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3.2.

2 Transport in Plants
The need for a Transport System
Diffusion is INADEQUATE for transport within multicellular plants as:

Their S.A.:V is SMALL

DISTANCES that materials have to travel are very LARGE.

QUANTITY of materials moving is LARGE.

This renders DIFFUSION an INEFFICIENT method to distribute substances


around the body. Thus TRANSPORT SYSTEMS evolved.

NOTE: The same reasons apply in the case of animals, which evolved
circulatory systems.

Three levels of transport in plants


Transport in plants occurs on 3 levels:

THE CELLULAR LEVEL: the UPTAKE and RELEASE of WATER and SOLUTES by
INDIVIDUAL CELLS, e.g. water and mineral absorption by root cells.

THE TISSUE LEVEL: SHORT DISTANCE TRANSPORT of substances from cell to


cell at the LEVEL of TISSUES and ORGANS e.g. loading sugars from
mature cells of leaves to sieve tube elements of phloem.

THE WHOLE PLANT LEVEL: LONG DISTANCE TRANSPORT within XYLEM and PHLOEM
over the whole plant.

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Transport at the cellular level
Revise relevant parts in topic 2 (esp. osmosis and secondary active
transport)

Short-distance transport at the tissue level


LATERAL MOVEMENT refers to movement of water and solutes from one place
to another in a PLANT TISSUE or ORGAN. It usually occurs along the RADIAL
AXIS of the plant e.g. from the outside (circumference of root) to inside
(centre of root), or vice versa.

Most mature plant cells have 3 compartments:

the CELL WALL,


the CYTOPLASM,
the LARGE CENTRAL VACUOLE.

The PLASMA MEMBRANE controls the passage of substances between the CELL
WALL and the CYTOPLASM.

The TONOPLAST controls the passage of substances between the CYTOPLASM


and the LARGE CENTRAL VACUOLE.

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This control is ACHIEVED by means of SPECIFIC PROTEINS embedded in the
plasma membrane and the tonoplast.

In most plant tissues:

The CELL WALLS of ADJACENT CELLS form a continuous compartment,


called the APOPLAST.

The CYTOPLASM of ADJACENT CELLS may form a continuous compartment


if PLASMODESMATA are present. This is called the SYMPLAST.

This organisation renders 3 routes for lateral transport:

1. TRANS-MEMBRANE PATHWAY: substances move out of one cell through


the plasma membrane, across the cell walls, and into the
neighbouring cell through its plasma membrane. These substances
continue moving along adjacent cells by means of the same
mechanism.

2. APOPLAST PATHWAY: water flows through cell walls and extra cellular
spaces and carries solutes with it. Substances will only move
through the plasma membrane when the destination is reached.

3. SYMPLAST PATHWAY: Water and solutes travel from cell to cell via
plasmodesmata. Requires only one crossing through the plasma
membrane (by osmosis). After entering one cell, solutes and water
move from cell to cell via plasmodesmata.

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4
Long distance transport at the whole plant level
Long distance transport involves VERTICAL MOVEMENT that refers to
movement of water (and solutes) and products of photosynthesis from
one place to another in a WHOLE PLANT. It usually occurs along the VERTICAL
AXIS of the plant, in xylem and phloem respectively.

Diffusion is too slow for transporting substances over long distances e.g.
from roots to leaves. Therefore water and solutes move through xylem
vessels and phloem sieve tubes by BULK FLOW i.e. the movement of a fluid
driven by PRESSURE.

Transport in the Root


Water and mineral salts from soil enter the plant through the EPIDERMIS of
the ROOTS, across the ROOT CORTEX, pass into the STELE, and then flow up
XYLEM VESSELS to the SHOOT SYSTEM.

Transport of water from soil to epidermis


Most of WATER ABSORPTION occurs:

Near the ROOT TIP as the epidermis here is PERMEABLE to water.

Through the ROOT HAIRS, which are EXTENSIONS of epidermal cells that
increase the SURFACE AREA of the root.

Through the MYCORRHIZAE i.e. symbiotic relationships between fungi


and roots. The HYPHAE ABSORB water and selected minerals and then
TRANSFER them to the roots of the HOST PLANT.

