Transport in Plants

Chapter 37

Transport Mechanisms
How does water move up to the top of a 10-story high tree?
Water first enters the roots
Then moves to the xylem
Innermost vascular tissue
Water rises through the xylem because of a combination of factors
Most of that water exits through the stomata in the leaves

Local changes result in long-distance movement of materials

Water and dissolved minerals travel great distances in xylem

Some pushing comes from pressure of water entering roots
Most of the force is pulling created by transpiration

Evaporation from thin films of water in the stomata

Occurs due to cohesion (water molecules stick to each other) and
adhesion (stick to walls)
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Water exits
CO2 and light through stomata
Photosynthesis
produces
carbohydrates,
H2O
which travel in
O2 phloem

Stoma
H2O
CO2
O2

H2O Carbohydrates Water goes up

xylem
Carbohydrates
and water go up
and down
phloem

Carbohydrates Water and

H2O
minerals enter
through roots

H2O and H2O and

minerals minerals

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Xylem Phloem

Movement of water at cellular level

Aquaporins

Water can diffuse through plasma membranes

- Osmosis is enhanced by aquaporins Osmosis
If a single plant cell is placed into water
Other substances depend on protein transporters Concentration of solutes inside cell
- Facilitated diffusion or active transport greater than solution
- ATP-dependent hydrogen ion pumps often fuel Water moves into cell by osmosis
active transport Cell expands and becomes turgid
- Unequal concentrations of solutes drive osmosis
If cell placed in high concentration of
Aquaporins sucrose
- water-selective pores in plasma membrane that Water leaves cell
increase the rate of osmosis because they allow bulk Cell shrinks plasmolysis
flow

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 Water Potential
Potentials are a way to represent free energy
Water potential (w) is used to predict which way water will move
Water moves from higher to lower w
Measured in units of pressure called megapascals (MPa)

Water potential has 2 components

1. Physical forces such as plant cell wall or gravity
Contribution of gravity usually not considered
Turgor pressure resulting from pressure against cell wall is pressure
potential p
As turgor pressure increases, p increases
2. Concentration of solute in each solution

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a.

Solutions that are not contained Concentration of solute in each w = p + s

within a vessel or membrane
cannot have turgor pressure,
and they always have a p of 0
MPa
solution The total water potential of
Solute potential (s) smallest a plant cell is the sum of its
amount of pressure needed to pressure potential (p) and
stop osmosis solute potential (s)
Pure water has a s of 0 Represents the total
As solute concentration potential energy of the
increases, s decreases (< 0 MPa) water in the cell
When the w inside the
A solution with a higher solute cell equals that of the
concentration has a more solution, there is no net
negative s movement of water.
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Cell wall and exterior

Once thought water moved across
plasma membranes only by osmosis
through the lipid bilayer Diffusion
Cell membrane
Water moved more rapidly than Bulk flow

predicted

Osmosis is enhanced by membrane

water channels called aquaporins
Speed up osmosis but do not
change direction of water Aquaporin

movement Cytoplasm Water molecules

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 Most of the water absorbed by the
plant comes in through the region
of the root with root hairs
Surface area further increased
by mycorrhizal fungi

Once absorbed through root hairs,

water and minerals must move
across cell layers until they reach
the vascular tissues

Water and dissolved ions then

enter the xylem and move
throughout the plant

Three transport routes exist through cells

1. Apoplast route movement through the cell walls and the space
between cells
Avoids membrane transport
2. Symplast route cytoplasm continuum between cells connected by
plasmodesmata
3. Transmembrane route membrane transport between cells and
across the membranes of vacuoles within cells
Permits the greatest control

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 Eventually on their journey inward, molecules
reach the endodermis

Any further passage through the cell walls is

blocked by the Casparian strips

Molecules must pass through the cell

membranes and protoplasts of the endodermal
cells to reach the xylem

Because the mineral ion concentration in the

soil water is usually much lower than it is in
the plant, an expenditure of energy (ATP) is
required for these ions to accumulate in root
cells

Plasma membranes of endodermal cells

contain a variety of protein transport channels,
through which proton pumps transport
specific ions against even larger concentration
gradients
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Xylem Transport
Root pressure is caused by the continuous accumulation of ions in the roots
When transpiration from leaves is low or absent at night
Causes water to move into plant and up the xylem despite the absence of transpiration
Guttation (production of dew) is loss of water from leaves when root pressure is high
Root pressure alone, however, is insufficient to explain xylem transport
Transpiration provides the main force
Water potential regulates the movement of water through a whole plant
Water moves from the soil into the plant only if water potential of the soil is greater than in
the root
Water in a plant moves along a w gradient from the soil to successively more negative water
potentials in the roots, stems, leaves, and atmosphere

