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Plant Physiology talk Six
Solute transport
Solute transport
• Plant cells separated from their environment
by a thin plasma membrane (and the cell
wall)
• Must facilitate and continuously regulate the
inward and outward traffic of selected
molecules and ions as the cell
– Takes up nutrients
– Exports wastes
– Regulates turgor pressure
– Send chemical signals to other cells
Two perspectives for
membrane transport
• Cellular level
– Contribution to cellular functions
– Contribution to ion homeostasis (i.e., balance)
• Whole-plant level
– Contribution to water relations
– Contribution to mineral nutrition
– Contribution to growth and development
Moving into cells and between compartments
requires membrane to be crossed
• Composed of a
phospholipid bilayer and
proteins.
• The phospholipid sets up
the bilayer structure
• Phospholipids have
hydrophilic heads and
fatty acid tails.
• The plasma membrane is
fluid--that is proteins
move in a fluid lipid
background
Membrane potential
• Arise because charged
solutes cross membranes
at different rates
• Create a driving force for
ionic transport
• Maintained by energy-
dependent electrogenic
pumps
Electrogenic pumps and membrane
potential
• Electrogenic pumps are
ATPases (enzymes that split
ATP)
• ATPases use ATP energy to
“pump” out protons (H+) to
create charge gradients
• H+ gradients create a type
of “battery” to power
transport and maintain ion
homeostasis
Electrogenic pumps and membrane
potential
• To prove this
• Add cyanide (CN)
– Rapidly poisons
mitochondria, so cells ATP
is depleted
– Membrane potential falls
to levels seen with
diffusion
• So membrane potential has
too parts
– Diffusion
– Electrogenic ion transport
• Requires energy
Ion homeostasis within plant cells
• Plant cells segregate
ions based upon:
– Function or role
– Potential toxicity
• This segregation
creates a balance
• Creating and maintaining
the balance may require
energy
Ion homeostasis within plant
cells
• Ion concentrations in cytosol and
vacuole are controlled by passive
(dashed) and active (solid)
transport processes
• In most plant cells vacuole takes
up 90% of the cell volume
– Contains bulk of cells solutes
• Control of cytosol ion concs is
important for the regulations of
enzyme activity
• Cell wall is not a permeability
barrier
– It is NOT a factor in solute
transport
Passive vs active transport
• Passive or active transport depends on the
gradient in electrochemical potential
• The electrochemical potential has 2 parts
– Concentration
– Charge (Electrical)
• The two parts together dictate the
electrochemical potential for a compartment
of a cell
Passive v. active transport
• Passive transport
– Movement down the electrochemical gradient
– From a more positive electrochemical potential
– to a more negative electrochemical potential
• Active transport
– Movement against electrochemical gradient
– From a more negative electrochemical potential
– to a more positive electrochemical potential
Electrochemical potential versus
water potential
• Just like water potential, solutes alone must
follow the rules of the electrochemical
potential and move passively
• If this is not what the cell or plant tissue
needs, two components are required
somewhere to counteract this natural
tendency
– Energy
– Membrane transport proteins
Summary of membrane
transport
• Three types of membrane transporters enhance the
movement of solutes across plant cell membranes
– Channels – passive transport
– Carriers – passive transport
– Pumps- active transport
Simple diffusion
• Movement down the gradient in
electrochemical potential
• Movement between phospholipid
bilayer components
• Bidirectional if gradient
changes
• Slow process
Channels
• Transmembrane proteins that
work as selective pores
– Transport through these
passive
• The size of the pore determines
its transport specifity
• Movement down the gradient in
electrochemical potential
• Unidirectional
• Very fast transport
• Limited to ions and water
Channels
• Sometimes channel transport
involves transient binding of the
solute to the channel protein
• Channel proteins have
structures called gates.
– Open and close pore in
response to signals
• Light
• Hormone binding
• Only potassium can diffuse
either inward or outward
– All others must be expelled
by active transport.