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A Soil particle surrounded bsorp
tion by film of water of
water
and Root hair
Water

Air space

minerals generally occurs in the YOUNGER ROOTS, as the OLDER ROOTS are LESS
PERMEABLE due to SUBERIN deposits in the outer cells of the root cortex.

Soil particles are usually coated with water and dissolved minerals. The
CELL WALLS of the epidermal cells ATTRACT water.

As the soil solution coating the soil crumbs has a higher ψ than the
epidermal cells. Water moves into the cells by OSMOSIS. These cells are
always turgid due to a high concentration of mineral ions inside them.

Two pathways may be followed:

Water flows into the walls of epidermal cells and start flowing to the
parenchyma cells of the cortex via the APOPLAST PATHWAY. This
process is NOT SELECTIVE as the cell wall is permeable to all
substances. It is the PREFERRED PATHWAY as it offers the LEAST
RESISTANCE.

Water flows across the plasma membrane of the epidermal cells and
is transported to the parenchyma cells of the cortex via the SYMPLAST
PATHWAY. As the plasma membrane is selectively permeable,
SELECTIVE ABSORBTION OF IONS is possible.

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Uptake of mineral ions from soil to epidermis
As the root RESPIRES, CO2 gas is released. It dissolves in the water film
around the soil crumbs, rendering the solution SLIGHTLY ACIDIC. This causes
the CATIONS (+vely charged ions) to be released and hence the root will
ACTIVELY absorb them.

The epidermal cells ACTIVELY PUMP H+ ions out of them due to a PROTON
PUMP. This has two effects:

The extracellular environment of the epidermal cells becomes


POSITIVELY CHARGED compared to the intracellular environment.

A DIFFERENCE in PROTON CONCENTRATION results across the cell wall.

Positively charged ions such as K+ move into the cell through the specific
ion channels on the membrane. (+ve ions move into a region of -ve
charge)

The difference in the proton concentration drives the active transport of


-vely charged ions e.g. Cl-, SO42-, NO3-. In fact as H+ ions flow back into
the epidermal cells, they drag in with them the –ve ions (example of co-
transport).

(Note: The mineral ions are actually the plant nutrients).

K
K K
Cl K
H
K
K K
K

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Transport from epidermis to root cortex
Water and mineral ions move across the ROOT CORTEX up to the ENDODERMIS
mainly via the:

SYMPLAST ROUTE: water crosses one cell wall and one plasma
membrane and then runs through the plasmodesmata in the
symplast.

APOPLAST ROUTE: water runs through the extracellular continuum


formed by the cell walls, crosses into one cell wall, crosses no
plasma membrane

(NOTE: As water and minerals flow through the cortex they may switch
from the apoplast to the symplast pathway.)

Transport from root cortex to xylem


When water and mineral ions reach the ENDODERMIS (innermost layer of the
root cortex), the CASPARIAN STRIP (or band) BLOCKS the APOPLAST pathway.
This is a belt of an IMPERMEABLE waxy material called SUBERIN that runs all
around the endodermal cell.

Therefore all water and minerals that will be transported up the plant
must cross into a cell and pass through a plasma membrane i.e. enter the
SYMPLAST pathway.

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ADVANTAGES OF THE CASPARIAN STRIP

It SLOWS DOWN the flow of water through the root.

It CONTROLS the entry of toxic substances and fungal pathogens as


water passes through the plasma membrane.

The transport proteins in the plasma membrane of these cells


determine which mineral ions pass into the stele and at what rate.
This enables the plant to CONTROL ITS CHEMICAL COMPOSITION thus
striking a balance in the concentrations of essential nutrients.

Water and mineral ions move from the endodermal cells into the
PARENCHYMA CELLS within the STELE.
The mineral IONS are ACTIVELY PUMPED into the xylem vessels when these are
reached. This LOWERS the ψ of the solution inside the xylem. Thus WATER
moves into the xylem by OSMOSIS.

The water and mineral ions absorbed from the soil are now ready for
UPWARD TRANSPORT into the shoot system.

