called the leaf
boundary layer resistance
THE TRANSPIRATION RATE MEASURE THE RELATIONSHIP BETWEEN
WATER AND CARBON GAIN

The effectivness of plants in moderating water loss while allowing

sufficient CO2 uptake for photosynthesis can be assessed by a
parameter called the Transpiration Ratio. This value is defined as
amount of water transpired by the plant divided by the amount of
carbon dioxide assimilated by photosynthesis
For plants in which the first stable product of carbon fixation is a 3-
carbon compound as manynas 400 molecules of water are lost for
every molecule of CO2 fixed by photosynthesis , giving a
transpiration ratio of 400.Plants with a transpiration ratio of 400
have a water use efficiency of 1/400 or 0.0025
THE LARGE RATIO OF H2O EFFLUX TO CO2 INFFLUX RESULT FROM THREE
FACTORS :

1. The concentration gradient driving water loss is about 50 times

larger than that driving the infflux of CO2 In the large part, this
difference is due to the low concentration of CO2 in the air (about
0.038%) and the relatively high concentration of water vapor within
the leaf.
2. CO2 diffuses about 1.6 times more slowly through air than water
does (the CO2 molecule is larger than H2O and has a smaller
diffusion coefficient
3. CO2 must cross the plasma membrane , the cytoplasm, and the
chloroplast envelope before it is assimilated in the chloroplast.
These membranes add to the resistance of the CO2 diffusion
pathway.