Remember the aquaporin channel
protein?
• There is some diffusion of
water directly across the bi-
lipid membrane.
• Aquaporins: Integral
membrane proteins that form
water selective channels –
allows water to diffuse faster
– Facilitates water
movement in plants
• Alters the rate of water flow
across the plant cell
membrane – NOT direction
Carriers
• Do not have pores that extend
completely across membrane
• Substance being transported is initially
bound to a specific site on the carrier
protein
– Carriers are specialized to carry a
specific organic compound
• Binding of a molecule causes the carrier
protein to change shape
– This exposes the molecule to the
solution on the other side of the
membrane
• Transport complete after dissociation
of molecule and carrier protein
Carriers
• Moderate speed
– Slower than in a channel
• Binding to carrier protein is
like enzyme binding site
action
• Can be either active or
passive
• Passive action is sometimes
called facilitated diffusion
• Unidirectional
Active transport
• To carry out active transport:
– The membrane transporter must couple the
uphill transport of a molecule with an energy
releasing event
• This is called Primary active transport
– Energy source can be
• The electron transport chain of mitochondria
• The electron transport chain of chloroplasts
• Absorption of light by the membrane transporter
• Such membrane transporters are called
PUMPS
Primary active transport-Pumps
• Movement against the
electrochemical gradient
• Unidirectional
• Very slow
• Significant interaction with
solute
• Direct energy expenditure
pump-mediated transport against the
gradient (secondary active transport)
• Involves the coupling of the
uphill transport of a
molecule with the downhill
transport of another
• (A) the initial conformation
allows a proton from outside
to bind to pump protein
• (B) Proton binding alters the
shape of the protein to allow
the molecule [S] to bind
pump-mediated transport against the
gradient (secondary active transport)
• (C) The binding of the
molecule [S] again alters
the shape of the pump
protein. This exposes the
both binding sites, and the
proton and molecule [S] to
the inside of the cell
• (D) This release restores
borh pump proteins to their
original conformation and
the cycle begins again
pump-mediated transport against the
gradient (secondary active transport)
• Two types:
• (A) Symport:
– Both substances move in
the same direction across
membrane
• (B) Antiport:
– Coupled transport in which
the downhill movement of a
proton drives the active
(uphill) movement of a
molecule
–
– In both cases this is
against the concentration
gradient of the molecule
(active)
pump-mediated transport against the
gradient (secondary active transport)
• The proton gradient
required for secondary
active transport is
provided by the activity of
the electrogenic pumps
• Membrane potential
contributes to secondary
active transport
• Passive transport with
respect to H+ (proton)
ABC transporters
• Also known as the (ATP-binding
cassette) superfamily.
• ABC transporters all have a
similar structure, consisting of
two ATP binding domains facing
the cytosol and two
transmembrane domains
• Similar to the situation seen with
ATP-driven ion pumps, the binding
and hydrolysis of ATP by ABC
transporters is thought to drive
conformational changes that
transport molecules across the
membrane.
Kretzschmar et al (2011). Biochemical Society
Essays Biochem. 50, 145–160
ABC transporters
• ABC transporters in Plant cells are
specialized for pumping small
compounds out of cells.
• In general, ABC transporters seem
to be crucial for getting foreign
substances (drugs and other toxins)
out of cells, making them extremely
important clinically
• ABC transporters shuttle
substrates as diverse as lipids,
phytohormones, carboxylates, heavy
metals, and chlorophyll catabolites
• ABC transporters participate in a
multitude of physiological processes
that allow the plant to adapt to
changing environments and cope
with biotic and abiotic stresses. Kretzschmar et al (2011). Biochemical Society
Essays Biochem. 50, 145–160
Overview of Ion homeostasis in
plant cells
The Vacuole
• Can be 80 – 90% of the plant
cell
• Contained within a vacuolar
membrane (Tonoplast)
• Contains:
– Water, inorganic ions,
organic acids, sugars,
enzymes, and secondary
metabolites.
