Transport Systems in Plants

Plants don’t have a circulatory system like animals, but they do have a sophisticated transport system for
carrying water and dissolved solutes to different parts of the plant, often over large distances.

Stem Structure Root Structure 

 Epidermis. One cell thick. In young

 Epidermis. A single layer of cells often with long
plants the epidermis cells may secrete
extensions called root hairs, which increase the surface area
a waterproof cuticle, and in older plants
enormously. A single plant may have 1010 root hairs.
the epidermis may be absent, replaced
 Cortex. A thick layer of packing cells often containing
by bark.
stored starch.
 Cortex. Composed of various “packing”
 Endodermis. A single layer of tightly-packed cells
cells, to give young plants strength and
containing a waterproof layer called the casparian strip. This
flexibility, and are the source of plant
prevents the movement of water between the cells. 
fibres such as sisal and hemp.
 Vascular Tissue. This contains
the phloemand xylem tissue, which
grow out from the cambium. In dicot
plants (the broad-leafed plants), the
vascular tissue is arranged in vascular
bundles, with phloem on the outside  Pericycle. A layer of undifferentiated meristematic

and xylem on the inside. In older plants (growing) cells.

the xylem bundles fuse together to form

 the bulk of the stem.  Vascular Tissue. This contains xylem and phloem cells,
 Pith. The central region of a stem, used which are continuous with the stem vascular bundles. The
for food storage in young plants. It may arrangement is different, and the xylem usually forms a star
be absent in older plants (i.e. they’re shape with 2-6 arms.
hollow).

Xylem Tissue

Xylem tissue is composed of dead

cells joined together to form long
empty tubes. Different kinds of cells
form wide and narrow tubes, and the
end cells walls are either full of holes,
or are absent completely. Before death
the cells form thick cell walls
containing lignin, which is often laid
down in rings or helices, giving these
cells a very characteristic appearance
under the microscope. Lignin makes
the xylem vessels very strong, so that
they don’t collapse under pressure,
and they also make woody stems
strong.
 Phloem Tissue

Phloem tissue is composed of sieve tube cells,

which form long columns with holes in their
end walls called sieve plates. These cells are
alive, but they lose their nuclei and other
organelles, and their cytoplasm is reduced to
strands around the edge of the cells.
These cytoplasmic strands pass through the
holes in the sieve plates, so forming
continuous filaments. The centre of these
tubes is empty. Each sieve tube cell is
associated with one or more companion cells,
normal cells with nuclei and organelles. These
companion cells are connected to the sieve
tube cells by plasmodesmata, and provide
them with proteins, ATP and other nutrients.

Water Transport in Plants 

Vast amounts of water pass through plants. A large tree can use water at a rate of 1 dm³ min -1. Only 1%
of this water is used by the plant cells for photosynthesis and turgor, and the remaining 99% evaporates
from the leaves and is lost to the atmosphere. This evaporation from leaves is called transpiration.

The movement of water through a plant can be split into three sections: through the roots, stem and
leaves:

1.    Movement through the Roots 

 Water moves through the root by two paths:

 The Symplast pathway consist of the living cytoplasms of the cells in the root (10%). Water is
absorbed into the root hair cells by osmosis, since the cells have a lower water potential that the water in
the soil. Water then diffuses from the epidermis through the root to the xylem down a water potential
gradient. The cytoplasms of all the cells in the root are connected by plasmodesmata through holes in the
cell walls, so there are no further membranes to cross until the water reaches the xylem, and so no further
osmosis.
 The Apoplast pathway consists of the cell walls between cells (90%). The cell walls are quite thick
and very open, so water can easily diffuse through cell walls without having to cross any cell membranes
by osmosis. However the apoplast pathway stops at the endodermis because of the waterproof casparian
strip, which seals the cell walls. At this point water has to cross the cell membrane by osmosis and enter
the symplast. This allows the plant to have some control over the uptake of water into the xylem.

The uptake of water by osmosis actually produces a force that pushes water up the xylem. This force is
called root pressure, which can be measured by placing a manometer over a cut stem, and is of the order
of 100 kPa (about 1 atmosphere). This helps to push the water a few centimetres up short and young
stems, but is nowhere near enough pressure to force water up a long stem or a tree. Root pressure is the
cause of guttation, sometimes seen on wet mornings, when drops of water are forced out of the ends of
leaves.

