Plant Transport Mechanisms

Summary of Lecture
 The Movement of Water and Minerals
o Root Pressure
o Water Potential and Vascular Plants
o Leaves: Transpiration and Pulling of Water
o Water Movement in Xylem through TACT Mechanisms
o Water Transport in the Root
o Transpiration and photosynthesis - a compromise
o Guard Cells and Water Transport
 The physical structure of guard cells
 Why guard cells change shape
 Factors triggering the change in shape
o Strategies for maximizing the availability of CO2
 C4 Photosynthesis
 CAM Photosynthesis
 Food Transport in Plants
o Mechanisms of Phloem Transport
o Flow from Source to Sink
o The Pressure Flow Hypothesis
 Gas Transport in Plants
 Mineral Nutrients and their Transport

Syllabus Transport in Plants

The Movement of Water and Minerals
Review:

 the role of water in metabolism

 the water cycle in ecology.

Root Pressure

As various ions from the soil are actively secreted into the root's vascular tissue water
follows (its potential gradient) and osmotic pressure increases.

This osmotic pressure is called root pressure.

Root pressure is observable at

night when evaporation is low
and excess water collects in
droplets around special openings
near the tip of grass blades.

Such water loss in its liquid phase

is known as guttation.
Note: the root endoderm because of a waxy layer of subrin has the ability to actively
transport ions in one direction only.

Root pressure can only provide a modest push in the overall process of water
transport. Its greatest contribution maybe to reestablish the continuous chains of water
molecules in the xylem which often break under the enormous tensions created by
transpiration.
Water Potential and Vascular Plants

When a water potential gradient is established between two areas, water will
spontaneously diffuse from the high end (soil) to the low end (air). This gradient is
necessary for plants to transport water.

Water potential may be established by:

 increasing the concentration of solutes. Pure water has the highest

potential while a saturated solution of ions etc. would have the
lowest potential.
 converting water to a gas. Water potential is highest when water is
a liquid and lowest when water is a gas in air.

Visit Biology 184 or an essay by Anne Bruce for a through explanation of Water
Transport

See below for my explanation of water transport.

Leaves: Transpiration and Pulling of Water

Photosynthesis requires water. The system of xylem vessels from root to leaf vein can
supply the needed water.

What force does a plant use to move water molecules into the leaf parenchyma cells where
they are needed? Read on!

Several forces combine to overcome the pull of gravity.

These combined forces culminate in a process called transpiration.

Ultimately water is pulled, molecule by molecule into the leaf. The pulling forces and
energy needed involves:

1. free energy of the water potential gradient

2. free energy of evaporation
3. force of surface tension
4. force of hydrogen bonding between water molecules
Each force can be communicated to the next because water forms a strong continuous
chain from root to leaf.

1. Water moves in the direction it does (root to leaf) because of the water
potential gradient. The gradient is highest in the water surrounding the
roots and lowest in the air space within the spongy parenchyma of the
leaf. (liquids have higher potential than gases and the purer the liquid the
higher its potential)

2. The energy of evaporation is needed to to pull molecules away from the

film of water coating air spaces within the spongy parenchyma. (see
diagram below)

3. As more molecules evaporate from the film coating the air spaces the
curvature of the meniscus increases which increases the surface tension.
Water from surrounding cells and air spaces will then be pulled towards
this area to reduce the tension.
 4. Finally these forces are communicated to water molecules within the
xylem because each water molecule is bound to the next by hydrogen
bonds.

Water Potential and the leaf

Evaporation from the leaf sets up a

water potential gradient between the
outside air and the leaf's air spaces.
The gradient is transmitted into the
photosynthetic cells and on to the
water-filled xylem in the leaf vein.

(Wallace)

Measurements reveal that the forces generated by transpiration can create pressures up
to 12 atmospheres, sufficient to lift a xylem sized column of water over 350 feet high
(130 meters).

Water Movement in Xylem through TACT Mechanism

Four important forces combine to transport water solutions from the roots, through the
xylem elements, and into the leaves. These TACT forces are:

 transpiration
 adhesion
 cohesion
 tension
Transpiration involves the pulling of water up through the xylem of a plant utilizing
the energy of evaporation and the tensile strength of water. The previous section
describes transpiration more fully.

