Biology: Transport in Vascular Plants

BIOLOGY TENTH EDITION
Global Edition
Campbell • Reece • Urry • Cain • Wasserman • Minorsky • Jackson
36
Transport in
Vascular Plants
Lecture Presentation by
Nicole Tunbridge and
Kathleen Fitzpatrick
© 2015 Pearson Education Ltd

A Whole Lot of Shaking Going On
a) Plants have various adaptations that aid in the

acquisition of resources, including water, minerals,
carbon dioxide, and light
a)For example, Aspen leaves have a peculiar adaption
that causes their leaves to tremble even
in light wind

Figure 36.1

Figure 36.1a

a) The success of plants depends on their ability to
gather resources from their environment and
transport them to where they are needed

Concept 36.1: Adaptations for acquiring resources
were key steps in the evolution of vascular plants
a)The algal ancestors of land plants absorbed water,

minerals, and CO2 directly from the surrounding
water
b)Early nonvascular land plants lived in shallow
water and had aerial shoots
c) Natural selection favored taller plants with flat
appendages, multicellular branching roots, and
efficient transport

a)The evolution of xylem and phloem in land plants
made possible the long-distance transport of
water, minerals, and products of photosynthesis
b)Xylem transports water and minerals from roots
to shoots
c)Phloem transports photosynthetic products from
sources to sinks

Figure 36.2-3
CO2 O2
Sugar Light
H2O
H2O and O2
minerals CO2

a) Adaptations in each species represent compromises
between enhancing photosynthesis and minimizing
water loss

Shoot Architecture and Light Capture
a) Stems serve as conduits for water and nutrients and

as supporting structures for leaves
b) Shoot length and branching pattern affect light
capture
c) There is a trade-off between growing tall and
branching

a)There is generally a positive correlation between
water availability and leaf size
b)Phyllotaxy, the arrangement of leaves on a stem,
is a species-specific trait important for light
capture
c) Most angiosperms have alternate phyllotaxy with
leaves arranged in a spiral
d)The angle between leaves is 137.5 and likely
minimizes shading of lower leaves
Figure 36.3
24 32
42 29 40
16
11 19
21 3 27
34 8
6
13 14
26 Shoot 1
5 apical 22
meristem 9
Buds 18
10 2 4
31 17
23 7 12
15
20 25
28
1 mm

a)The depth of the canopy, the leafy portion of all
the plants in a community, affects the productivity
of each plant
b)Self-pruning, the shedding of lower shaded
leaves, occurs when they respire more than
photosynthesize

a) Light absorption is affected by the leaf area index, the
ratio of total upper leaf surface of a plant divided by
the surface area of land on which it grows

Figure 36.4
Ground area
covered by plant
Plant A Plant B
Leaf area = 40% Leaf area = 80%
of ground area of ground area
(leaf area index = 0.4) (leaf area index = 0.8)
a) Leaf orientation affects light absorption
b) In low-light conditions, horizontal leaves capture more
sunlight
c) In sunny conditions, vertical leaves are less damaged
by sun and allow light to reach lower leaves

Root Architecture and Acquisition of Water and
Minerals
a) Soil contains resources mined by the root system

b) Root growth can adjust to local conditions
a)For example, roots branch more in a pocket of high
nitrate than low nitrate
c) Roots are less competitive with other roots from the

same plant than with roots from different plants

a)Roots and the hyphae of soil fungi form
mutualistic associations called mycorrhizae
b)Mutualisms with fungi helped plants colonize land
c) Mycorrhizal fungi increase the surface area for
absorbing water and minerals, especially
phosphate

Concept 36.2: Different mechanisms transport
substances over short or long distances
a) There are two major pathways through plants

a)The apoplast
b)The symplast

The Apoplast and Symplast: Transport Continuums
a)The apoplast consists of everything external to

the plasma membrane
b)It includes cell walls, extracellular spaces, and the
interior of vessel elements and tracheids
c) The symplast consists of the cytosol of all the
living cells in a plant, as well as the
plasmodesmata

a) Three transport routes for water and solutes are
a)The apoplastic route, through cell walls and
extracellular spaces
b)The symplastic route, through the cytosol
c)The transmembrane route, across cell walls

Figure 36.5
Cell wall
Apoplastic route
Cytosol
Symplastic route
Transmembrane route
Key
Plasmodesma
Plasma membrane Apoplast
Symplast

Short-Distance Transport of Solutes Across Plasma
Membranes
a)Plasma membrane permeability controls short-

distance movement of substances
b)Both active and passive transport occur in plants
c) In plants, membrane potential is established
through pumping H by proton pumps
d)In animals, membrane potential is established
through pumping Na by sodium-potassium
pumps