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Root pressure
ROOT PRESSURE refers to the pressure EXERTED BY ROOT tissues that FORCES
liquid up the xylem.

This pressure arises due to the xylem water and solutes having a lower
(more –ve) ψ than the soil. This arises from the accumulation of ions in
the stele. The –ve potential draws water into the stele, which exerts
pressure upwards.

Evidence for root pressure

GUTTATION: liquid water is forced


out through special openings called
HYDATHODES in the leaves. This is
best observed:
1. When humidity in atmosphere is
high.
2. When water is plentiful in the
soil.
3. In early morning as at night
transpiration is very low or zero.

Causes water (and solutes) to OOZE


out of the cut stumps of some
plants e.g. Coleus.
However as root pressure is very low (rarely above 1atm) it only plays a
MINOR ROLE in the transport of solution in plants.

Vertical transport
Water rises up the xylem from the root to the stem and shoots AGAINST
GRAVITY. The xylem water (and solutes) flows upward at 15 meters/hr or
faster.

Water rises up the xylem due to:

ROOT PRESSURE (minor role) that PUSHES the xylem water (and solutes)
upwards.

The TRANSPIRATION-COHESION-TENSION MECHANISM that PULLS the xylem


sap upwards.

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The Transpiration-Cohesion-Tension Theory mechanism

Transpiration
Transpiration refers to the EVAPORATION OF WATER at the stomata in the
leaves (or evaporation of water from the aerial parts of the plant).

In fact only a very small percentage of water reaching the leaves is used
for photosynthesis. The rest is lost by evaporation.

Stomata lead to AIR SPACES within the LEAF MESOPHYLL TISSUE. The air in these
spaces is SATURATED with water vapour because it is in contact with the
moist walls of the cells.

On most days the air outside the leaf is drier than the air inside the leaf.
Therefore water vapour diffuses down its concentration gradient and exits
the leaf via the stomata.

As the THIN FILM OF WATER coating the mesophyll cells EVAPORATES to saturate
the air spaces A NEGATIVE PRESSURE is created (a pressure LESS than
atmospheric pressure).

This negative pressure, or


TENSION, is the PULLING FORCE that
draws water OUT OF the leaf
XYLEM, through the MESOPHYLL,
and toward the cells and air
SPACES near the STOMATA.

Flow of water occurs via the


SYMPLAST and the APOPLAST
PATHWAYS. As water is lost from
mesophyll cells, their ψ
becomes more NEGATIVE. Thus
more water enters from the
nearest xylem vessel (in vein).

As water is removed from the


xylem of the veins in the
leaves, a TENSION is established
on the ENTIRE COLUMN OF WATER
contained within the xylem, so
the COLUMN IS DRAWN UPWARD all the way from the roots.

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12
FORCES OF COHESION AND ADHESION

The tension generated in the leaves (TRANSPIRATIONAL PULL) is transmitted


all the way from the leaves to the root tips. This is due to:

COHESION

This refers to the TENDENCY OF WATER MOLECULES TO STICK TOGETHER [due


to the presence of hydrogen bonds].

Cohesion is responsible for an UNBROKEN COLUMN of water being


pulled all the way from roots to the leaves.

ADHESION

This refers to the TENDENCY of WATER MOLECULES TO STICK TO THE CELL


WALLS. The smaller the diameter of the column, the greater the
tensile strength.

Adhesion helps water to rise up the plant against gravity, as water


adheres to the hydrophilic walls of the xylem vessels.

As water evaporates from the cells in the leaves, the resulting tension
pulls up more water to replace water that has been lost.

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14
Passive nature of transpiration
All these processes are PASSIVE. Evidence supporting this statement:

Xylem is a DEAD TISSUE. Therefore no active transport can occur


here.

If CYANIDE is applied to xylem, movement of water CONTINUES.

As xylem is DEAD it offers LESS RESISTANCE to the water flow. If part of


the stem of a tree is cut in half thus damaging only some xylem
vessels, the WATER FLOW in the UNDAMAGED XYLEM VESSELS is not affected.
This would not be possible if active transport was observed in
xylem.