 
2.    Movement through the Stem 

The xylem vessels form continuous pipes from the roots to the leaves. Water can move up through
these pipes at a rate of 8m h-1, and can reach a height of over 100m. Since the xylem vessels are
dead, open tubes, no osmosis can occur within them. The driving force for the movement is
transpiration in the leaves. This causes low pressure in the leaves, so water is sucked up the stem to
replace the lost water. The column of water in the xylem vessels is therefore under tension (a
stretching force). Fortunately water has a high tensile strength due to the tendency of water
molecules to stick together by hydrogen bonding (cohesion), so the water column does not break
under the tension force. This mechanism of pulling water up a stem is sometimes called thecohesion-
tension mechanism.

The very strong lignin walls of the xylem vessels stops them collapsing under the suction pressure, but in
fact the xylem vessels (and even whole stems and trunks) do shrink slightly during the day when
transpiration is maximum.

3.    Movement through the Leaves [back to top]

The xylem vessels ramify in the leaves to form a branching system of fine vessels called leaf veins.
Water diffuses from the xylem vessels in the veins through the adjacent cells down its water potential
 gradient. As in the roots, it uses the symplast pathway through the living cytoplasm and the apoplast
pathway through the non-living cell walls. Water evaporates from the spongy cells into the sub-
stomatal air space, and diffuses out through the stomata.

Evaporation is endothermic and is driven by solar energy, which is therefore the ultimate source of energy
for all the water movements in plants:

  

Factors affecting Transpiration [back to top]

The rate of transpiration can be measured in the lab using a potometer (“drinking meter”):

A potometer actually measures the rate of water uptake by the cut stem, not the rate of transpiration; and
these two are not always the same. During the day plants often transpire more water than they take up
(i.e. they lose water and may wilt), and during the night plants may take up more water than they transpire
(i.e. they store water and become turgid). The difference can be important for a large tree, but for a small
shoot in a potometer the difference is usually trivial and can be ignored.

The potometer can be used to investigate how various environmental factors affect the rate of
transpiration.
   Light. Light stimulates the stomata to open allowing gas exchange for photosynthesis, and as a
side effect this also increases transpiration. This is a problem for some plants as they may lose
water during the day and wilt.
 Temperature. High temperature increases the rate of evaporation of water from the spongy cells,
and reduces air humidity, so transpiration increases.
  Humidity. High humidity means a higher water potential in the air, so a lower water potential
gradient between the leaf and the air, so less evaporation.
  Air movements. Wind blows away saturated air from around stomata, replacing it with drier air,
so increasing the water potential gradient and increasing transpiration.

Many plants are able to control their stomata, and if they are losing too much water and their cells are
wilting, they can close their stomata, reducing transpiration and water loss. So long periods of light, heat,
or dry air could result in a decrease in transpiration when the stomata close.

Adaptations to dry habitats [back to top]

Plants in different habitats are adapted to cope with different problems of water availability.

Mesophytes      plants adapted to a habitat with adequate water

Xerophytes        plants adapted to a dry habitat

Halophytes        plants adapted to a salty habitat

Hydrophytes      plants adapted to a freshwater habitat

Some adaptations of xerophytes are:

Adaptation How it works Example

thick cuticle stops uncontrolled evaporation most dicots
through leaf cells
small leaf surface area less area for evaporation conifer needles, cactus spines
low stomata density fewer gaps in leaves  
stomata on lower surface of more humid air on lower surface, so most dicots
leaf only less evaporation
shedding leaves in dry/cold reduce water loss at certain times of deciduous plants
season year
sunken stomata maintains humid air around stomata marram grass, pine
stomatal hairs maintains humid air around stomata marram grass, couch grass
folded leaves maintains humid air around stomata marram grass,
succulent leaves and stem stores water cacti
extensive roots maximise water uptake cacti

Mineral Ion transport in Plants [back to top]

Ions are absorbed from the soil by both passive and active transport. Specific ion pumps in the
membranes of root hair cells pump ions from the soil into the cytoplasms of the epidermis cells. Two lines
of evidence indicate that active transport is being used:

 The concentrations of ions inside root cells are up to 100 time greater than in the soil, so they are
being transported up their concentration gradient.
  If respiratory inhibitors such as cyanide are applied to living roots, ion uptake is greatly reduced,
since there is no ATP being made to drive the membrane pumps. Any remaining uptake must be
passive.

The active uptake of ions is partly responsible for the water potential gradient in roots, and therefore for
the uptake of water by osmosis.

Ions diffuse down their concentration gradient from the epidermis to the xylem. They travel up the xylem
by mass flow as the water is pulled up the stem (in other words they are simply carried up in the flow of
the xylem solution). In the leaves they are selectively absorbed into the surrounding cells by membrane
pumps.