Adhesion is the attractive force between water molecules and other substances.
Because both water and cellulose are polar molecules there is a strong attraction for
water within the hollow capillaries of the xylem.

Cohesion is the attractive force between molecules of the same substance. Water has
an unusually high cohesive force again due to the 4 hydrogen bonds each water
molecule potentially has with any other water molecule. It is estimated that water's
cohesive force within xylem give it a tensile strength equivalent to that of a steel wire
of similar diameter.

A combination of adhesion, cohesion, and

surface tension (see below) allow water to
climb the walls of small diameter tubes like
xylem. This is called capillary action. The
U shaped surface formed by water as it
climbs the walls of the tube is called a
meniscus.

Tension can be thought of as a stress placed on an object by a pulling force. This

pulling force is created by the surface tension which develops in the leaf's air spaces.

As water molecules leave the surface film by evaporating

into the air spaces the remaining film forms menisci (a, b, c)
which become more and more concave. A meniscus has a
tension that is inversely proportional to the radius of the
curved water surface.

In other words, as the water surface becomes more curved tension increases. "Tension
is a negative pressure -- a force that pulls water from locations where the water
potential is greater." Campbell

The bulk flow of water to the top of a plant is driven by solar energy since
evaporation from leaves is responsible for transpiration pull.
Water Transport in the Root

The flow of water and minerals from the soil to the cells of the root is accomplished
by transpirational pull, active transport and a special layer of cells called the
casparian strip.

Active transport establishes a lower water potential and helps the root hairs take in the
necessary minerals dissolved in soil water. A lower water potential allows water to be
drawn into the root cells by osmosis.

In order to regulate the quantity and type of minerals and ions reach the xylem, the
root has a waxy layer between the endodermis and pericycle called the casparian strip.
Water and mineral normally can travel through the porous cell walls of the root cortex
-- this is the apoplastic route. But in order for water and minerals to reach the stele
(xylem) the highly regulated (cytoplasmic) symplastic route must be taken. The
apoplastic route is blocked by the casparian strip.

The symplastic route involves special openings between adjacent cell walls called
plasmodesmata. (see below)

Paths that solutions can take through plant cells

Symplastic vs Apoplastic paths

Transpiration and photosynthesis - a compromise
An actively photosynthesizing plant has an insatiable need for water. The efficiency
of photosynthesis increases as the surface area for CO 2, the second reactant, increases.
This is the purpose of the spongy mesophyll which is honeycombed with air sacs. The
spongy mesophyll has 10 to 30 times more surface area than the corresponding
external surface of a leaf.

Photosynthesis is limited by available water which can be swiftly depleted by

transpiration. It has been estimated that for every gram of CO 2 fixed by the Calvin
cycle, anywhere from 300 to 600 grams of water escape through transpiration. The
humidity of rainforests is due in large part to this vast cycling of water from root to
leaf to atmosphere and back to the soil.

Transpiration has more that one purpose however, it:

 supplies water for photosynthesis

 transports minerals from the soil to all parts of the plant
 cools leaf surfaces some 10 to 15 degrees by evaporative cooling
 maintains the plant's shape and structure by keeping cells turgid

Guard Cells and Water Transport

 The physical structure of guard cells

A stoma is a physical gap between two special epidermal cells called

guard cells. When the pair of guard cells are turgid -- full of water -- they
bow in such a way as to increase the gap -- stoma -- between them.

If the plant experiences water deprivation it will wilt. To compensate the

guard cells become flaccid and the stoma is closed.

The structure of guard cells explains why they bow apart when turgid.

1. The two guard cells are fused at their ends.

2. The inner cell walls which form the stoma are thicker than the
outer walls.
3. Cellulose microfibrils are oriented radially rather than
longitudinally.

 
 

 The sequence of events which result in stomatal opening

The immediate cause is an increase in turgor pressure - water enters the

central vacuole by osmosis

Turgor pressure increases because of a negative water potential due to an

influx of potassium ions (K+). The cell becomes hypertonic to its
environment

The reversible uptake of K+ ions takes place because of the membrane

potential created when H+ are actively pumped out of the cell -
consuming ATP. The cell's interior becomes negative compared to the
surroundings.