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Cavitation
Cavitation (or embolism) occurs when an AIR BUBBLE gets BLOCKED in the
xylem vessel. This breaks the column of water, stop water transport and
can lead to dehydration and death or part or all of the plant. In nature
cavitation never occurs except:

When the plant is exposed to a lot of RADIATION.


When the TEMPERATURE falls below zero or it is very high.
When cells are deformed.

Anatomical adaptations compensate for cavitation. When it occurs, WATER


moves through alternative routes through pores connecting to other
tracheids or vessels. This SLOWS the water flow up a tree. Cavitation is
one of the reasons that older xylem no longer conducts water.

Xylem Structure
Xylem tissue has TWO major roles in plants:

TRANSPORT of water and mineral ions.


SUPPORT.

It consists of 4 cell types:

Tracheids
Vessel elements
Parenchyma
Fibres

XYLEM VESSELS are mainly responsible for the transport of water in flowering
plants.

They are very LONG TUBULAR STRUCTURES formed when several VESSEL ELEMENTS
fuse together.

Xylem vessels exhibit a number of ADAPTATIONS for the efficient transport of


water:

VESSEL DIAMETER is wide to MINIMISE FRICTIONAL RESISTANCE.

Vessels also LACK CYTOPLASM that would impede water flow.

LIGNIFICATION prevents walls from collapsing because of tension.

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Lignin deposition may be ANNULAR, SPIRAL or RETICULATE.

ABSENCE of CROSS WALLS.

Formation of a continuous tube helps reduce frictional resistance.

Control of Transpiration
Transpiration occurs from:

STOMATA: 90% of total water lost. Most water is lost from the LOWER
SURFACE of the LEAF due to a larger number of stomata here.

LENTICELS: these are spaces between loosely arranged CORK CELLS in


the BARK OF TREES.

CUTICLE: water is lost from the cuticle when it is very THIN e.g. in
FERNS.

Transport of water in the leaf


Water arriving in the XYLEM VESSELS in the leaf has a HIGHER ψ than the
MESOPHYLL CELLS.

Thus water moves OUT of the xylem into the mesophyll largely through the
APOPLAST PATHWAY. Water evaporates and diffuses out of the leaf through
the stomata.

[Note: The evaporation of water itself provides a pulling force which


pulls water out of the xylem into the leaf.]

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The need for stomata
A stoma is a GAP in the epidermis. A PAIR of SPECIALIZED epidermal cells
called GUARD CELLS controls its OPENING and CLOSING.

The stomata lead to a honeycomb of AIR SPACES. This INCREASES the SURFACE
AREA through which:

CO2, diffuses to the PHOTOSYNTHETIC CELLS in the mesophyll.

H2O evapoRAtes and is then lost through the stomata.

Water loss through the rest of the leaf surface is limited by the WAXY CUTICLE
that is secreted by the leaf epidermis.

THEREFORE STOMATA MUST ENABLE EFFICIENT CO2 UPTAKE BUT REDUCE WATER
LOSS TO A MINIMUM.

The TRANSPIRATION TO PHOTOSYNTHESIS RATIO evaluates how EFFICIENTLY a plant


uses water:

WATER LOSS [cm3]: CO2 ASSIMILATION [g]

Most plants 600: 1


Corn 300:1

Functions of Stomata
GASEOUS EXCHANGE of CO2 and O2 between the leaf and the
atmosphere.

When water is lost, the plant COOLS down as heat is eliminated.

Due to uptake of water, minerals are TRANSPORTED along the plant.

Structure of Stomata
Each stoma (sing.) consists of a pore between a PAIR of GUARD CELLS. These
are surrounded by SUBSIDIARY CELLS.

DICOTS generally have SAUSAGE SHAPED guard cells. GRASSES and SEDGES have
DUMB-BELL SHAPED guard cells.

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ADAPTATIONS OF GUARD CELLS

INNER WALL (i.e. cell wall closer to stoma) is THICKER due to more
cellulose deposited on it. Therefore it is LESS ELASTIC than the outer
wall.

Unlike other epidermal cells they contain CHLOROPLASTS.

Mechanism causing stomata to open and close


In the majority of plants stomata are open during the day and closed at
night.

