Transport in flowering plants

Transport systems in plants

Plants may not have blood vessels and a heart, but they nevertheless have transport systems of
cells which form tubular vessels to transport molecules and ions in solution from one place to
another. The xylem tissue carries water and dissolved ions from the roots to the aerial parts of the
plant. In the tallest trees this can be over 100 metres. Phloem carries water and dissolved food
molecules from the leaves to all parts of the plant.
For more about the solvent properties of water see Soil water.

Turgor and plasmolysis

The shape of plant cells is defined by their cell wall. This is normally slightly stretched and rigid,
due to the uptake of water by osmosis. Water will move from a less concentrated solution into a
more concentrated solution through a partially permeable membrane, as there will be a diffusion
gradient from where there is more water to where there is less water.
The ability of water to move from one place to another is called water potential. Water will move
from higher to lower water potential. The water potential of a cell,, is given by:
= s + p
Where s is the solute potential, due to particles dissolved in the water in the cytoplasm (which
lowers water potential), and p is the pressure potential due to the stretching of the cell wall
(which in increases water potential).
The water potential of pure water at one atmosphere pressure is zero, so adding solutes has a
negative effect on water potential, tending to make water enter a system. Increasing the pressure
has a positive effect, tending to make water leave.
In a solution of higher water potential, cells will take up water, the cell wall will stretch and the cell
will become turgid. The pressure potential will increase until it cancels out the effect of having a
more dilute solution outside the cell. At this point the solute potential and the pressure potential are
equal and opposite, no more water enters and the cell is said to be fully turgid.
Turgid cells press against each other and provide support to the tissues in a plant.
In a solution of lower water potential, cells lose water by osmosis until the cell wall is no longer
stretched and the pressure potential becomes zero. After this further water loss causes the cell
vacuole to shrink and to pull the cell membrane away from the cell wall. Gaps appear between the
cell wall and the cell wall, and the cell is said to be plasmolysed. Plasmolysed cells do not press
against each other and tissues become soft. Leaves and stems will wilt.
It is therefore important that plants have adequate water to maintain turgidity of their cells, so they
can support the leaves to present the maximum surface area to sunlight, to promote
photosynthesis and maximum growth rates. Flowers must also be open for pollinating insects, to
promote full fertilisation for maximum seed and fruit development.

The transpiration stream

Transpiration is the evaporation of water from the surface of the mesophyll cells (especially spongy
mesophyll) in leaves. It diffuses out into the atmosphere through the stomata. The transpiration
stream is the flow of water and dissolved inorganic ions from the roots up to the leaves. As water
evaporates in the leaves, more water is pulled through the plant, in a continuous stream from the
roots up through the xylem and into the leaves, due to the cohesion of the water molecules.

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Cold, dull, damp or humid days, or soil
water shortage slow down transpiration. Figure 1
Hot, dry, windy conditions favour Guard cells in
transpiration, but also cause significant the epidermis
water loss from the plant. The guard cells of a leaf
therefore regulate the diffusion of water control the
from leaves by opening and closing the opening and
stomata to different extents, depending closing of the
stomata. This
on the water supply being taken from the
regulates loss
soil and the rate of photosynthesis. of water by
The waxy cuticle of the leaf greatly transpiration.
reduces water loss from the outside of
the leaf, except through the stomata.
The thicker the waxy layer, the less water is lost through this surface.
The stomata must be open to allow carbon dioxide to diffuse in for photosynthesis to occur. ATP
synthesised during the light reactions of photosynthesis in chloroplasts in the guard cells provides
energy for the active uptake of potassium ions.
Water passes from the neighbouring epidermal cells (which have no chloroplasts) by osmosis into
the guard cells, causing them to become turgid. The inner walls of the guard cells around the
stomata are thickened and inelastic. The outer cell walls stretch as water enters, so the cells bend
into a banana shape, opening the stomata. This can be modelled by putting sticky tape down one
side of a sausage-shaped balloon. When the balloon inflates it will expand on the side away from
the tape and bend.
There is a daily cycle. As daylight increases in the morning, photosynthesis also increases and the
stomata open to allow carbon dioxide to enter, but also transpiration to occur. In the evening light
fades, photosynthesis ceases, the potassium ions migrate out of the guard cells so that they lose
water and turgor, and the stomata close to prevent transpiration and conserve water.
Water travels from the xylem through the leaves by three routes:
 Apoplast route. Water is able to pass out of the xylem and move freely through the spaces in
the highly porous cellulose cell walls in the leaves.
 Symplast route. As water evaporates it also causes a concentration gradient between
neighbouring cells. Evaporation of water causes the cytoplasm and vacuoles of cells to
become slightly more concentrated. Water can diffuse freely from cell to cell as the cytoplasm
of neighbouring cells is connected by plasmodesmata.
 Vacuolar route. Water can also pass through cell membranes and through the tonoplasts -the
surrounding membranes of cell vacuoles. Water therefore passes from cell to cell by osmosis
down the concentration gradient from the xylem (higher water concentration, higher water
potential) to the surfaces of the mesophyll cells (lower water concentration, lower water
potential), where water is evaporating.

