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Transcript
Chapter 36
Transport in Vascular Plants
A. Physical Forces
CO2
H2O
O2
light
sugar
O2
H2O
minerals
CO2
A. Physical Forces

major substances transported are:

H2O and minerals
 transport in xylem



transpiration moves water because of
evaporation, cohesion and adhesion
sugars

transport in phloem

bulk flow
gas exchange

transport occurs on three scales
 cellular


from environment into
plant cells
transport of H2O and solutes
into root hairs

short-distance transport


from cell to cell

loading of sugar from
photosynthetic leaves into
phloem sieve tubes
long-distance transport

transport in xylem and phloem
throughout whole plant

membranes
 selective permeability

diffusion, passive transport, active transport

phospholipid bilayer, protein channels
Cellular Transport
 solutes are moved into plant cells by active transport

proton pumps
 active transport protein




in cell membrane
chemiosmosis – mechanism that uses the
energy stored in a concentration gradient to
drive cellular work
use ATP to pump H+ (hydrogen) ions
against the concentration gradient out of
the cell
sets up membrane potential – a separation of
opposite charge across a membrane
The Proton Pump

both the proton pump and membrane potential
have stored energy
which is used to drive the
transport of many different solutes
Water Potential
 water uptake and loss must be balanced


water moves by osmosis
cell walls add physical pressure which
affects osmosis
water potential, Ψ , takes both solute (dissolved
substances) concentration and physical
into account
pressure
 measured in megapascals, MPa (or bars)

Ψ = ΨS + ΨP
where:

Ψ = water potential
ΨS = solute potential (osmotic potential)
ΨP = pressure potential
the ΨS of pure water is zero
Pure
water
 = 0 MPa

adding solute lowers
the water potential
(because there is less free
water molecules less
capacity to do work) and
ΨS is negative
Addition of
solutes
0.1 M
solution
Pure
water
H2O
 = 0 MPa
P = 0
S = –0.23
 = –0.23 MPa

ΨP can be positive or negative
atmospheric pressure
Applying
physical
pressure
Pure
water
relative to
Applying
physical
pressure

H2O
P = 0.23
S = –0.23
 = 0 MPa
 = 0 MPa
Pure
water
H2O
 = 0 MPa
P = 0.30
S = –0.23
 = 0.07 MPa

water under tension (pulling) gives negative
pressure eg) water in xylem
Negative
pressure
Pure
water
P = –0.30
S = 0
 = –0.30 MPa
H2O
P = 0
S = –0.23
 = –0.23 MPa

water pushing out gives
eg) turgor pressure

water always moves from areas of high Ψ to areas of
low Ψ

water moves through the phospholipids bilayer and
through transport proteins called aquaporins

cells will be plasmolyzed or turgid depending on
the environment
plasmolyzed
positive pressure
turgid

loss of turgor causes wilting
Short-Distance Transport
 plant cells are compartmentalized
 cell wall

cell membrane – cytosol

vacuole
Cell wall
Cytosol
Vacuole
Plasmodesma
Vacuolar membrane
(tonoplast)
Plasma membrane

transport routes for water and solutes
 transmembrane route

repeated crossing
Transmembrane route
of plasma membrane
symplast route



movement within cytosol
plasmodesmata junctions connect cytosol of
neighboring cells
Key
Symplast
Transmembrane route
Symplast
Symplastic route
apoplast route



movement through the continuum of cell walls from
cell to cell
no cell membranes are crossed
Key
Symplast
Apoplast
Transmembrane route
Apoplast
Symplast
Symplastic route
Apoplastic route
Long-Distance Transport
 bulk flow which is the movement of fluid driven
by pressure
 flow in xylem tracheids and
vessels



transpiration creates negative
pressure which pulls xylem
sap upwards from roots
loading of sugar from
photosynthetic leaf cells
generates high positive pressure
which pushes phloem sap
through sieve tubes
lack of some organelles in phloem cells and the complete
lack of cytoplasm in xylem cells makes them very
efficient tubes for transport
B. Roots
 much of the absorption of water and minerals
takes place at the root tips

root hairs
 extensions of epidermal cells

walls are
hydrophilic

huge amount of surface area

soil solution moves into apoplast

flows through walls into cortex

solution moves into symplast
of root cells

water moves from high Ψ in
soil to low Ψ in root

active transport concentrates
certain molecules in the root
cells
eg) K+ ions

mycorrhizae
 symbiotic structures



plant roots with fungus
greatly increases
surface area for water
and mineral absorption
greatly increases
volume of soil reached
by plant

endodermis
 layer surrounding vascular cylinder
of root
 lined with impervious
Casparian strip
 forces solution through selective cell
membrane and into symplast