During the DAY, light is available. The stomata OPEN allowing CO2 to
diffuse into the leaf enabling photosynthesis.

At NIGHT, in the absence of light, photosynthesis cannot proceed.


Hence stomata remain CLOSED thereby conserving water.

Opening mechanism

When stomata are about to open, H+ ions are ACTIVELY pumped out of
guard cells by means of a proton pump.

[Note: H+ are obtained from MALIC ACID, an organic acid formed due to
conversion of starch.]

This leads to the entry of K+ ions into the GUARD CELLS, via specific ion
channels, from the surrounding epidermal cells.

The guard cells’ increase in positive charge due to the entry of K+ is


-
BALANCED by the entry of negatively charged ions e.g. Cl . These are
+
dragged into the cells along with the H ions that diffuse back into the
guard cells down a concentration gradient.

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This LOWERS the ψ within guard cells. Therefore water ENTERS by OSMOSIS,
rendering them more TURGID.

When water enters the guard cells their length remains constant but their
volume increases. (Volume may increase by up to 50%.)

[Note: Most of the K+ and water are stored in the vacuole, and thus the
tonoplast also plays a role in stomatal activity.]

However the THICKNESS of the CELL WALL of the guard cells is not even:

OUTER WALL is THINNER


and more FLEXIBLE.

INNER WALL is THICKER


and not so flexible.

As a result the guard cells


BEND, the ends swell and
the thickened regions are
drawn apart.

Closing mechanism

The stomata close by


REVERSE PROCESS.

K+ ions DIFFUSE OUT of the


guard cells PASSIVELY,
decreasing the ψ of the
neighbouring epidermal
cells. WATER flows out of
the guard cells by OSMOSIS.

Turgidity is lost; guard


cells collapse thus closing
the stoma in between.

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Factors influencing stomatal movements
a] Water

Some guard cells MONITOR their own ψ. If it is too LOW (i.e. water is
lacking), they release ABSCISIC ACID (ABA), a plant hormone. This causes
K+ ions to move out of the guard cells leading to the closure of the
stomata, preventing loss of H2O from the plant.
b] CO2

During the DAY CO2 level in air spaces is LOW as the cells use it for
PHOTOSYNTHESIS. This favours the OPENING of the stomata, allowing CO2 to
diffuse in.

At NIGHT only RESPIRATION occurs in the plant. This increases the CO2 level
in the air spaces in the leaf, leading to the CLOSURE of the stomata and
conservation of water.

c] Temperature

At LOW TEMPERATURES guard cells behave SLUGGISHLY and tend to remain


CLOSED.

As the temperature RISES they act more rapidly, thus stomata tend to
remain OPEN.

Yet if temperature RISES FURTHER, the rate of respiration increases releasing


a lot of CO2 that causes stomata to CLOSE.

The stomata may close when the temperature exceeds 30-34°C and water
relations are unfavourable. To ensure sufficient gas exchange, these
stomata open when it is dark and the temperature has dropped.
Succulents (cacti) are able to collect CO2 in a modified form (organic
compounds) at night when the temperature is low. These compounds are
decarboxylated during the day, providing a source of CO2 for fixation
when stomata are closed. This helps in the conversion of water.

d] Light

Light activates the BLUE-LIGHT RECEPTOR on the plasma membrane of guard


cells. This stimulates the active transport of H+ through the proton
pumps leading to OPENING of stomata.

Light also triggers photosynthesis in chloroplasts of guard cells, making


ATP available for active transport of H+.

e] Circadian Rhythms

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All eukaryotic organisms have INTERNAL CLOCKS that somehow keep track of
time and regulate cycles. Cycles that are REPEATED DAILY (i.e. every 24
hours) are called CIRCADIAN RHYTHMS.

Guard cells have a built in circadian rhythm. In fact if a plant is kept in


the dark, its stomata will continue their daily rhythm of opening and
closing.

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12.00

09.00 15.00

18.00
06.00

21.00
3.00
24.00

Light
Low CO2
Moderate temperature
Sufficient water

Stoma closed Stoma open

Darkness
High CO2
Extreme temperature
Water deficiency

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Pore size and stomatal density
Water vapour escaping through a stoma spreads out into CONCENTRIC zones
called DIFFUSION SHELLS.