The xylem
The xylem of flowering plants (angiosperms) contains two types of cells that carry water. The cells
die, leaving only a woody cell wall (thickened with lignin). These cells have great mechanical
strength and are able to withstand the forces that develop inside them and they also give support
to the plant.
Tracheids are elongated cells with tapered ends that overlap. Pits in the cell walls allow movement
of water and dissolved inorganic ions from cell to cell. Xylem vessels grow end to end and lose
their end walls to form continuous tubes, so that there is no resistance to water flow in these
vessels.

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Roots
The xylem is found in a vascular
bundle in the centre of roots. Here it
provides resistance to stretching
Figure 2 Cross section
forces, thereby helping to anchor of a dicot root. Xylem
the plant in the ground. A sheath of vessels can be seen
specialised cells called the inside the ring of cells
endodermis surrounds the vascular that form the
bundle. Each cell is bound to its endodermis in the
neighbours by a strip of corky centre of the root.
material (suberin) forming the
Casparian strip. This forces all the
water that passes into the xylem in
the root to pass through selective
cell membranes that control which
inorganic ions reach the xylem.
Important ions that are transported
include nitrates, phosphates, potassium and magnesium.
Protein carrier molecules in the cell membranes use energy from ATP to actively transport
inorganic ions through the endodermis. This lowers the water potential of the cells inside the
endodermis, so water follows by osmosis.
This creates root pressure; water is forced into the xylem even if there is no transpiration. Water is
also taken up into the xylem by capillarity - adhesion of water molecules to xylem cell walls and
surface tension effects cause water to rise up the narrow tubes, especially when they are very
narrow like xylem vessels, with a diameter of the order of 0.02 mm.
As water is being removed into the xylem, a water potential gradient is set up and water will pass
into the root and through to the endodermis by the same three roots found in the leaves: apoplast,
symplast and vacuolar.
Water can only be taken up in parts of the root near the growing tips called the piliferous region.
Here the epidermis is freely permeable to water. The rate of water and inorganic ion uptake is
enhanced by greatly increasing the surface area of the root cells in contact with soil water, through
the use of the long thin extensions found in root hair cells in the piliferous regions.

Mycorrhizae
The roots of most plants are colonised by symbiotic fungi which greatly increase the surface area
available for water and mineral uptake by the plant. Phosphate uptake in particular is boosted. In
return, the fungi are able to obtain carbohydrates from the plant. These fungi are most beneficial in
nutrient poor soils. Companies such as The Royal Horticultural Society and Myccorrhizal
Applications Inc. offer products like RHS Rootgrow which boost the myccorhizae in soil and claim
significant improvements in plant growth rates and FDSAC crop yields.

Water-logging
The ability of the plant to actively uptake inorganic ions depends on having a supply of ATP for the
protein carrier molecules. Energy must be supplied by aerobic respiration in the mitochondria.
Waterlogging excludes air and oxygen from the soil, without which plant roots are unable to take in
the minerals that the plant needs. Prolonged water logging will lead to the death of roots.

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Water potential gradient
Through the active uptake of
Figure 3 Cross section
ions in the endodermis and the
through a flax stem. Key:
transpiration stream the plant is
able to maintain a water 1 Pith (parenchyma)
potential gradient from the soil 2 Protoxylem
to the atmosphere.
3 Xylem
This provides a supply of water
4 Phloem
for photosynthesis and to
maintain cell turgidity. 5 Sclerenchyma (fibres)
Photosynthesis uses carbon 6 Cortex
dioxide and water to synthesise 7 Epidermis
sugars, starch and fats.
Inorganic ions taken from the
soil are used for the manufacture of other organic molecules, including proteins, chlorophyll, ATP,
DNA and other molecules needed to sustain life.

Translocation
Food molecules that have been synthesised in the leaves have to be moved to all the other parts
of the plant in a process called translocation. Nutrients like sugars (glucose and starch are
converted to sucrose for translocation) and amino acids are moved up and down in the plant in the
phloem. Meristems (growing regions), fruits and seeds, and root storage systems in particular need
good supplies of nutrients. When necessary, molecules must be retrieved from storage and
redistributed. The place of origin of a nutrient is called its source and its destination is called the
sink.

The phloem
Phloem is found in vascular bundles, closely associated with the xylem. It is a living tissue.
Translocation requires energy. If phloem cells are killed, translocation will cease. Phloem tissue
contains sieve tube cells which lie end to end to form continuous tubes. The cellulose cells walls at
the ends of the cells are perforated to form sieve plates that allow cytoplasm to run from one cell
into the next. These cells translocate nutrients, but lose their nuclei and most of their organelles
when they mature.
Companion cells run alongside the sieve tube cells. These are connected to them by
plasmodesmata. They do not translocate nutrients, but control the activity of the sieve tube cells.
They have cytoplasm with many cell organelles.

Finding out
The mechanism of movement of nutrients is not well understood. The mass flow (or pressure flow)
hypothesis suggests that phloem is loaded with sugars by active transport, using energy from
respiration in mitochondria in companion cells. Water follows by osmosis, the subsequent pressure
driving fluid through the phloem. This hypothesis does not account for movement of nutrients both
up and down the plant in phloem. More recent suggestions include the electroosmosis (postulating
an electrical imbalance) and cytoplasmic streaming offer hypotheses, but neither provides a
complete explanation of observations.
What is electroosmosis?
What is cytoplasmic streaming?