• Required for plant cell
enlargement
• The turgor pressure
generated by vacuoles
provides the structural
rigidity needed to keep
herbaceous plants upright.
The Vacuole
In general, the functions of the vacuole include:
• Isolating materials that might be harmful
or a threat to the cell
• Containing waste products
• Containing water in plant cells
• Maintaining internal hydrostatic pressure
or turgor within the cell
• Maintaining an acidic internal pH
• Containing small molecules
• Exporting unwanted substances from the
cell
• Allows plants to support structures such
as leaves and flowers due to the pressure
of the central vacuole
• In seeds, stored proteins needed for
germination are kept in 'protein
bodies', which are modified vacuole
Ion homeostasis in plant cells
• Tonoplast antiporters
move sugars, ions and
contaminants to the
cytoplasm from the
vacuole
• Anion channels maintain
charge balance between
the cytoplasm and vacuole
• Ca channels work to
control second messenger
levels & cell signaling
paths between vacuole and
cytoplasm
Plasma membrane transporters
Plasma membrane transporters
Ion transport in roots
• As all plant cells are
surrounded by a cell wall,
Ions can be carried
through the cell wall space
with out entering an actual
cell
– The apoplast
• Just as the cell walls form
a continuous space, so do
the cytoplasms of
neighboring cells
– The symplast
Ion transport in roots
• All plant cells are connected
by plasmodesmata.
• In tissues where large
amounts of intercellular
transport occurs neighboring
cells have large numbers of
these.
– As in cells of the root tip
Ion transport in roots
• Ion absorption in the root is
more pronounced in the root
hair zone than other parts
of the root.
• An Ion can either enter the
root apoplast or symplast
but is finally forced into the
symplast by the casparian
strip.
Ion transport in roots
• Once the Ion is in the
symplast of the root it must
exit the symplast and enter
the xylem
– Called Xylem Loading.
• Ions are taken up into the
root by an active transport
process
• Ions are transported into
the xylem by passive
diffusion
Summary
• The movement of molecules and Ions from
one location to another is known as
transport.
• Plants exchange solutes and water with
their environment and among tissues and
organs
• Both local and long distance transport are
controlled by cellular membranes
Any Questions?

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SoluteTransport.ppt

  • 1. Plant Physiology talk Six Solute transport
  • 2. Solute transport • Plant cells separated from their environment by a thin plasma membrane (and the cell wall) • Must facilitate and continuously regulate the inward and outward traffic of selected molecules and ions as the cell – Takes up nutrients – Exports wastes – Regulates turgor pressure – Send chemical signals to other cells
  • 3. Two perspectives for membrane transport • Cellular level – Contribution to cellular functions – Contribution to ion homeostasis (i.e., balance) • Whole-plant level – Contribution to water relations – Contribution to mineral nutrition – Contribution to growth and development
  • 4. Moving into cells and between compartments requires membrane to be crossed • Composed of a phospholipid bilayer and proteins. • The phospholipid sets up the bilayer structure • Phospholipids have hydrophilic heads and fatty acid tails. • The plasma membrane is fluid--that is proteins move in a fluid lipid background
  • 5. Membrane potential • Arise because charged solutes cross membranes at different rates • Create a driving force for ionic transport • Maintained by energy- dependent electrogenic pumps
  • 6. Electrogenic pumps and membrane potential • Electrogenic pumps are ATPases (enzymes that split ATP) • ATPases use ATP energy to “pump” out protons (H+) to create charge gradients • H+ gradients create a type of “battery” to power transport and maintain ion homeostasis
  • 7. Electrogenic pumps and membrane potential • To prove this • Add cyanide (CN) – Rapidly poisons mitochondria, so cells ATP is depleted – Membrane potential falls to levels seen with diffusion • So membrane potential has too parts – Diffusion – Electrogenic ion transport • Requires energy
  • 8. Ion homeostasis within plant cells • Plant cells segregate ions based upon: – Function or role – Potential toxicity • This segregation creates a balance • Creating and maintaining the balance may require energy