The rate of diffusion depends on:

The DIAMETER of the STOMA: diffusion rate is GREATER through SMALL


STOMATA than through large stomata.

The DISTANCE BETWEEN STOMATA: when stomata are CLOSE together the
diffusion shells of adjacent stomata OVERLAP. This forms one large
area from which water diffuses to the atmosphere and REDUCES
TRANSPIRATION RATE.

Factors affecting transpiration


LIGHT: INDIRECTLY affects transpiration as it leads to opening of
stomata due to PHOTOSYNTHESIS in GUARD CELLS.

HUMIDITY: high transpiration rate favoured by LOW HUMIDITY as


CONCENTRATION GRADIENT of water vapour between leaf air spaces and
air is LARGE.

TEMPERATURE: high temperature favours high transpiration rate as


temperature LOWERS HUMIDITY of air, and provides LATENT HEAT of
VAPORISATION, and INCREASES K.E. of water molecules.

AIR MOVEMENTS: favour a high transpiration rate. Air movements


sweep away water vapour in the air around stomata, INCREASING the
CONCENTRATION GRADIENT of water vapour between the leaf air spaces
and air.

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ATMOSPHERIC PRESSURE: the LOWER the atmospheric pressure (e.g. in
high altitudes) the GREATER the RATE of transpiration.

WATER SUPPLY: if water in soil is LACKING, the plant wilts and STOMATA
CLOSE, thus transpiration stops.

LEAF STRUCTURE: influences transpiration depending on whether it


shows ADAPTATIONS against water loss or not.

Translocation of sap (sugars in solution) in


phloem
Chemical analyses of phloem sap indicated that its MAIN COMPONENT is
SUCROSE (may reach 30%). It may also contain MINERALS, AMINO ACIDS and
HORMONES.

Phloem sap samples are extracted by means of APHIDS, small insects. The
aphid drives its STYLET, which is a SHARP MOUTHPART that functions like a
hypodermic needle, between the epidermal cells and withdraws sap from
a sieve tube cell.

If the aphid is anaesthetized using ether, its body can be carefully cut
away, leaving the stylet. Phloem sap is then collected and analysed.

Translocation
Translocation refers to the TRANSPORT of the PRODUCTS OF PHOTOSYNTHESIS
(food) through PHLOEM to the rest of the plant.

Translocation may occur along the VERTICAL AXIS of the plant or along its
RADIAL AXIS.

In this process SIEVE TUBES carry food from a SUGAR SOURCE to a SUGAR SINK.

A SUGAR SOURCEis a PLANT ORGAN where SUGARS are PRODUCED by


PHOTOSYNTHESIS or by the BREAKDOWN OF STARCHES. Mature LEAVES are the
primary sugar sources.

A SUGAR SINK (destination) is an ORGAN where SUGARS are STORED (as


starches in tubers) or USED (roots, stems, growing parts of plant).

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Proof that phloem is site of translocation
BARK RINGING: a ring of bark is removed from a region in the plant
i.e. all tissues external to xylem are removed. After photosynthesis
sucrose accumulates in the region just above where the bark was
removed showing that sucrose is transported in phloem

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P ARASITIC
ORGANISM
s:

DODDER:
a parasitic
plant.
Sucker-
like
process
that

penetrate the host attach to the latter’s phloem.

APHID: parasitic insect inserts stylet into phloem.

Honeydew Stylet
droplet
of aphid

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AUTORADIOGRAPHY

This technique may be used to determine the passage of substances


through specimens, sites of reactions and their rates.

A plant is enclosed in a perspex box in an atmosphere containing 14CO2.


The path of the 14C used to build sucrose is then traced by means of
autoradiography.

In this process plants of the same species are exposed to radioactive C-


14 for varying amounts of time. Then they are fixed and processed. They
are placed in contact with a photographic plate e.g. X-ray film.

After sufficient time passed, the images obtained showed the distribution
of the radioactive sucrose in phloem tissue because the regions where
radioactivity was present appear ‘fogged’.