Solute Transport in Plants [back to top]

The phloem contains a very concentrated solution of dissolved solutes, mainly sucrose, but also  other
sugars, amino acids, and other metabolites. This solution is called the sap, and the transport of solutes in
the phloem is called translocation.
Unlike the water in the xylem, the contents of the phloem can move both up or down a plant stem, often
simultaneously. It helps to identify where the sugar is being transported from (the source), and where to
(the sink).

 During the summer sugar is mostly transported from the leaves, where it is made by
photosynthesis (the source) to the roots, where it is stored (the sink).
 During the spring, sugar is often transported from the underground root store (the source) to the
growing leaf buds (the sink).
 Flowers and young buds are not photosynthetic, so sugars can also be transported from leaves
or roots (the source) to flowers or buds (sinks).

Surprisingly, the exact mechanism of sugar transport in the phloem is not known, but it is certainly far too
fast to be simple diffusion. The main mechanism is thought to be the mass flow of fluid up the xylem and
down the phloem, carrying dissolved solutes with it. Plants don’t have hearts, so the mass flow is driven
by a combination of active transport (energy from ATP) and evaporation (energy from the sun). This is
called the mass flow theory, and it works like this:

1.  Sucrose produced by photosynthesis is actively pumped into the phloem vessels by the
companion cells.
2. This decreases the water potential in the leaf phloem, so water diffuses from the neighbouring
xylem vessels by osmosis.
3. This is increases the hydrostatic pressure in the phloem, so water and dissolved solutes are
forced downwards to relieve the pressure. This is mass flow: the flow of water together with its
dissolved solutes due to a force.
4. In the roots the solutes are removed from the phloem by active transport into the cells of the root.
 5. At the same time, ions are being pumped into the xylem from the soil by active transport,
reducing the water potential in the xylem.
6. The xylem now has a lower water potential than the phloem, so water diffuses by osmosis from
the phloem to the xylem.
7. Water and its dissolved ions are pulled up the xylem by tension from the leaves. This is also
mass flow.

This mass-flow certainly occurs, and it explains the fast speed of solute translocation. However there
must be additional processes, since mass flow does not explain how different solutes can move at
different speeds or even in different directions in the phloem. One significant process is cytoplasmic
streaming: the active transport of molecules and small organelles around cells on the cytoskeleton.

Translocation Experiments [back to top]

1.                  Puncture Experiments

If the phloem is punctured with a hollow tube then the sap oozes
out, showing that there is high pressure (compression) inside the
phloem (this is how maple syrup is tapped). If the xylem is
punctured then air is sucked in, showing that there is low
pressure (tension) inside the xylem. This illustrates the main
difference between transport in xylem and phloem: Water
is pulled up in the xylem, sap is pushed down in the phloem.

2.                  Ringing Experiments

Since the phloem vessels are outside the xylem vessels,

they can be selectively removed by cutting a ring in a stem
just deep enough to cut the phloem but not the xylem. After
a week there is a swelling above the ring, reduced growth
below the ring and the leaves are unaffected. This was
early evidence that sugars were transported downwards in
the phloem.
 

3.                  Radioactive Tracer Experiments

Radioactive isotopes can be used trace precisely where different compounds are being transported from
and to, as well as measuring the rate of transport. The radioactivity can be traced using photographic film
(an autoradiograph) or a GM tube. This techniques can be used to trace sugars, ions or even water.

In a typical experiment a plant is grown in the lab and one leaf is exposed for a short time to carbon
dioxide containing the radioactive isotope 14C. This 14CO2 will be taken up by photosynthesis and the 14C
incorporated into glucose and then sucrose. The plant is then frozen in liquid nitrogen to kill and fix it
quickly, and placed onto photographic film in the dark. The resulting autoradiograph shows the location of
compounds containing 14C.

This experiment shows that organic compounds (presumably sugars) are transported downwards
from the leaf to the roots. More sophisticated experiments using fluorescently labelled compounds
can locate the compound specifically to the phloem cells.

4.                  Aphid Stylet Experiments

Aphids, such as greenfly, have specialised mouthparts called stylets,

which they use to penetrate phloem tubes and sup of the sugary sap
therein. If the aphids are anaesthetised with carbon dioxide and cut off,
the stylet remains in the phloem so pure phloem sap can be collected
through the stylet for analysis. This surprising technique is more accurate
than a human with a syringe and the aphid’s enzymes ensure that the
 stylet doesn’t get blocked.