7.

Transport in plants
Dicotyledonous (dicots) plants

 Dicotyledonous (dicots) plants have:

o Seeds that contain two cotyledons (seed leaves)
o Network of veins
o Leaves that typically have broad blades (leaf surface) and petioles (stalks)
o Tap root with lateral branches
 Herbaceous dicots have a relatively short life cycle (one growing season) and non-woody
tissue

Transport systems

 Plants need transport systems to meet their metabolic demands (glucose, hormones,
mineral ions are required for various processes within plants), to efficiently move
substances up and down and to compensate for their relatively small SA:V ratio (generally
plants cannot rely on diffusion alone)
 Plants have a vascular system which involves a network of vessels (vascular tissue) running
through the leaves, stem and roots. These three parts are the main organs involved in
transport
 The vascular system is comprised of two distinct types:
o Xylem (transports water and mineral ions from the roots to the rest of the plant)
o Phloem (transports substances from the source (eg. leaf) to the sink (eg.root))
 The xylem and phloem are arranged together in vascular bundles
 The bundles are laid out differently in the leaves, stem and roots

Tissue plan diagrams of a dicotyledonous leaf, stem and root

Xylem & Phloem Distribution

In roots and stem of a dicotyledonous plant, the xylem tissue is found on the inside –
however, in leaves, xylem is found above phloem tissue.

Xylem
 The functions of xylem tissue in a plant are:
o Vascular tissue that carries dissolved minerals and water up the plant
o Structural support
o Food storage
 Xylem tissue is found, along with phloem tissue and other tissues, in vascular bundles
o In the roots the vascular bundle is found in the centre and the centre core of this
is xylem tissue. This helps the roots withstand the pulling strains they are subjected to
as the plant transports water upwards and grows
o In the stems the vascular bundles are located around the outside and the xylem
tissue is found on the inside (closest to the centre of the stem) to help support the
plant
o In the leaves the vascular bundles form the midrib and veins and therefore spread
from the centre of the leaf in a parallel line. The xylem tissue is found on the upper
side of the bundles (closest to the upper epidermis)

Phloem
 The function of phloem tissue in a plant is to transport organic compounds, particularly
sucrose, from the source (eg. leaf) to the sink (eg. roots). The transport of these compounds
can occur up and down the plant
 Phloem is a complex tissue also made up of various cell types; its bulk is made up of sieve
tube elements which are the main conducting cells and the companion cells
 Other cell types of phloem tissue also include parenchyma for storage and strengthening
fibres
o In the roots the vascular bundle is found in the centre and on the edges of the centre
core is the phloem tissue
o In the stems, the vascular bundles are located around the outside and the phloem
tissue is found on the outside (closest to the epidermis)
o In the leaves, the vascular bundles form the midrib and veins and therefore spread
from the centre of the leaf in a parallel line. The phloem tissue is found on the lower
side of the bundles (closest to the lower epidermis)
Xylem Vessel Elements: Structure & Function

 The functions of xylem tissue in a plant are:

o Vascular tissue that transports dissolved minerals and water around the plant
o Structural support
o Food storage
 Xylem tissue is made up of four cell types that function together:

1. Tracheids (long, narrow tapered cells with pits)

2. Vessel elements (large with thickened cell walls and no end plates when mature)
3. Xylem parenchyma
4. Sclerenchyma cells (fibres and sclereids)

 Most of the xylem tissue is made up of tracheids and vessel elements, which are both types
of water-conducting cell

Images of xylem vessel elements, (a) photomicrograph in longitudinal section (lignin is stained
red), (b) scanning electron micrograph in transverse section and (c) microscope image in
transverse section and drawing (lignin is stained red)
 Relating structure & function in xylem vessel elements table

You must be able to recognise the xylem vessel elements in images so look for the thicker cell walls
and the larger diameter. You also need to know the difference between xylem and phloem tissue.

Phloem Sieve Tube Elements & Companion Cells: Structure & Function

 The function of phloem tissue in a plant is to:

o Transport organic compounds (assimilates), particularly sucrose, from the source
(eg. leaf) to the sink (eg. roots). The transport of these compounds can
occur up and down the plant
 The organic compounds are dissolved in water to form sap
 Phloem is a complex tissue made up of various cell types; its bulk is made up of sieve tube
elements which are the main conducting cells and companion cells
 Other cell types of phloem tissue also include parenchyma for storage and strengthening
fibres
 Mature phloem tissue contains living cells, unlike xylem tissue
Structure of phloem tissue. (a) Microscope slide image and drawing of a sieve tube element and
companion cell in transverse section (TS), (b) photomicrograph image and drawing of a sieve
tube element and companion cell in longitudinal section (LS).