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Phloem loading
SUCROSE formed in the LEAF MESOPHYLL CELLS by photosynthesis, may reach
the sieve tube elements via the SYMPLAST and/ or the APOPLAST pathways.

When sucrose reaches the COMPANION CELLS (sometimes called TRANSFER


CELLS) and the SIEVE TUBE ELEMENTS, it switches from the apoplast to the
symplast pathway by SECONDARY ACTIVE TRANSPORT.

A PROTON PUMP ACTIVELY pumps H+ ions out of the companion cell or


sieve tube element.

This creates a CONCENTRATION GRADIENT of H+ ions across the


membrane.

As H+ ions FLOW BACK into the phloem DOWN a concentration gradient


(i.e. PASSIVELY), sucrose is DRAGGED into the phloem.

Pressure flow model (bulk flow)

As sucrose enters the phloem it LOWERS the ψ of the sieve tube elements.
Therefore WATER follows into them by OSMOSIS because their ψ is lower than
that of the surrounding cells.

29
This INCREASES the HYDROSTATIC PRESSURE (turgor) at the SOURCE end of the
tube.

However, at the SINK end sucrose LEAVES the sieve tube elements into the
neighbouring cells where it is stored.
This movement of sucrose develops a lower ψ OUTSIDE the sieve tube
elements. Thus water LEAVES the sieve tube elements by OSMOSIS.

This DECREASES the


HYDROSTATIC PRESSURE at
the SINK end of the
tube.

THE BUILDING OF
PRESSURE AT ONE END OF
THE TUBE (SOURCE) AND
REDUCTION OF THAT
PRESSURE AT THE OTHER
END (SINK) CAUSE WATER
TO FLOW FROM SOURCE TO
SINK, CARRYING SUGAR
ALONG.

(Note: Water moves


back to xylem where
it is recycled back to
the source.)

This model may be


used to explain why
in the phloem
movement is NOT UNI-
DIRECTIONAL like xylem
flow. In fact the
phloem sap can move
in any direction, as
long as a hydrostatic
pressure gradient
exists.

Proof for the pressure flow model

30
Aphids are allowed to feed on a plant and then their body is removed
leaving the stylet embedded in the phloem.

The CLOSER the stylet is to a SUGAR SOURCE, the FASTER the sap will flow out
and the GREATER its sugar concentration. These results support the
pressure flow model.

Phloem structure
The main role of phloem tissue in plants is the transport of organic
solutes.

Phloem tissue consists mainly of 5 types of cells:


Sieve tube elements
Companion cells
Phloem parenchyma
Phloem fibres
Phloem sclereids

Sieve-tube elements are aligned to form long tube-like structures called


SIEVE TUBES.

SIEVE-TUBE ELEMENTS are LIVING CELLS,


interconnected by PERFORATIONS in
their end walls formed from
enlarged and modified
plasmodesmata (SIEVE PLATES).
Sieve-
tube Lateral walls are THICKENED due to
member
higher amount of cellulose
deposited.

These cells RETAIN their PLASMA


Sieve MEMBRANE, but they have LOST their
plate
NUCLEI and much of their CYTOPLASM.

However, mature sieve elements


RETAIN a modified endoplasmic
reticulum, mitochondria, numerous
types of proteins and specialized
sieve element plastids. Most of
these components are arranged along the lateral walls of the sieve
elements.

The absence of a nucleus implies that sieve tube elements rely on


associated COMPANION CELLS for their maintenance. In fact they are
connected together by means of PLASMODESMATA

31
Companion cells are SMALLER than sieve tube elements. They have THIN
CELLULOSE WALLS, a NUCLEUS and are very rich in ORGANELLES.

This shows that companion cells are involved in the ACTIVE TRANSPORT of
soluble food molecules into and out of sieve-tube elements through
porous sieve areas in the wall.

Adaptations of Plants to particular


environments
Adaptations of Xerophytes

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Morphological adaptations to living in dry regions and confer ability to
survive long periods of drought.

SMALL, THICK LEAVES (reduces S.A.:V)

Secrete a HEAVIER layer of CUTICLE over the leaf epidermis to retard


water loss.

