Transport in plants occurs on three levels:
the uptake and release of water and solutes by individual cells
o absorption of water and minerals from he soil by root cells
short-distance transport of substances from cell to cell
o loading of sucrose from photosynthetic cells into the sieve tube cells of the
phloem
long-distance transport of sap within the xylem and phloem
o this is a whole plant phenomena - transport of photosynthate from leaf to root
Cellular-level Transport
A key component of cellular-level transport is the movement of solutes and ions across the
plasma membrane. We have already covered this, so I won't repeat it. If you are unsure, review
Lecture 9.
Survival of the plant depends on balancing water uptake and water loss.
In an animal cell, water flows from hypotonic to hypertonic solutions, but in a plant cell, there is
the added presence of the pressure created by the cell wall
The combination of solute concentration differences and physical pressure are incorporated
into water potential, abbreviated with the Greek letter psi ( )
Water will flow through a membrane from a solution of high water potential to a solution
of low water potential
Water potential is measured in units of megapascals (MPa)
Pure water has a water potential of 0 MPa ( = 0 MPa)
o The addition of solutes lowers water potential ( = -0.2 MPa for instance)
o An increase in pressure (by lowering a piston for example) will raise water
potential
These two forces combine to form the following equation:
o = p + s
o = total water potential
o p = water potential due to pressure
May be positive or negative
o s = water potential due solute concentration (also known as Osmotic Potential)
Always negative or zero
Movement of Water Through Cells - Two Routes, the Symplast and the Apoplast
Symplastic Movement
Movement of water and solutes through the continuous connection of cytoplasm (though
plasmodesmata)
No crossing of the plasma membrane (once it is in the symplast - however, if the solute
was initially external to the cell, then it must have crossed one plasma membrane to enter
the symplast)
Apoplastic Movement
Movement of water and solutes through the cell walls and the intercellular spaces
No crossing of the plasma membrane
More rapid - less resistance to the flow of water
Absorption of Water and Minerals by Roots
Absorption is a surface area phenomenon - the more surface area there is, the more absorption there will
be.
Root hairs - extensions of the root epidermal cells to increase surface area
Mycorrhizae - fungal associations with roots - greatly increase surface area
o as much as three meters of fungal hyphae can extend from each centimeter of root
o this is an ancient association - some of the oldest terrestrial plant fossils have fungal
associations
o click here to download a pdf of a paper on this if you're interested (its on page 6)
As water is drawn into the root, dissolved minerals are also brought into the root
Water flows through the apoplast and the symplast on its way to the xylem
o The majority of the water, however, travels through the apoplast
The Endodermis - The Root's Border Guard
Water flowing through the apoplast contains many minerals that the plant needs - it may also
contains toxins and substances that the plant may not want. However, since the water is flowing
through the apoplast, there is no way to prevent the passive transport of these toxins, until the
water hits the endodermis.
Endodermis
Cells of the endodermis possess cell walls that are ringed by the Casparian Strip, a waxy layer
(composed of suberin).
The Casparian Strip is a wax and therefore prevents the apoplastic flow of water
Water must pass through the plasma membrane and enter the symplast
The plasma membrane of the endodermal cells contain many transport proteins to actively
transport some molecules in and others to pump other molecules out
Once water passes under the Casparian Strip in the endodermal cells, it is free to enter the
apoplast again on its way to the xylem.
Transport of Xylem Sap
Xylem sap rises against gravity, without the help of any mechanical pump, to reach heights of more than
100m in the tallest trees. How can this occur?
Transpiration-Cohesion-Tension: A Mechanism to Pull Xylem Sap up the Plant
Stomata open up during the day to let CO2 in and inadvertently let H2O escape
There is a gradient in water potential, high water potential in the soil and very low water potential
in the air
Water vapor leaves the air spaces of the plant via the stomates
This water is replaced by evaporation of the thin layer of water that clings to the mesophyll cells
Remember, water has strong adhesive and cohesive properties - as the water leaves, it is replaced
by water clinging to the inside of the cell walls
o This creates a tension (pulling) on the water in the xylem and gently pulls the water
toward the direction of water loss
o The cohesion of water is strong enough to transmit this pulling force all the way down to
the roots
o Adhesion of water to the cell wall also aids in resisting gravity
As we said before, the water column in the tallest trees can be 100m - the tension created by
evaporation of water coupled with the cohesive and adhesive forces is enough to support this
column against the forces of gravity
Root Pressure: A Mechanism to "Push" Xylem Sap Up the Plant
At night, transpiration is almost nil. However, the root cells continue to actively transport
minerals into the stele (the root stele is basically everything surrounded by the endodermis -
primarily the xylem and the phloem).
This active transport lowers the water potential within the stele
Water passively flows into the roots, pushing the water up against gravity
Water that reaches the leaves is often forced out, causing a beading of water upon the leaf tips
known as guttation
In most plants, however, root pressure is not the primary mechanism for transporting the xylem
o Tall trees generate almost no root pressure (the weight of the water pushing down on the
xylem more than counteracts any generated root pressure)
The Control of Transpiration
Water is needed for photosynthesis - it is also lost as a product of obtaining carbon by this very same
process. How does the plant balance is requirement for water with its requirement for carbon in
photosynthesis?