Structure of phloem tissue. (c) Transmission electron micrograph image and drawing of a sieve
tube element and companion cell in transverse section (TS)
Sieve tube elements

 Sieve tube elements line up end to end to form a continuous tube

Phloem sieve tube elements structure & function table

Companion cells
 Each sieve tube element has a companion cell associated with it as companion cells control
the metabolism of their associated sieve tube member
 They also play a role in loading and unloading of sugars into the phloem

Phloem companion cells structure & function table

 Comparison of xylem & phloem tissue table

Understand the difference between sieve tube elements and companion cells, and how they are
different to xylem tissue. Remember that mature xylem tissue is dead, so there is no evidence of
organelles, and they have lignified cell walls, whereas sieve tube elements have no lignin, do have
sieve plates, and their companion cells contain nuclei and dense cytoplasm.

Water & Mineral Ion Transport: Pathways & Mechanisms

 Within a plant mineral ions and organic compounds (eg. sucrose) are transported by
being dissolved in water. The dissolved mineral ions are transported in the xylem tissue
and the dissolved organic compounds are transported in the phloem tissue
 The plant roots are responsible for the uptake of water and mineral ions and can have root
hairs to increase the surface area for absorption of the substances
 The uptake of water is a passive process and occurs by osmosis (the diffusion of water from
a higher (less negative) water potential to a lower (more negative) water potential
 The uptake of minerals can be passive or active and occurs by diffusion or active
transport respectively
 Plants must take in a constant supply of water and dissolved minerals to compensate for the
continuous loss of water via transpiration in the leaves, and so that they can photosynthesise
and produce proteins

There are two pathways that water (and the dissolved solutes) can take to move across the cortex
(and molecules can change between routes at any time):

o Apoplastic
o Symplastic
Apoplast pathway

 Most water travels via the apoplastic pathway (when transpiration rates are high), which is
the series of spaces running through the cellulose cell walls, dead cells, and the hollow tubes
of the xylem
 The water moves by diffusion (as it is not crossing a partially permeable membrane)
 The water can move from cell wall to cell wall directly or through the intercellular spaces
 The movement of water through the apoplastic pathway occurs more rapidly than the
symplastic pathway
 When the water reaches the endodermis the presence of a thick, waterproof, waxy band
of suberin within the cell wall blocks the apoplastic pathway
 This band is called the Casparian strip and forms an impassable barrier for the water
 When the water and dissolved minerals reach the Casparian strip they must take the
symplastic pathway. The presence of this strip is not fully understood but it is thought that
this may help the plant control which mineral ions reach the xylem and generate root
pressure
 As the plant ages the Casparian strip thickens (as more suberin is deposited) except in cells
called the passage cells, allowing for further control of the mineral ions

Symplast pathway

 A smaller amount of water travels via the symplastic pathway, which is

the cytoplasm and plasmodesmata or vacuole of the cells
 The water moves by osmosis into the cell (across the partially permeable cell surface
membrane), possibly into the vacuole (through the tonoplast by osmosis) and between cells
through the plasmodesmata
 The movement of water in the symplastic pathway is slower than the apoplastic pathway

Water (and any dissolved substances) can travel via the apoplastic or symplastic pathways. As
the plant ages the apoplastic pathway can be blocked by the presence of the casparian strip
helping the plant control which mineral ions can move into the xylem vessels

Remember water moves through the apoplastic and symplastic pathways in the leaves as well as the
roots. Water does not move by osmosis in the apoplastic pathway as the molecules are in the cell wall
which is freely permeable.
Transpiration:

 The movement of water through a plants xylem is largely due to the evaporation of water
vapour from the leaves and the cohesive and adhesive properties exhibited by water
molecules
 It is the gradient in water potential that is the driving force permitting the movement of water
from the soil (high water potential), to the atmosphere (low water potential), via the
plant’s cells
 Plants are constantly taking water in at their roots and losing water via the stomata (in the
leaves)
 Around 99% of the water absorbed is lost through evaporation from the plant’s stem and its
leaves in a process called transpiration
 Transpiration refers to the loss of water vapour from a plant to its environment
by diffusion and the transpiration stream refers to the movement of water from the roots to
the leaves
 The advantage of transpiration is that:
o It provides a means of cooling the plant via evaporative cooling
o The transpiration stream is helpful in the uptake of mineral ions
o The turgor pressure of the cells (due to the presence of water as it moves up the plant)
provides support to leaves (enabling an increased surface area of the leaf blade) and
the stem of non-woody plants

The loss of water vapour from the leaves of plants (transpiration) results in a lower water
potential creating a concentration gradient between the roots and leaves causing water to move
upwards
Movement of water through leaves

 Certain environmental conditions (eg. low humidity, high temperatures) can cause a water
potential gradient between the air inside the leaves (higher water potential) and the air
outside (lower water potential) which results in water vapour diffusing out of the leaves
through the stomata (transpiration)
 The water vapour lost by transpiration lowers the water potential in the air spaces
surrounding the mesophyll cells
 The water within the mesophyll cell walls evaporates into these air spaces resulting in a
transpiration pull
 This transpiration pull results in water moving through the mesophyll cell wall (apoplastic
pathway) or out of the mesophyll cytoplasm (symplastic pathway) into the cell wall
 The pull from the water moving through the mesophyll cells results in water leaving the
xylem vessels through pits (non-lignified areas), which then causes water to move up the
xylem vessels (due to the cohesive and adhesive properties of the water). This movement is
called transpiration stream
o When rates of transpiration are high the walls of the xylem are pulled inwards by the
faster flow of water

The role of the stomata

  Transpiration is mainly controlled by the pairs of guard cells that surround stomata (plural,
stoma is singular)
 Guard cells open the stomata when they are turgid and close the stomata when they lose
water
 When the stomata are open there is a greater rate of transpiration and of gaseous exchange
 When the stomata close transpiration and gaseous exchange decrease
 As stomata allow gaseous exchange (CO2 in and O2 out) they are generally open during the day

Remember that water vapour diffuses through the stomata and water evaporates from the
mesophyll cells into the air spaces in the leaf. Transpiration and transpiration pull/stream are
different – transpiration is the loss of water vapour from the leaves or stem, whereas transpiration
pull/stream is the movement of water through the xylem tissue and mesophyll cells.

Water & the Transpiration Pull

 Water has unique properties

o it is polar
o hydrogen bonds form between the water molecules
 Water moves from the roots to the leaves because of a difference in the water potential
gradient between the top and bottom of the plant. This gradient is created because of different
events occurring within the plant and due to the properties of water
 In the leaves, water evaporates from the mesophyll cells resulting in water (and any dissolved
solutes) being pulled from the xylem vessels (transpiration pull) into the mesophyll cells
 The water that is pulled into the mesophyll cells moves across them passively (either via the
apoplastic – diffusion or symplastic – osmosis, pathways) lowering the hydrostatic pressure
within the xylem vessels and creating a tension on these vessels
 Xylem vessels have lignified walls to prevent them from collapsing due to the pressure
differences being created from the mass flow (all the water molecules and any dissolved
solutes move together) of water upwards
 The mass flow is helped by the polar nature of water and the hydrogen bonds (H-bonds) that
form between water molecules which results in cohesion between water molecules
and adhesion between the cellulose in the cell walls and the water molecules
 So due to the evaporation of water from the mesophyll cells in the leaves a tension is created
in the xylem tissue which is transmitted all the way down the plant because of the
cohesiveness of water molecules. The cohesive force results in a continuous column of water
with high tensile strength (it is unlikely to break) and the adhesive force stops the water
column from pulling away from the walls of the xylem vessels so water is pulled up the xylem
tissue from the roots to replace what was lost in the leaves. This mechanism is called
the cohesion-tension theory

The transpiration stream

 The pathway of the water from the soil through the roots up the xylem tissue to the leaves is
the transpiration stream
 Plants aid the movement of water upwards by raising the water pressure in the roots (root
pressure)
o This is raised by actively secreting solutes (eg. mineral ions) into the xylem vessels in
the root which lowers the water potential within the xylem
 o This results in water from the surrounding cells being drawn into the xylem (by
osmosis) thus increasing the water pressure (root pressure)
 Root pressure helps move water into the xylem vessels in the roots however the volume
moved does not contribute greatly to the mass flow of water to the leaves in the transpiration
stream