Dense covering of TRICHOMES (EPIDERMAL HAIRS) to trap humid air


around them.

IN DECIDUOUS PLANTS LEAVES produced only if water is abundant, and


shed them when it is scarce.

CACTI have SPINES rather than typical leaves, so photosynthesis


confined to fleshy stems, also spines reflect incident radiation and
disperse heat.

SUCCULENTS have FLESHY LEAVES OR STEMS in which water is stored.

Ammophila arenaria (marram grass), corn and related grasses


ROLLS UP its LEAVES trapping the humid air around it and reducing leaf
surface area.

Some trees hang leaves VERTICALLY thus avoiding mid-day sun e.g.
eucalyptus.

STOMATA concentrated on the LOWER LEAF SURFACE.

FEWER STOMATA in NUMBER but HIGH STOMATAL DENSITY therefore less water
lost due to overlapping of diffusion shells.

SUNKEN STOMATA IN PITS e.g. in Pinus (pine tree), this reduces the
drying effects of air currents (Pinus is not a xerophytes)

Some plants have a TAP ROOT SYSTEM. Roots grow to great depth,
sufficient to reach underground water supplies e.g. mesquite tree,
acacia, oleander.

Cacti have shallow but extensive FIBROUS ROOT SYSTEMS that


effectively trap water at the surface of the soil following even light
rains.

Desert annuals evade periods of drought. They carry out their


entire life cycle from seed to seed during a brief period in which
rainfall has made the surrounding desert soil moist.

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Accumulate amino acid proline and other solutes in their vacuole.
This enables better osmotic uptake of water from soil.

These adaptations reduce water loss but also carbon dioxide uptake.
Therefore they limit photosynthesis, consequently most xerophytes are
slow growing.

Adaptations of Halophytes
Plants adapted to a SALINE HABITAT.

Saline environments pose AN OSMOTIC PROBLEM due to the HIGH NEGATIVE ψ of


the environment. To obtain water from such an environment, plants must
have even more negative ψ than that of plants in non-saline
environments.

ACCUMULATE Na+ and Cl- in leaves.

SALT GLANDS in leaves excrete salt thus reducing the danger of


poisoning by accumulated salt e.g. tamarisk.

SUCCULENCE.

OPEN stomata during the NIGHT and CLOSE them during the DAY. [CAM
plants]

Adaptations of Mesophytes
Plants adapted to live in habitats with adequate water supply.

Adapted to grow in well-drained soils with their aerial system


exposed to moderately dry air. Loss of water is substantial but it is
controlled by CLOSURE OF STOMATA.

Deficit of water created during the day is COMPENSATED as water


uptake continues during the night.

WOODY PERENNIAL mesophytes shed their leaves before winter sets in,
thus water loss is reduced before water freezes.

HERBACEOUS PERENNIALS lose aerial system during the unfavourable


season, surviving as UNDERGROUND ORGANS e.g. bulbs, corms and
rhizomes

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Survive winter as DORMANT SEEDS.

Adaptations of Hydrophytes
Plants adapted to live in habitats that have a large supply of water or
grow wholly or partly submerged in water.

Tend to form THIN LEAVES that LACK characteristics to conserve water.

Air spaces in leaf parenchyma (aerenchyma) this is important as a


store of gases and for buoyancy.

CUTICLE tends to be very THIN.

Have NO STOMATA (if leaves are COMPLETELY SUBMERGED in water.)

Vascular tissue is REDUCED, especially xylem.

Roots have no oxygen supply therefore they carry out alcoholic


fermentation (anaerobic) to produce ATP

Some (example black mangrove, Avicennia germinans) have roots and


with pneumatophores – extensions that grow out of the water. These
extensions have lenticels spongy tissue for diffusion of oxygen.

Plants whose LEAVES FLOAT on the surface, e.g. WATER LILY:

Have STOMATA on their UPPER EPIDERMIS only.

Tend to have LARGE AIR SPACES to provide BUOYANCY.

Have SEVERAL LAYERS of PALISADE mesophyll cells above the spongy


mesophyll cells.

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