A1 BIOLOGY 3

Unit 3: Organisms Exchange

Substances with their Environment:

Transport in Plants
3.4.2 Mass Transports in Plants
Xylem as the tissue that transports water in the stem and leaves of plants. The cohesion-tension
theory of water transport in the xylem.
Phloem as the tissue that transports organic substances in plants.
The mass flow hypothesis for the mechanism of translocation in plants. The use of tracers and
ringing experiments to investigate transport in plants.
Students should be able to:
 recognise correlations and causal relationships
 interpret evidence from tracer and ringing experiments and to evaluate the evidence for and
against the mass flow hypothesis.

3.2. Gas Exchange

Structural and functional compromises between the opposing needs for efficient gas exchange and
the limitation of water loss shown by terrestrial insects and xerophytic plants.

Notes 11
Gas exchange and water loss in the leaf
The spongy mesophyll layer of the leaf, with its large air spaces and thin-walled cells is the principal
gas exchange surface within the leaf. This is in close contact with numerous pores or stomata, across
which gases enter and leave the leaf via diffusion down steep concentration gradients. Carbon
dioxide will be low in the leaf by day as it is used in photosynthesis. The reverse is true for oxygen. At
night, oxygen concentration will be low in the leaf, as it is used by cells for respiration. Reverse for
carbon dioxide. The cells of the spongy mesophyll layer are loosely packed, creating numerous air
spaces that provide a large surface area for gas exchange.

waxy cuticle
upper epidermis

spongy mesophyll

stoma

A short diffusion pathway is ensured by the spongy cells having thin cell walls and being in direct
contact with the air, as well as the thinness of the leaf itself.
a. Name the process that occurs in the leaf cells throughout both the day and night and the
gases it uses and produces.
respiration uses oxygen and produces carbon dioxide
b. Name the process that only occurs in leaves during daylight hours and the gases it uses and
produces.
photosynthesis uses carbon dioxide and produces oxygen
c. How does the leaf minimise water loss while still maintaining effective diffusion?
stomata mainly on underside only. Thick(er) waxy cuticle on upper epidermis.
Action of guard cells can close stomata / reduce stomatal diameter e.g. when temps too high

1
Mass transport in plants
Flowering plants have two distinct mass transport systems, both of which consist of tubes or
vessels. Mass transport is the movement of materials over large distances, due to pressure
differences.
Phloem tissue is concerned with the transport of organic substances such as sugars and other
soluble products of photosynthesis from the leaves to other parts of the plant. Xylem tissue is
concerned with the transport of water and dissolved minerals, from the soil, through the roots, the
stem and to the leaves. We will firstly focus on the uptake and transport of water through the xylem.

Transpiration

Transpiration is the process of water movement through a plant and its evaporation from leaves. The
continuous columns of water that move from the soil through the roots, stems and leaves to the air
are known as the transpiration stream.

Transpiration is a consequence of gas exchange in leaves. When the stomata

are open to allow the exchange of oxygen and carbon dioxide, water will
inevitably be lost. Plants only absorb about 1% of the water entering their
roots. This is used in photosynthesis and in maintaining turgid cells. The
remaining 99% evaporates from the leaves and is lost from the plant.
Transpiration rates in large trees can be as high as 1 dm3 min-1.
The tallest known living tree is a coast redwood (Sequoia sempervirens) is
found in California and is 115.6 m (379 feet) high.

Any theory of how transpiration works must be able to explain how water can
be moved from the soil to the top of such large trees against gravity.

Movement of water through the leaf

1. When stomata are open, water 2. mesophyll cells
3. xylem
vapour diffuses from the air spaces
in the leaf, out through the stomata,
down a water potential gradient. This
loss of water is called transpiration.

2. To replace this, water evaporates

from the walls of mesophyll cells into guard cell
the air spaces forming water vapour,
1. water diffuses out of
which builds up in the air spaces.
stoma

3. The water in the mesophyll cells is

replaced by water from the xylem
vessels in the leaf which are
continuous with xylem in the stem.