  • 9. Ion homeostasis within plant cells • Ion concentrations in cytosol and vacuole are controlled by passive (dashed) and active (solid) transport processes • In most plant cells vacuole takes up 90% of the cell volume – Contains bulk of cells solutes • Control of cytosol ion concs is important for the regulations of enzyme activity • Cell wall is not a permeability barrier – It is NOT a factor in solute transport
  • 10. Passive vs active transport • Passive or active transport depends on the gradient in electrochemical potential • The electrochemical potential has 2 parts – Concentration – Charge (Electrical) • The two parts together dictate the electrochemical potential for a compartment of a cell
  • 11. Passive v. active transport • Passive transport – Movement down the electrochemical gradient – From a more positive electrochemical potential – to a more negative electrochemical potential • Active transport – Movement against electrochemical gradient – From a more negative electrochemical potential – to a more positive electrochemical potential
  • 12. Electrochemical potential versus water potential • Just like water potential, solutes alone must follow the rules of the electrochemical potential and move passively • If this is not what the cell or plant tissue needs, two components are required somewhere to counteract this natural tendency – Energy – Membrane transport proteins
  • 13. Summary of membrane transport • Three types of membrane transporters enhance the movement of solutes across plant cell membranes – Channels – passive transport – Carriers – passive transport – Pumps- active transport
  • 14. Simple diffusion • Movement down the gradient in electrochemical potential • Movement between phospholipid bilayer components • Bidirectional if gradient changes • Slow process
  • 15. Channels • Transmembrane proteins that work as selective pores – Transport through these passive • The size of the pore determines its transport specifity • Movement down the gradient in electrochemical potential • Unidirectional • Very fast transport • Limited to ions and water
  • 16. Channels • Sometimes channel transport involves transient binding of the solute to the channel protein • Channel proteins have structures called gates. – Open and close pore in response to signals • Light • Hormone binding • Only potassium can diffuse either inward or outward – All others must be expelled by active transport.
  • 17. Remember the aquaporin channel protein? • There is some diffusion of water directly across the bi- lipid membrane. • Aquaporins: Integral membrane proteins that form water selective channels – allows water to diffuse faster – Facilitates water movement in plants • Alters the rate of water flow across the plant cell membrane – NOT direction
  • 18. Carriers • Do not have pores that extend completely across membrane • Substance being transported is initially bound to a specific site on the carrier protein – Carriers are specialized to carry a specific organic compound • Binding of a molecule causes the carrier protein to change shape – This exposes the molecule to the solution on the other side of the membrane • Transport complete after dissociation of molecule and carrier protein
  • 19. Carriers • Moderate speed – Slower than in a channel • Binding to carrier protein is like enzyme binding site action • Can be either active or passive • Passive action is sometimes called facilitated diffusion • Unidirectional
  • 20. Active transport • To carry out active transport: – The membrane transporter must couple the uphill transport of a molecule with an energy releasing event • This is called Primary active transport – Energy source can be • The electron transport chain of mitochondria • The electron transport chain of chloroplasts • Absorption of light by the membrane transporter • Such membrane transporters are called PUMPS
  • 21. Primary active transport-Pumps • Movement against the electrochemical gradient • Unidirectional • Very slow • Significant interaction with solute • Direct energy expenditure
  • 22. pump-mediated transport against the gradient (secondary active transport) • Involves the coupling of the uphill transport of a molecule with the downhill transport of another • (A) the initial conformation allows a proton from outside to bind to pump protein • (B) Proton binding alters the shape of the protein to allow the molecule [S] to bind
  • 23. pump-mediated transport against the gradient (secondary active transport) • (C) The binding of the molecule [S] again alters the shape of the pump protein. This exposes the both binding sites, and the proton and molecule [S] to the inside of the cell • (D) This release restores borh pump proteins to their original conformation and the cycle begins again
