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.

Water conducting cells of the xylem

 The two types of water-conducting cells, tracheids and vessel elements, are tubular,
elongated cells that are dead at functional maturity.
 Tracheids occur in the xylem of all vascular plants. In addition to tracheids,
most angiosperms, as well as a few gymnosperms and a few seedless
vascular plants, have vessel elements.
 When the living cellular contents of a tracheid or vessel element
disintegrate, the cell’s thickened walls remain behind, forming a nonliving
conduit through which water can flow.
 The secondary walls of tracheids and vessel elements are often interrupted
by pits, thinner regions where only primary walls are present. Water can
migrate laterally between neighboring cells through pits.
 Tracheids are long, thin cells with tapered ends. Water moves from cell to
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cell mainly through the pits, where it does not have to cross thick
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secondary walls.
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 Vessel elements are generally wider, shorter, thinner walled, and less
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tapered than tracheids. They are aligned end to end, forming long pipes
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known as vessels that in some cases are visible with the naked eye.
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 The end walls of vessel elements have perforation plates that enable water
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to flow freely through the vessels.

 The secondary walls of tracheids and vessel elements are hardened with
lignin. This hardening provides support and prevents collapse under the tension of water transport.

 The primary xylem shows various patterns of

lignin deposition as shown in the diagrams on
the right

Describe the structure of the xylem vessel in relation to their functions

 Xylem vessels are elongated single tubes formed from the longitudinal fusion of several cells without horizontal end walls and
without cytoplasm
 The structure of various xylem vessels differs in degree and nature of the longitudinal side walls that are reinforced with lignin
 Protoxylem are xylem vessels that are ringed with lignin deposits (annular vessels) or have spirals of lignin.
 Metaxylem are xylem vessels that have more lignin deposits on the side walls arranged in patterns known as pitted, reticulate and
scalariform
 The structure to function of the xylem is as described below:
The vessels are joined end to end without cross walls to allow for unimpeded water flow and water flow in a continuous column
The pits present allow for lateral flow of water where necessary such as in the case of overcoming blockages and air locks
Lignin in the side walls helps to strengthen the walls to resist compressional forces and preventing them from collapse
The lumen is narrow so that capillarity forces can act to help the water to rise in the vessels
Lignification of the cellulose walls also increases adhesion of water molecules to help capillarity

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Movement of water
 Water Potential. Osmosis can be quantified using water potential, so we can calculate which way water will move, and how fast.
Water potential (Ψ, the Greek letter psi, pronounced "sy") is simply the
effective concentration of water. It is measured in units of pressure (Pa,
or usually kPa), and the rule is that water always "falls" from a high to
a low water potential (in other words it's a bit like gravity potential or
electrical potential).
 100% pure water has Ψ = 0, which is the highest possible water potential,
so all solutions have Ψ < 0, and you cannot get Ψ > 0.
There are two factors which affect the water potential.
a). Solute concentration:

 Pure water has a water potential of 0 KPa.

 Adding solute into pure water will decrease its water potential.
 For example, a solution containing 17g of sucrose in 1dm3 of water has a water potential of -130KPa. A solution containing 35g of
sucrose in 1dm3 of water would have a water potential of -260 KPa.
 This simply means that the water molecules will have a lesser tendency to move away from a more concentrated solution (with a
low ¥).
b). Pressure on both side of the membrane:
 Consider a plant cell placed in pure water. Water enters the cell by osmosis down a water potential gradient. However, this does
not go on forever. The inward movement of water will stop when the cell becomes turgid, even though a water potential gradient
still exists.
 This is because the cell wall exerts a pressure on the water molecules and decreases its tendency (water potential) to enter the
cells.

Water Transport in Plants

 Vast amounts of water pass through plants. A large tree can use water at a rate of 1 dm³ min-. 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:
 In the roots water movement is by three pathways namely apoplast, symplast and vacuolar
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.
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The Apoplast pathway

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 consists of the cell walls between cells (90%). The cell walls are quite thick and very open, so water can easily diffuse through
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cell walls without having to cross any cell membranes by osmosis.