2
Movement of water from roots to leaves
Water is carried in xylem tissue. This is a series of continuous tubes called xylem vessels. These are
modified dead cells arranged end-to-end. The diagram below shows the structure of xylem vessels.
Features of xylem vessels
Structural function
feature

Their cell walls lignin strengthens the xylem walls

contain lignin. against the tension within them,
and makes them waterproof.

The lignified this leaves a hollow lumen with no

vessel walls cytoplasm that offers little
cause the cell resistance to the mass flow of
contents to die. water and minerals.

The walls of If a vessel becomes blocked or

xylem vessels damaged, the water can be
contain tiny diverted laterally, so the upwards
holes called movement of water can continue in
pits an adjacent vessel.

The vessels they form a continuous column for

also lose their water movement from root to
end walls. leaves

Q.3. Suggest a feature of lignin other than strength that would be useful in ensuring that water was
carried up the plant.

Waterproof;

Q.4. Suggest one advantage of xylem vessels being dead cells in order to carry out their function
effectively.

Living cells have cell membranes and cytoplasm. Water movement would be slower if it had to
pass through these;

The thickening of the cell wall in xylem vessels is often spiral (see right).

Q.5. Suggest two advantages to the plant of having this arrangement rather than
continuous thickening.

Uses less material is therefore less wasteful;

Uses less material therefore allows xylem to be flexible/have lower mass;

3
 The Cohesion Tension Theory

This is the currently accepted theory of how

water moves up the plant from root to leaves.
The idea here is that the evaporation of water
from the leaves pulls up more water from the
xylem in the stem. To understand the theory it
is important to remember that water molecules
are polar and so very many weak hydrogen
bonds form between them making them stick
together. This is called cohesion. Water
molecules also form hydrogen bonds with the
walls of the xylem. This is called adhesion.

Key Points
 As transpiration occurs through open stomata, water is evaporated from the cell walls of
mesophyll cells of the leaf and is replaced by water from the xylem due to the cohesive forces
between water molecules.
 This creates low pressure at the top of the xylem in the stem.
 Water in the xylem is under tension and is pulled up towards the leaves.
 Continuous columns of water are maintained due to cohesion between water molecules
 There is also adhesion of water molecules to the walls of the xylem vessels. This creates an
inward pull on the vessel walls as the water is pulled up, causing the xylem vessels to
decrease in diameter.

This process is entirely passive as no ATP (metabolic energy) is required for this to occur. The
energy is needed to drive this process is the heat required to evaporate water from the leaves. This
theory is the best current explanation of how water moves up the stems of plants and can account for
the movement of water even to the top of very tall trees.
Evidence
 Tension has been measured in xylem as plants transpire.
 If a column of water in the xylem is broken, air bubbles in the xylem form and this stops any
further upward movement of water in that xylem vessel as the air bubbles prevent cohesion.
 Respiratory inhibitors, such as cyanide or a lack of oxygen, do not inhibit this process.
 The diameter of trees decrease when they are transpiring, (tension pulls xylem walls in), and
more so when temperatures and light intensities are higher. This can be measured using a
dendrometer.

4
When a xylem vessel is broken, water does not leak out, instead air is drawn in.
Q.7. Explain how this observation provides evidence in support of the cohesion-tension theory.

The fact that air is drawn into the xylem when it is broken means that the water in the xylem must
be under tension (pulled from the top) rather than under pressure (pushed from the bottom) as
predicted by the cohesion-tension theory.

A dendrometer A dendrometer trace

Q.8. Why is the trunk diameter lowest at noon?

Stomata are open, transpiration rates are high, increased tension, so water column pulled up xylem
faster, so walls of xylem pulled in and xylem vessels decrease in diameter

Q.9. Why is the trunk diameter highest at midnight?