  • 24. pump-mediated transport against the gradient (secondary active transport) • Two types: • (A) Symport: – Both substances move in the same direction across membrane • (B) Antiport: – Coupled transport in which the downhill movement of a proton drives the active (uphill) movement of a molecule – – In both cases this is against the concentration gradient of the molecule (active)
  • 25. pump-mediated transport against the gradient (secondary active transport) • The proton gradient required for secondary active transport is provided by the activity of the electrogenic pumps • Membrane potential contributes to secondary active transport • Passive transport with respect to H+ (proton)
  • 26. ABC transporters • Also known as the (ATP-binding cassette) superfamily. • ABC transporters all have a similar structure, consisting of two ATP binding domains facing the cytosol and two transmembrane domains • Similar to the situation seen with ATP-driven ion pumps, the binding and hydrolysis of ATP by ABC transporters is thought to drive conformational changes that transport molecules across the membrane. Kretzschmar et al (2011). Biochemical Society Essays Biochem. 50, 145–160
  • 27. ABC transporters • ABC transporters in Plant cells are specialized for pumping small compounds out of cells. • In general, ABC transporters seem to be crucial for getting foreign substances (drugs and other toxins) out of cells, making them extremely important clinically • ABC transporters shuttle substrates as diverse as lipids, phytohormones, carboxylates, heavy metals, and chlorophyll catabolites • ABC transporters participate in a multitude of physiological processes that allow the plant to adapt to changing environments and cope with biotic and abiotic stresses. Kretzschmar et al (2011). Biochemical Society Essays Biochem. 50, 145–160
  • 28. Overview of Ion homeostasis in plant cells
  • 29. The Vacuole • Can be 80 – 90% of the plant cell • Contained within a vacuolar membrane (Tonoplast) • Contains: – Water, inorganic ions, organic acids, sugars, enzymes, and secondary metabolites. • Required for plant cell enlargement • The turgor pressure generated by vacuoles provides the structural rigidity needed to keep herbaceous plants upright.
  • 30. The Vacuole In general, the functions of the vacuole include: • Isolating materials that might be harmful or a threat to the cell • Containing waste products • Containing water in plant cells • Maintaining internal hydrostatic pressure or turgor within the cell • Maintaining an acidic internal pH • Containing small molecules • Exporting unwanted substances from the cell • Allows plants to support structures such as leaves and flowers due to the pressure of the central vacuole • In seeds, stored proteins needed for germination are kept in 'protein bodies', which are modified vacuole
  • 31. Ion homeostasis in plant cells • Tonoplast antiporters move sugars, ions and contaminants to the cytoplasm from the vacuole • Anion channels maintain charge balance between the cytoplasm and vacuole • Ca channels work to control second messenger levels & cell signaling paths between vacuole and cytoplasm
  • 34. Ion transport in roots • As all plant cells are surrounded by a cell wall, Ions can be carried through the cell wall space with out entering an actual cell – The apoplast • Just as the cell walls form a continuous space, so do the cytoplasms of neighboring cells – The symplast
  • 35. Ion transport in roots • All plant cells are connected by plasmodesmata. • In tissues where large amounts of intercellular transport occurs neighboring cells have large numbers of these. – As in cells of the root tip
  • 36. Ion transport in roots • Ion absorption in the root is more pronounced in the root hair zone than other parts of the root. • An Ion can either enter the root apoplast or symplast but is finally forced into the symplast by the casparian strip.
  • 37. Ion transport in roots • Once the Ion is in the symplast of the root it must exit the symplast and enter the xylem – Called Xylem Loading. • Ions are taken up into the root by an active transport process • Ions are transported into the xylem by passive diffusion
  • 38. Summary • The movement of molecules and Ions from one location to another is known as transport. • Plants exchange solutes and water with their environment and among tissues and organs • Both local and long distance transport are controlled by cellular membranes