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 Cavitation

An air bubble can break the tensile

strength of a water column
A gas-filled bubble can expand and
block the tracheid or vessel
Damage can be minimized by
anatomical adaptations
Presence of alternative pathways
Pores smaller than air bubbles

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Tracheids and vessels are essential

for the bulk transport of minerals
Ultimately the minerals are relocated through the xylem from the roots
to other metabolically active parts of the plant
Phosphorus, potassium, nitrogen, and sometimes iron
Calcium, an essential nutrient, cannot be transported elsewhere once it
has been deposited in a particular plant part

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 Transpiration pull

Water has an inherent

tensile strength that
arises from the
cohesion of its
molecules
The tensile strength of a
water column varies
inversely with its
diameter
Because tracheids and
vessels are tiny in
diameter, they have
strong cohesive water
forces
The long column of
water is further
stabilized by adhesive
forces

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Rate of Transpiration

Over 90% of the water taken in by the plants roots is

ultimately lost to the atmosphere
At the same time, photosynthesis requires a CO2 supply from
the atmosphere
Closing the stomata can control water loss on a short-term
basis
However, the stomata must be open at least part of the time to
allow CO2 entry

Transportation in Plants
https://www.youtube.com/watch?v=bvPM6sfidY4

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 Guard cells
Only epidermal cells containing chloroplasts
Have thicker cell walls on the inside and thinner cell walls elsewhere
Bulge and bow outward when they become turgid
Causing the stomata to open
Turgor in guard cells results from the active uptake of potassium (K +),
chloride (Cl), and malate
Water enters osmotically

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Active pumping of sucrose out of guard cells in the evening leads to loss of
turgor and closes the guard cell

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 Rate of Transpiration
Transpiration rates increase with temperature and wind velocity because water molecules
evaporate more quickly
Several pathways regulate stomatal opening and closing
Abscisic acid (ABA) initiates a signaling pathway to close stomata in drought
Opens K+, Cl, and malate channels
Water loss follows
- Close when CO2 concentrations are high
- Open when blue wavelengths of light promote uptake of K+ by the guard cells
- Close when temperature exceeds 3034C and water relations unfavorable
Alternative photosynthetic pathways, such as Crassulacean acid metabolism (CAM),

reduce transpiration

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Water Stress Responses

Many morphological adaptations allow

plants to limit water loss in drought
conditions
Dormancy
Loss of leaves deciduous plants
Covering leaves with cuticle and
wooly trichomes
Reducing the number of stomata
Having stomata in pits on the leaf
surface

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 Plants have adapted to flooding conditions
which deplete available oxygen
Flooding may lead to abnormal
growth
Oxygen deprivation most
significant problem
Form larger lenticels and
adventitious roots
Plants have also adapted to life in
fresh water
Form aerenchyma, which is
loose parenchymal tissue with
large air spaces
Collect oxygen and transport it
to submerged parts of the plant

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Plants such as mangroves grow in

areas flooded with salt water
Must supply oxygen to submerged roots and control salt balance
Pneumatophores long, spongy, air-filled roots that emerge above the mud
Provide oxygen to submerged roots
Succulent leaves contain large amount of water to dilute salt
May secrete salt or block salt uptake

Halophytes
Plants that can tolerate
soils with high salt
concentrations
Some produce high
concentrations of organic
molecules in their roots
- This decreases the water
potential enhancing water
uptake from the soil

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 Phloem Transport
Most carbohydrates produced in leaves are distributed through phloem to
rest of plant
Translocation provides building blocks for actively growing regions of the
plant
Also transports hormones, mRNA, and other molecules
Variety of sugars, amino acids, organic acids, proteins, and ions
Using aphids to obtain the critical samples and radioactive tracers to mark
them, plant biologists have demonstrated that substances in phloem can
move remarkably fast, as much as
50 to 100 cm/h

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Pressure-flow theory is a model describing the

movement of carbohydrates in phloem

Dissolved carbohydrates flow from a source and are released at a sink

Sources include photosynthetic tissues
Food-storage tissue can be sources or sinks
Sinks include growing root and stem tips as well as developing fruits

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 Phloem-loading occurs at the source
Carbohydrates enter the sieve tubes in the smallest veins at the source
Sieve cells must be alive to use active transport to load sucrose
Water flows into sieve tubes by osmosis
Turgor pressure drives fluid throughout plant
At sink, sucrose actively removed and water follows by osmosis
Water may be recirculated in xylem or lost

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