Stomata are closed, transpiration rates are low, little tension so little inward pull on walls of xylem
vessels

While studying transpiration, a scientist

used a dendrometer to record the small
daily changes in the diameter of a tree
trunk at two different heights (2 and 3
metres) above the ground at the same
time. The diameter decreased in the
daytime.

Q.10. Explain why this decrease

happened first at the higher location.

This decrease happens earlier at 3 metres than at 2 metres as the transpiration pull occurs from
the top of the plant where water evaporates from leaves

5
The route water takes from root to leaf

6
The route water can take from xylem to stomata across the leaf.

7
8
Limiting water loss in plants
Xerophytic adaptations

To reduce the inevitable loss of water by transpiration, leaves are covered in a waxy waterproof
covering called the cuticle. If water loss is high plants can close their stomata. In habitats with a
restricted supply of water, certain plants have additional adaptations to further reduce water loss
through transpiration. These plants are known as xerophytes and are found in any region where
water is scarce, for example deserts.

Xerophytes have structural adaptations, which allow them to survive in dry conditions by either
reducing transpiration loss or by storing water.

Marram grass colonises sand and is the main species found on sand dunes. Marram grass is a good
example of a xerophyte as it shows a number of adaptations to reduce water loss from leaves.

Transverse section through leaf of marram grass

leaf rolled up

thick waxy cuticle

trapped humid air

with high water
potential

hairs on lower epidermis of leaf

stomata sunken in pits

9
Complete the table below showing how the xerophytic adaptations of marram grass reduce water
loss. Try to relate these adaptations to reducing the water potential gradient or increasing the
length of the diffusion pathway and hence the decreasing the diffusion rate of water from the
leaf.

Adaptation How the adaptation helps limit water loss

trapped humid air with high water Water vapour is trapped within the rolled leaf, this
potential reduces the water potential gradient between air
spaces inside leaf and the atmosphere. A lower rate
of diffusion from the stomata occurs.

hairs on lower epidermis of leaf Water vapour is trapped between the hairs so
reducing the water potential gradient between air
spaces inside leaf and the atmosphere. A lower rate
of diffusion and less transpiration occurs

stomata sunken in pits Water vapour is held above the stomatal pore so
reducing the water potential gradient between air
spaces inside leaf and the atmosphere. A lower rate
of diffusion and less transpiration occurs

thick waxy cuticle
The cuticle is waxy so reduces water loss (from the
epidermis). Greater thickness increases the length
of the diffusion pathway for water to reach the
atmosphere, and so decreases rate of diffusion of
water through the cuticle.

Other adaptations of xerophytes How the adaptation helps limit water loss
reduction in the surface area to volume The smaller the SA:VOLUME ratio the slower
ratio of leaves the rate of diffusion. Some xerophytes like
cacti have spines rather than broad, flat leaves
stomata confined to underside of leaf Underside of leaves is often cooler so less
heat energy to evaporate water.
daylight closure of stomata reduces transpiration during the hottest part of
the day
succulent stems or leaves storage of water

10
Q.11. Give one reason why plants growing on sand dunes need to have xerophytic features even
though there is plentiful rain.
Rain rapidly drains through the sand out of the reach of roots
Sand dunes are in windy situations, this reduces the water potential of the air and so increases the
water potential gradient leading to increased water loss from the soil (wind dries sand out)

Q.12. Describe the similarities in preventing water loss seen in xerophytes and insects.
Stomata and spiracles can be closed to prevent water loss
Waterproofing of cuticle in plants, chitin in insects
Other?