The transpiration stream – the mass flow of water from the roots to the leaves. This is
possible due to the cohesion-tension theory
When answering questions about transpiration it is important to include the following keywords:

 Water potential gradient (between leaves and roots),

 Diffusion (water vapour through the stomata)
 Transpiration pull (evaporation of water from the mesophyll cells pulls other water
molecules from the xylem tissue)
 Cohesion (between water molecules)
 Adhesion (between water molecules and cellulose within the cell walls)
 Cohesion-tension theory (tension present in xylem vessels causes a continuous column of
water and is due to cohesive and adhesive forces)
 Osmosis (water via the apoplastic or symplastic pathways in the roots and leaves)

Xerophytic Plant Leaf Adaptations

 Xerophytes (from the Greek xero for ‘dry’) are plants that are adapted to dry and arid
conditions
 Xerophytes have physiological and structural (xeromorphic) adaptations to maximise water
conservation

Photomicrograph and annotated drawing showing the xeromorphic features of a leaf of

Ammophilia arenaria (Marram grass)
Xeromorphic features table

Remember not all leaves will have every feature listed above so if you are looking at an unfamiliar
image consider whether the adaptations you can see will help reduce water being lost from the leaf.
Movement in the Phloem

 Although translocation could refer to the transport of substances in the xylem and phloem, as
it means ‘moving from one place to another,’ it is more commonly connected with the
transport of assimilates in the phloem tissue
 Thus translocation within phloem tissue can be defined as the transport of assimilates
from source to sink and requires the input of metabolic energy (ATP)
 The liquid that is being transported (found within phloem sieve tubes) is called phloem sap
 This phloem sap consists not only of sugars (mainly sucrose) but also of water and other
dissolved substances such as amino acids, hormones and minerals
 The source of the assimilates could be:
o Green leaves and green stem (photosynthesis produces glucose which is transported
as sucrose, as sucrose has less of an osmotic effect than glucose)
o Storage organs eg. tubers and tap roots (unloading their stored substances at the
beginning of a growth period)
o Food stores in seeds (which are germinating)
 The sinks (where the assimilates are required) could be:
o Meristems (apical or lateral) that are actively dividing
o Roots that are growing and / or actively absorbing mineral ions
o Any part of the plant where the assimilates are being stored (eg. developing seeds,
fruits or storage organs)

 The loading and unloading of the sucrose from the source to the phloem, and from the phloem
to the sink is an active process
 It can be slowed down or even stopped at high temperatures or by respiratory inhibitors

Translocation of assimilates is not fully understood yet by scientists. The understanding they
do have has come from studies such as:

o On plants whose sap does ‘clot’ so that it is still possible to collect and study the sap
(eg. castor oil plants)
o Using aphids to collect the sap – after the aphid inserts its stylet (tubular mouthpart)
scientists remove the aphids head and collect the sap that continues to flow
o Using radioactively labelled metabolites (eg. Carbon-14 labelled sugars) which can be
traced during translocation
o Advances in microscopes enabling the adaptations of companion cells to be seen
o Observations about the importance of mitochondria to the process of translocation

Assimilates can move upwards or downwards in the phloem sieve tubes as they move from source to
sink.

The Sucrose Loading Mechanism

 Assimilates such as sucrose are transported from source to sink through the phloem sieve
tubes
 Carbohydrates are generally transported in plants in the form of sucrose because:
o It allows for efficient energy transfer and increased energy storage (sucrose is a
disaccharide and therefore contains more energy)
o It is less reactive than glucose as it is a non-reducing sugar and therefore no
intermediate reactions occur as it is being transported
Loading of assimilates (eg. sucrose)