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 However, the apoplast pathway stops at the endodermis because of the waterproof Casparian strip, which seals the cell walls.
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At this point water has to cross the cell membrane by osmosis and enter the symplast. This allows the plant to have some
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control over the uptake of water into the

xylem.

1. Movement through the Roots

The apoplast pathway:

 Water enters the protoplast of the
endodermal cell as it is prevented from
moving completely through the cell walls by
the water proof suberin of the Casparian
strip in the endodermis. Salts may be
secreted by active transport into the xylem
from the endodermal cells to lower the water potential of the xylem. This enables water to enter the xylem from the endodermis
The symplast pathway
 The water potential of the pericycle cells becomes lower as the water molecules enter the xylem. In turn the pericycle cells draw
in water through the symplast from the neighbouring endodermal cells, causing them to also have lower water potential. The
endodermal cells then draw in water through the symplast pathway from neighbouring cortex cells. As cortex cells now have low
water potential, water is drwan in via the root hair cells
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 the cohesion-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

 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:
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Transpiration
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 Is the process of water vapour being lost from the inside of a leaf to the outside atmosphere, via the stomata (mostly), cuticle
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and lenticels.
 Transpiration rate can be indirectly measured by a potometer which directly measures rate of water absorption
 Various factors affect rate of transpiration in the ways discussed below
i. 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.
ii. Temperature. High temperature increases the rate of evaporation of water from the spongy cells, and reduces air humidity, so
transpiration increases.
iii. 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.
iv. Air movements. Wind blows away saturated air from around stomata, replacing it with drier air, so increasing the water
potential gradient and increasing transpiration.

Explain the relationship between transpiration and gaseous exchange;

 Stomata are the pores through which gaseous exchange occurs, the same way they are the pathway for transpiration
 So as the stomata open to allow gases for photosynthesis to enter, water also diffuse out
Adaptations to dry habitats shown by plants (xerophytic adaptations)

Outline how the structure of the phloem sieve tube is related to its function

 The main function of phloem is the translocation of organic substances such as

sucrose
 This translocation is made possible via sieve tube cells laid end to end with pores
measuring 2-6µm in diameter at the end walls
 The end walls contain callose (a polysaccharide) and forms a sieve plate maintaining
continuous, longitudinal strands of cytoplasm from cell to cell known as transcellular
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strands
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 The sieve plate is also important in preventing the sieve tube from bulging or
exploding outwards due to high internal pressures of the solutes
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 Because sieve tubes are living cells, this allows the maintanance of the cell surface
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membranes to prevent leakage of sucrose from the sieve tubes during translocation
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 In the event of damage of the sieve tube, callose is deposited across the sieve tube
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to block the sieve pores and seal off the tube

 The transcellular strands also provide another mechanism of translocation called
cytoplasmic streaming
 The companion cell attached to the sieve tube cell is a modified parenchyma cell with
a thin cellulose cell wall. It contains cytoplasm, a nucleus, many mitochondria, golgi
apparatus rough ER and vacuoles. They are connected to sieve tube cells via
plasmodesmata
How does the Phloem load and unload?
 Loading: Loading of sieve tubes from the cell walls requires energy
which is derived indirectly by the proton gradient.
 1. ATP and H+-carrier in the cell membrane are used to pump protons
out of the sieve tube.
 2. A proton gradient forms across the membrane with high H+
concentration on the outside of the membrane, K+ enters to keep
charge balance.
 3. Diffusion of proton back into the sieve tube, through ATP-ase, is
coupled to a carrier and powers the transport of sugar into the sieve
tube.
 ATP is important in this process because: - It increases Phloem
loading
 After loading, the sucrose is then translocated mainly via the mass
flow mechanism as postulated by Munch
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 neighboring 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
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force.
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4. In the roots the solutes are removed from the phloem by

active transport into the cells of the root.
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5. At the same time, ions are being pumped into the xylem
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from the soil by active transport, reducing the water

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potential in the xylem.

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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