Guard cells control the size of the stomatal openings and thus regulate gas and water exchange
Water loss by a plant through stomatal openings is known as transpiration
The efficiency of a plant can be measured by its transpiration-to-photosyntesis ratio
o The amount of water lost per gram of CO2 assimilated into organic material created by
photosynthesis
o A typical ratio for a C3 plant is 600:1 - for a typical C4 plant it is more like 300:1
As long as plants can pull water from the soil as fast as it leaves from the leaves, there is no
problem
When water loss exceeds water uptake, the plants will wilt as the leaves lose turgor pressure
o The conditions that favor wilting are hot, sunny, and windy days
How Stomates Open and Close
Each stoma is flanked by a pair of guard cells that are capable of changing shape, thereby
widening or narrowing the gap between the two cells
When dicot guard cells take in water by osmosis, they become turgid and swell
o Guard cells are not uniformly thick - this, along with a series of radically oriented
cellulose microfibrils in the cell wall, cause the guard cell to buckle outwards.
As they swell, the gap between the guard cells widens
If the plant loses water, the guard cells become flaccid and the gap closes
The changes in turgor pressure result primarily through the reversible uptake of K+ ions
Stomata open when guard cells accumulate K+ from neighboring epidermal cells
o How does this change the water potential ( ) in the guard cells?
Stomata close when K+ leaves the guard cells into the neighboring epidermal cells
o The transport of K+ is probably coupled to the transport of H+ in an antiport system (see
Fig 36.2)
Stomatal opening is triggered by light
o Blue light receptors are present on the membranes of guard cells
o Stimulation of the blue-light receptors stimulates an ATP-poweed proton pump on the
plasma membrane
o The pumping of H+ out of the cell creates and electrical potential which drives in cations
like K+
Plants also observe a 24 hour cycle (a circadian rhythm)
o If placed in total darkness, the plant will still open its stomates when it normally would if
there was light
Adaptations to reduce transpiration loss in plants growing in dry conditions (xerophytes)
Thick cuticles - prevent water loss from epidermal cells
Succulent (thick) leaves - store water
Loss of leaves/reduction of leaves to form spines - light is not limiting, so photosynthesis can be
carried out by the shoot
o What type of plant am I describing?
White leaves/spines - light colors reflect light and heat, thereby cooling the plant
Trichomes (hairs) - create a more humid microenvironment to reduce evaporative water loss
Sunken stomates - like trichomes, a more humid microenvironment is created
CAM photosynthesis - stomates open during the night (when it is cooler) and fix CO 2 into four-
carbon acids
o The light reaction occurs during the day, generating NADPH and ATP
The Translocation of Phloem
Translocation - the process of moving photosynthetic product through the phloem
In angiosperms, the specialized cells that transport food in the plant are called sieve-tube
members, arranged end to end to form large sieve tubes
Phloem sap is very different from xylem sap
o sugar (sucrose) can be concentrated up to 30% by weight
Phloem transport is bidirectional
o Phloem moves from a sugar source (a place where sugar is produce by photosynthesis or
by the breakdown of sugars) to a sugar sink (an organ which consumes or stores sugar)
o What are some organs which would be sugar sinks?
Phloem Loading and Unloading
Sucrose manufactured in the mesophyll cells can travel via the symplast to sieve-tube members
In some species, sugar can leave the symplast and enter the apoplast, where is it pumped back
into the sieve-tube members and the companion cells
o Some companion cells have cell wall ingrowths that facilitate apoplatic transport of
sucrose into the symplast
Sucrose is loaded into the phloem via a chemiosmotic ATPase mechanism coupled with a
H+/sucrose symport (taken from Lecture 8 notes)
Other Active and Transport Mechanisms - The H+ / Sucrose Pump
o H+ is actively pumped out by hydrolyzing ATP
o H+ accumulated outside the membrane, generating a concentration and electrochemical
gradient
o The H+ cannot cross the membrane, but there is a carrier protein
o H+ binds to carrier protein, but sucrose must also bind. When both are bound, the
configuration changes and the protein opens to the membrane interior.
o Downstream, sucrose must be unloaded, again utilizing an H+ / Sucrose pump
The Mechanism of Translocation in Angiosperms
Phloem loading results in a high solute concentration at the source end of the
o This creates hypnotic conditions in the phloem, causing water to flow into the phloem
o Hydrostatic pressure builds in the sieve tube, but it is greatest in the source
At the sink, osmosis occurs with the unloading of sugar - water flows out of the phloem
The buildup of pressure at the source and the reduction of that pressure at the sink causes water to
flow from source to sink, carrying the sugar along with it.
o Water is recycled via transport in the xylem
This explanation is very simplified - scientists are just now discovering the subtle details of
phloem movement in plants
http://www.scribd.com/doc/4801010/TRANSPORTATION-IN-PLANTS