11
12
Investigating the effect of air movement on the rate of transpiration by
using a potometer

Procedure:

1. The potometer was immersed in water to remove air bubbles.

2. The screw clip was opened to fill the capillary tube with water.
3. The branch of the plant was cut in the water.
4. The end of branch was pushed through the hole in the rubber stopper under water.
5. A beaker of water was placed at the end of the capillary tube.
6. The screw clip was closed and the potometer was removed from the water.
7. The surface of the leaves and stems was wiped with tissue paper.
8. Vaseline was spread around every joint to ensure that they are airtight.
9. The potometer was left until the air bubble in the capillary tube moved steadily.
10. Two points, P and Q, of length 10 cm apart were marked on the capillary tube.
11. The time taken for the air bubble to move from P to Q was recorded.
12. The position of the air bubble was reset at point P by opening the screw clip.
13. Steps 11 and 12 were repeated twice.
14. Steps 11 to 13 were repeated by placing the potometer below a moving fan.
15. The rate of transpiration in this case was calculated by using the following formula:
rate of transpiration = 10/t cm min -1 where t = time taken for air bubble to move from P to Q

Results
Air movement Time / minute mean rate of
1 2 3 mean transpiration
/ cm min-1
Still air 20.0 18.5 19.5
Fast moving air 1.6 1.9 1.6

Questions
1. Explain why steps 3 and 4 are carried out in the water.

…………………………………………………………………………………………………………………………….......

…………………………………………………………………………………………………………………………….......
(1 mark)
2. Calculate the mean times and rates of transpiration to three significant figures in the table above.
(2 marks)
3. Identify the independent and the dependent variables in the investigation.

Independent
variable ..............................................................................................................................................

13
Dependent
variable ................................................................................................................................................
(2 marks)
4. Identify two variables that the student should have controlled in this investigation.

…………………………………………………………………………………………………………………………….......

…………………………………………………………………………………………………………………………….......
(2 marks)
5. Describe how your two other variables could have been controlled.

…………………………………………………………………………………………………………………………….......

…………………………………………………………………………………………………………………………….......

…………………………………………………………………………………………………………………………….......

…………………………………………………………………………………………………………………………….......
(4 marks)
6. Draw a suitable graph to display the results of this investigation.

(4 marks)

14
Translocation: transport in the phloem
Plants have another transport tissue called phloem. The table below shows a comparison of xylem
and phloem sap in a typical plant.
substance concentration in xylem sap concentration in phloem sap
-3
/mg dm /mg dm-3
sucrose 0 154000
amino acids 700 13000
potassium ions 90 1540
sodium ions 60 120
magnesium ions 27 85
calcium ions 17 21

Q. 13. What conclusions can you make from the table about the transport of substances in the xylem
and the phloem?
 Only phloem transports sucrose
 Phloem transports more organic solutes such as sucrose and amino acids than the xylem
 Both transport ions but more so in the phloem
 Total concentration in the phloem greater than in the xylem sap, so xylem carries more water

The process by which soluble organic molecules (e.g. sucrose and amino acids) and some mineral
ions (e.g potassium, chloride and phosphate) are transported around the plant as sap is known as
translocation.

Q.14. Why is it necessary to transport sucrose and amino acids from one part of the plant to another?
Only leaves photosynthesis to produce sugars but all parts of the plant requires sugars for
respiration.
Amino acids are required for protein synthesis.

Q.15. Sucrose is a disaccharide and a non-reducing sugar.

a) Write the word equation for the hydrolysis of sucrose.
Sucrose → α-glucose + β-fructose

b) Why might non-reducing sugars such as sucrose be transported in the phloem rather than
reducing sugars such as glucose?
Reducing sugars are too reactive and would be chemically altered before arriving at their destination

Translocation occurs in a tissue known as phloem. Phloem tissue is made up of sieve tube
elements, long thin cells arranged end to end. Companion cells are associated with sieve tubes.

15
Structure of the phloem tissue Sieve tube
element(few
organelles)
 Each sieve tube element links to the
next via a sieve plate which is
perforated with pores.
 The sieve tube has little cytoplasm,
no nucleus, no vacuole and few
organelles other than a small
number of mitochondria.
 The sieve tubes are alive because of
cytoplasmic connections
(plasmodesmata) with the
companion cell.
 Each companion cell has a nucleus,
many mitochondria and other
organelles.