 The molecules may move by the:

o symplastic pathway (through the cytoplasm and plasmodesmata) which is a passive
process as the sucrose molecules move by diffusion
o apoplastic pathway (through the cell walls) which is an active process
 If the sucrose molecules are taking the apoplastic pathway then modified companion
cells (called transfer cells) pump hydrogen ions out of the cytoplasm via a proton
pump and into their cell walls. This is an active process and therefore requires ATP as an
energy source
 The large concentration of hydrogen ions in the cell wall of the companion cell results in the
hydrogen ions moving down the concentration gradient back to the cytoplasm of the
companion cell
 The hydrogen ions move through a cotransporter protein. While transporting the hydrogen
ions this protein also carries sucrose molecules into the companion cell against the
concentration gradient for sucrose
 The sucrose molecules then move into the sieve tubes via the plasmodesmata from the
companion cells
 Companion cells have infoldings in their cell surface membrane to increase the available
surface area for the active transport of solutes and many mitochondria to provide the energy
for the proton pump
 This mechanism permits some plants to build up the sucrose in the phloem to up to three
times the concentration of that in the mesophyll
Unloading of assimilates (eg. sucrose)

 The unloading of the assimilates (eg. sucrose) occurs at the sinks

 Scientists believe that the unloading of sucrose is similar to the loading of sucrose, with the
sucrose being actively transported out of the companion cells and then moving out of the
phloem tissue via apoplastic or symplastic pathways
 To maintain a concentration gradient in the sink tissue, sucrose is converted into other
molecules. This is a metabolic reaction so requires enzymes (eg. invertase which hydrolyses
sucrose into glucose and fructose)

Phloem: Mass Flow

 The Mass Flow Hypothesis was the model initially used to explain the movement of
assimilates in the phloem tissue
 The mass flow hypothesis was modelled by Ernst Münch. His simple model consisted of:
o Two partially permeable membranes containing solutions with different
concentrations of ions (one dilute the other concentrated)
o These two membranes were placed into two chambers containing water and were
connected via a passageway
o The two membranes were joined via a tube
o As the membranes were surrounded by water, the water moved by osmosis across the
membrane containing the more concentrated solution which forced the solution
towards the membrane containing the more dilute solution (where water was being
forced out of due to hydrostatic pressure)
 Scientists now support a modified version of this hypothesis – the pressure flow gradient

An illustration of Münch’s model for mass flow in phloem tissue

Pressure (hydrostatic) flow gradient

 Phloem sap (containing sucrose and other organic solutes) moves by mass flow up and down
the plant
 Carbohydrates are generally transported in plants in the form of sucrose because:
o It allows for efficient energy transfer and increased energy storage (sucrose is a
disaccharide and therefore contains more energy)
o It is less reactive than glucose as it is a non-reducing sugar and therefore no
intermediate reactions occur as it is being transported
 The advantage of mass flow is that it moves the organic solutes faster than diffusion
 In xylem tissue the pressure difference that causes mass flow occurs because of a water
potential gradient between the soil and leaf (this requires no energy input by the plant)
 However in phloem tissue energy is required to create pressure differences for the mass
flow of the organic solutes
 The pressure difference is generated by actively loading sucrose into the sieve elements at
the source which lowers the water potential in the sap
 This results in water moving into the sieve elements as it travels down the water potential
gradient by osmosis
 The presence of water within the sieve elements increases the hydrostatic or
turgor pressure at the source and as solutes (eg. sucrose) are removed / unloaded from the
sieve elements causing water to follow by osmosis at the sink (creating a low hydrostatic
pressure), a hydrostatic pressure gradient occurs
 The pressure difference between the source and the sink results in the mass flow of water
from the high hydrostatic pressure area to the low hydrostatic pressure area
 The mass flow of organic solutes within the phloem tissue occurs above and below the
sources. Therefore sap flows upwards and downwards within a plant

Drawing tissue plan diagrams: You need to

 Read the instructions carefully

 Draw a large diagram
 Use a sharp pencil and do not shade (including the nucleus)
 Use clear, continuous lines
 When using an eye-piece graticule, use it to ensure you have correct proportions

 If drawing from a low-power image:

o Do not draw individual cells
o Read the question carefully as you may only have to draw a portion of the image
o Include the magnification on the drawing

 If drawing from a high-power image:

o Draw only a few of the required cells
o Draw the cell wall of the plant cells
o Include the magnification on the drawing

 When labelling, remember:

o Use a ruler for label lines (and scale line if appropriate)
o Label-lines should stop exactly at the structure (do not use arrows)
o Don’t cross label-lines over each other
o Label all tissues and relevant structures (those requested)