The stoma is closed at night when
the large central vacuole is
isotonic, even hypotonic to
surrounding fluids. K+ ions are
outside of the cell, and H+ ions by
and large remain attached to the
weak organic acids within the cell.
With H+ on the outside K+ readily
Blue light is absorbed by a
diffuse into the cell to compensate
membrane protein which
for the negative electrical
somehow causes an
potential. The hypertonic
increase in the activity of
conditions within the cell attract
proton pumps which use
water molecules and the stoma
ATP to transport H+ out of
opens as turgor pressure
the cell.
increases.

 What factors trigger the change of shape in guard cells

1. increase in blue light at dawn - a blue light sensitive receptor
activates proton pumps and turgor pressure increases. Light also
stimulates the photosynthetic production of ATP
2. Absence of CO2
3. Circadian rhythms - All eukaryotic cells have chemical based
metabolic clocks entrained to the day-night cycle. A common 24
hour biological clock is responsible for circadian rhythms (circa,
about - dies, day)

 
Strategies for maximizing the availability of CO2 while minimizing water loss have
evolved in land plants.

 C4 Photosynthesis

C4 plants are twice as efficient as C 3 varieties in terms of fixing carbon

(making sugar). A C4 plant will lose "only" 300 grams of water by
evaporation for every gram of CO2 fixed by photosynthesis whereas C 3
plants lose 600 grams of water for the same grams of CO 2 fixed.

 CAM Photosynthesis

A CAM plant is adapted to the hot dry conditions prevalent in desert

climes. These plants have a unique strategy in which their stomata
remain closed most of the day when water loss is highest, but can
maintain a healthy rate of photosynthesis even though CO 2 is not
supplied by gas exchange through these pores. How can this be?

These plants absorb and store CO2 at night when stomata are open.
Water loss is reduced and the acids which are used to sequester the CO 2
readily release it during the day as needed.

 
Food Transport in Plants
Mechanisms of Phloem Transport - Links

 Aphid Investigation
 Techniques to measure plant cell water and solute relations
 Biology 11 - Lecture 21 Fertilizers, Soil, Water

Flow from Source to Sink

Food, primarily sucrose is transported by the vascular tissue called phloem from a
source to a sink.

Unlike transpiration's one-way flow of water sap, food in phloem sap can be
transported in any direction needed so long as there is a source of sugar and a sink
able to use, store or remove the sugar.

The source and sink may be reversed depending on the season, or the plant's needs.
Sugar stored in roots may be mobilized to become a source of food in the early spring
when the buds of trees, the sink, need energy for growth and development of the
photosynthetic apparatus.

Phloem sap is mainly water and sucrose, but other sugars, hormones and amino acids
are also transported. The movement of such substances in the plant is called
translocation.

The Pressure Flow or Mass Flow Hypothesis

The accepted mechanism needed for the translocation of sugars from source to sink is
called the pressure flow hypothesis. (see diagram below)

As glucose is made at the source (by photosynthesis for example) it is converted to

sucrose (a dissacharide). The sugar is then moved into companion cells and into the
living phloem sieve tubes by active transport. This process of loading at the source
produces a hypertonic condition in the phloem.

Water in the adjacent xylem moves into the phloem by osmosis. As osmotic pressure
builds the phloem sap will move to areas of lower pressure.

At the sink osmotic pressure must be reduced. Again active transport is necessary to
move the sucrose out of the pholem sap and into the cells which will use the sugar --
converting it into energy, starch, or cellulose. As sugars are removed osmotic pressure
decreases and water moves out of the phloem.

The movement of sugars in the phloem begins at the source, where (a) sugars are loaded
(actively transported) into a sieve tube. Loading of the phloem sets up a water potential gradient
that facilitates the movement of water into the dense phloem sap from the neighboring xylem (b).
As hydrostatic pressure in the phloem sieve tube increases, pressure flow begins (c), and the sap
moves through the phloem. Meanwhile, at the sink (d), incoming sugars are actively transported
out of the phloem and removed as complex carbohydrates. The loss of solute produces a high
water potential in the phloem, and water passes out (e), returning eventually to the xylem.