16
The mechanism of translocation
Experiments have demonstrated that translocation can be bidirectional not just unidirectional like
transpiration, and that rates of translocation are around 1 ms -1. This is too rapid to simply be
explained by diffusion. Translocation moves organic solutes from ‘sources’ to ‘sinks’.

Sources are:
 Where the organic solutes are produced and are therefore at a high concentration.

 The source for sucrose is usually the mesophyll cells of the leaves where it is formed by the
condensation of fructose and glucose

Sinks are:
 Where organic solutes are used up and are therefore at a low concentration.

 The sinks are the other parts of the plant, especially the growing points (meristems) of
roots, stems, flowers and leaves where the sucrose is hydrolysed to glucose and fructose
and then respired to provide metabolic energy in the form of ATP.

 Fruits, seeds, roots and other storage organs act as sinks when sucrose is converted into
starch and stored.

17
The mass flow theory is the currently accepted explanation of translocation

1. The transfer of sucrose from photosynthesising tissue to sieve tubes

 Active transport is used to pump the sucrose (and other solutes) from mesophyll cells of the
leaf into companion cells and then into the sieve tube elements of the phloem.
 This requires ATP and special carrier proteins in the cell membranes of companion cells.
 This increases the concentration of sucrose (and other solutes) in the sieve tubes in this
region.

2. Mass flow of sucrose through sieve tube elements

Mass flow is the bulk movement of substances from high to low pressure. The mass flow of sucrose
in sieve tube elements takes place as follows.
 The increased concentration of sucrose (and other solutes) in the sieve tubes lowers the water
potential of the sieve tubes, so water enters by osmosis from the xylem and the companion
cells.
 This creates a high hydrostatic pressure within the sieve tubes around the source.
 At sinks, sucrose is either used up in respiration or converted to starch for storage, so the
sucrose concentration of sinks is low.
 Sucrose is actively transported from sieve tube elements through companion cells into sink
cells.
 This reduces the water potential of these cells and so water leaves the sieve tubes by osmosis
entering sink cells.
 The hydrostatic pressure of sieve tubes in this region is therefore lowered.
 As a result of water entering the sieve tube elements at the source and leaving at the sink,
there is a hydrostatic pressure gradient between the source and the sink.
 Therefore, there is a mass flow of sucrose solution down this gradient in the sieve tubes.

3. Transfer of sucrose from the sieve tube elements into sink cells

 Sucrose is actively transported from sieve tube elements through companion cells into sink
cells.

18
The process of translocation of sucrose in the phloem is summarised in the diagram
below. Annotate the diagram.
High sucrose concentration/low WP

Water enters active transport

phloem by of sucrose
osmosis creating
high hydrostatic
pressure at source

mass flow of sucrose solution down a

hydrostatic gradient in the sieve
tubes.
Water leaves phloemby
osmosis creating low
hydrostatic pressure at
sink

active transport
of sucrose

Low sucrose concentration/high WP

19
 The table below contrasts the structure and roles of xylem and phloem.
Complete rows 1-4 in the table using the most appropriate word or words.
For row 5 make your own contrast.

Xylem Phloem
1. Xylem transports water and named Phloem transports organic solutes
mineral ion such as sucrose/amino acids

2. Xylem vessels have no cross walls Sieve tubes contain perforated cross
walls

3. Xylem vessel walls are impregnated Sieve tubes have no additional

with lignin support

4. Xylem vessel walls contain pits that There are many gaps in the cell walls
allow water to pass into adjacent between companion cells and sieve
vessels tube elements called plasmodesmata

5. Xylem cells are dead Phloem cells are living

Evidence for and against the mass flow theory of translocation in the phloem.