also prevents leakage of xylem sap
back into soil
solution in endodermis and parenchyma cells is
discharged into cell walls (apoplast) by
active and passive transport
this allows the solution to then move to the xylem
cells
Pathway along
apoplast
Casparian strip
Endodermal cell
Pathway
through
symplast
Casparian strip
Plasma
membrane
Apoplastic
route
Symplastic
route
Vessels
(xylem)
Root
hair
Epidermis
Endodermis Vascular cylinder
Cortex
C. Ascent of Xylem Sap
 root pressure
 mineral ions in xylem of roots lowers
the Ψ




water flows in
causing root pressure
positive pressure
upward push of xylem sap
accounts for very small part of ascent of
sap

transpiration pull
 generated by leaf




solar
powered
Ψ in leaf is higher than Ψ in atmosphere
water vapour leaves the leaf through the
stomata (transpiration)
water
pulled up

Ψ is high in roots and low in leaves, moves water
up plant

adhesion, cohesion, hydrogen bonding
Xylem
sap
Outside air Ψ
= –100.0 MPa
Mesophyll cells
Stoma
Leaf Ψ(air spaces)
= –7.0 MPa
Trunk xylem Ψ
= –0.8 Mpa
Water potential gradient
Leaf Ψ(cell walls)
= –1.0 MPa
Water molecule
Transpiration Atmosphere
Xylem Adhesion
cells
Cell wall
Cohesion, by hydrogen
bonding
Cohesion and
adhesion in
the xylem
Water molecule
Root xylem Ψ
= –0.6 MPa
Soil Ψ
= –0.3 MPa
Root hair
Soil particle
Water
Water uptake from soil
D. Stomata
 photosynthesis and transpiration
 compromise
CO2

CO2 in and O2 out but also H2O out

leaf transpires more than its weight in a day

xylem sap can flow at 75 cm/min
O2,
H2O

H2O evaporation takes place even with closed stomata

drought will cause wilting

transpiration causes evaporative cooling of the
leaves

regulation of stomata
 microfibril mechanism

guard cells attached at tips

contain microfibrils in cell walls

guard cells elongate and bow out when turgid

guard cells shorten and become less bowed when flaccid
Cells turgid/Stoma open
Cells flaccid/Stoma closed
Radially oriented
cellulose microfibrils
Cell wall
Vacuole
Guard cell

ion mechanism

proton pumps are used to move K+ ions
cells (stored in vacuoles)

Ψ in cells lower than surrounding cells H2O
moves in

guard cells become turgid and open

loss of K+ ions causes H2O to move out of guard
cells
become flaccid and close

into guard
Cells turgid/Stoma open Cells flaccid/Stoma closed
H 2O
H 2O
H 2O
H 2O
H2O
+
H 2O
K
H 2O
H 2O
H 2O
H 2O

other cues
 light



blue-light receptors in plasma membrane
triggers ATP-powered proton pumps causing
K+ uptake
stomata open
depletion of CO2

CO2 in air spaces in mesophyll is used for
photosynthesis

depletion causes stomata to open

circadian rhythm

automatic 24-hour cycle

stomata open in day, close at night

xerophytes
 plants adapted for
arid regions

adapted to reduce water loss
 small, thick leaves
 reflective leaves
 hairy leaves
 stomata in pores on underside of leaves
 alternative photosynthetic pathway (CAM)
E. Organic Nutrients
 translocation is the transport of organic nutrients
 phloem sap contains:
 water

sugar (sucrose) (30% by weight)
minerals

amino acids

hormones


sieve tubes carry sap from sugar source (leaves)
to sugar sink (growing roots, buds, stems and
fruit) variable direction of flow

sap flow rate can be as high as 1 m/hr

sugars are loaded into the phloem
 flow through symplast
via plasmodesmata

active cotransport of sucrose into phloem
cells with H+ ions in proton pump
Key
Apoplast
Symplast
Cotransporter
Mesophyll cell
Cell walls (apoplast)
Plasma membrane
High H+ concentration
Companion
Sieve-tube
(transfer) cell
member
Proton
pump
Plasmodesmata
Sucrose
Mesophyll cell
Bundlesheath cell
Phloem
parenchyma cell
Low H+ concentration

pressure flow





Ψ in phloem is lower than in the xylem at
sugar source because of the sugar loading
that takes place
H2O diffuses from xylem into phloem
positive pressure is generated which
causes the sap to move through phloem
sieve tubes
Ψ in phloem is higher than in the
xylem at sugar sinks
because of the
sugar being removed from the phloem
H2O diffuses back into xylem from
phloem
Vessel
(xylem)
high Ψ
low Ψ
Sieve tube
(phloem)
low Ψ
H2O
H2O
Sucrose Source cell
(leaf)
H2O
Sink cell
(storage
Sucrose
root)
high Ψ