Evidence supporting the mass flow Evidence against the mass flow
theory theory

 There is hydrostatic pressure in the  The theory leaves the function of

phloem as shown by the release of
sap when they are cut.
 The concentration of sucrose is higher
in the leaves (source) than in the roots
(sink).
 Downward flow in the phloem occurs
in daylight, but ceases when leaves
are shaded, or at night.
 Increases in sucrose levels in the leaf  Sucrose is delivered at more or less
are followed by increases in the
phloem a little later.
 Metabolic inhibitors and/or a lack of
oxygen inhibit translocation of sucrose
in the phloem.
 Companion cells have many
mitochondria.
sieve plates unclear as they would
hinder mass flow of sucrose – they
may have a function in preventing
bursting of sieve tubes under
pressure.
 Not all solutes move at the same
speed – they should if moved by mass
flow.
the same rate to all sinks, rather than
going faster to those with the lowest
sucrose concentration as the mass
flow theory would suggest.

20
Q.15. Explain in terms of the mass flow theory why downward flow in the phloem ceases when
leaves are shaded, or at night. (3)
1. No/little photosynthesis and so production of sucrose stopped/reduced;
2. No/little sucrose actively transported into the sieve tubes/phloem;
3. Concentration of sucrose not increased/water potential not decreased in sieve tubes/phloem;
4. No/little water enters sieve tubes/companion cells/phloem from xylem;
5. No/less hydrostatic pressure gradient produced;

Q.16. Explain in terms of the mass flow theory why metabolic inhibitors and/or a lack of oxygen inhibit
translocation of sucrose in the phloem. (3)
1. Respiration inhibited, so no/little ATP produced;
2. No/little active transport of sucrose from mesophyll to sieve tubes/phloem;
3. Concentration of sucrose not increased/water potential not decreased in sieve tubes/phloem;
4. No/little water enters sieve tubes/companion cells/phloem from xylem;
5. No/less hydrostatic pressure gradient produced;

Investigating transport in plants

Ringing experiments
Mason and Maskell (1928)
demonstrated that removing a
complete ring of bark while leaving
the wood (xylem) intact prevented
downward movement of sugars, the
phoem is removed with the bark. This
ringing or girdling has no immediate
effect on transpiration, since water
mainly moves in the xylem, interior to
the bark. However, sugar transport in
the trunk is blocked at the site where
the bark has been removed. Sugars
accumulate above the ring or girdle,
that is, on the side toward the leaves
and are depleted below the treated
region.
Eventually the bark below the girdle dies, while the bark above swells and remains healthy. It was
concluded that sugar is transported in the bark of the tree and that the phloem was the route of sugar
transport.

21
Mason and Maskell carried out further ringing
experiments on cotton shrubs. A varying
amount of bark was left connecting the upper
and lower parts of the stems. The plants were
left for 24 hours and then the mass of
carbohydrate in the lower part of the stem was
measured.

Q.17. Describe the trend is seen in the data.

(1)

There is a positive correlation between the two

sets of data;

A possible conclusion of this experiment is that

removing the bark reduced the mass of
carbohydrate transported to the lower part of
the stem.

Q.18. Evaluate this conclusion based on the

results from the experiment. (3)

There is a positive correlation;

BUT correlation does not infer causation;
The results do not prove that there is a downward movement of carbohydrate in the phloem;
There could be other factors affecting the results. For example it could be that the carbohydrate is
transported in the xylem, but the xylem was damaged in the ringing process;
The experiment has also been carried out on only one species of plant;

Radioactive Tracer Experiments

If a plant is exposed to an atmosphere containing radioactively labelled carbon dioxide 14CO2,

carbohydrates are synthesised using the radioactively labelled carbon. The movement of these
carbohydrates within the plant can be traced using photographic film. The film blackens where it is
exposed to the radioactive isotope, revealing the location of the labelled carbohydrates. This process
is called autoradiography.

In the photomicrograph, to the right,

carbohydrates including sucrose from
the leaves appear in the sieve tube
elements of phloem in the stem. The
location of the label is revealed in the
tissue cross sections by the presence of
dark grains on the film. The label is
confined almost entirely to the sieve
elements and companion cells of the
phloem, demonstrating the organic
solutes such as carbohydrates travel in
the phloem.

22