Figure 36.6
CYTOPLASM EXTRACELLULAR − +
− + S H+ − H+
FLUID + H+
+H H+
ATP − + Hydrogen − + H+
H+ H+
− + ion S
H+ +
H
H+
H+ H+ H+
H+
H+ H+ S −
− + + H+
Proton pump −
H+ − + Sucrose
+ H+/sucrose
(a) H+ and membrane potential cotransporter − + (neutral solute)
(b) H+ and cotransport of neutral solutes
− +
H+ H+
− + − +
− + H+ K+ Potassium ion
H+ H+ − +
K+
H+ Nitrate K+ − +
H+ K+
H+
K+
− + K+ K+
− + H+ − +
H+/NO3− H+
H+ Ion channel − +
cotransporter − +
(c) H+ and cotransport of ions (d) Ion channels

a)Plant cells use the energy of H gradients to
cotransport other solutes by active transport
b)Plant cell membranes have ion channels that
allow only certain ions to pass

Short-Distance Transport of Water Across Plasma
Membranes
a)To survive, plants must balance water uptake and

loss
b)Osmosis is the diffusion of water into or out of a
cell that is affected by solute concentration and
pressure

a)Water potential is a measurement that combines
the effects of solute concentration and pressure
b)Water potential determines the direction of
movement of water
c) Water flows from regions of higher water potential
to regions of lower water potential
d)Potential refers to water’s capacity to perform
work

a)Water potential is abbreviated as  and
measured in a unit of pressure called the
megapascal (MPa)
b)   0 MPa for pure water at sea level and at room
temperature

How Solutes and Pressure Affect Water Potential
a) Both solute concentration and pressure affect water

potential
b) This is expressed by the water potential equation:  
S  P
c) The solute potential (S) of a solution is directly
proportional to its molarity
d) Solute potential is also called osmotic potential

a) Pressure potential (P) is the physical pressure on a
solution
b) Turgor pressure is the pressure exerted by the
plasma membrane against the cell wall, and the cell
wall against the protoplast
c) The protoplast is the living part of the cell, which
also includes the plasma membrane

Water Movement Across Plant Cell Membranes
a) Water potential affects uptake and loss of water by

plant cells
b) If a flaccid (limp) cell is placed in an environment
with a higher solute concentration, the cell will lose
water and undergo plasmolysis
c) Plasmolysis occurs when the protoplast shrinks and
pulls away from the cell wall

Figure 36.7
Environment
Initial flaccid cell: 0.4 M sucrose solution:
ψP = 0 ψP = 0
ψS = −0.7 ψS = −0.9
ψ = −0.7 MPa ψ = −0.9 MPa
Final plasmolyzed cell at osmotic

equilibrium with its surroundings:
ψP = 0
ψS = −0.9
ψ = −0.9 MPa
(a) Initial conditions: cellular ψ > environmental ψ
Environment
Initial flaccid cell: Pure water:
ψP = 0 ψP = 0
ψS = −0.7 ψS = 0
ψ = −0.7 MPa ψ = 0 MPa
Final turgid cell at osmotic

equilibrium with its surroundings:
ψP = 0
ψS = −0.7
ψ = 0 MPa
(b) Initial conditions: cellular ψ < environmental ψ
a)If a flaccid cell is placed in a solution with a lower
solute concentration, the cell will gain water and
become turgid
b)Turgor loss in plants causes wilting, which can
be reversed when the plant is watered

Figure 36.UN02
Turgid

Figure 36.UN03
Wilted

Aquaporins: Facilitating Diffusion of Water
a)Aquaporins are transport proteins in the cell

membrane that facilitate the passage of water
b)These affect the rate of water movement across
the membrane

Long-Distance Transport: The Role of Bulk Flow
a)Efficient long-distance transport of fluid requires

bulk flow, the movement of a fluid driven by
pressure
b)Water and solutes move together through
tracheids and vessel elements of xylem, and
sieve-tube elements of phloem
c) Efficient movement is possible because mature
tracheids and vessel elements have no
cytoplasm, and sieve-tube elements have few
organelles in their cytoplasm
Concept 36.3: Transpiration drives the transport of
water and minerals from roots to shoots via the xylem
a) Plants can move a large volume of water from their

roots to shoots

Absorption of Water and Minerals by Root Cells
a) Most water and mineral absorption occurs near root

tips, where root hairs are located and the epidermis is
permeable to water
b) Root hairs account for much of the surface area of
roots
c) After soil solution enters the roots, the extensive
surface area of cortical cell membranes enhances
uptake of water and selected minerals

a) The concentration of essential minerals is greater in
the roots than soil because of active transport

Transport of Water and Minerals into the Xylem
a)The endodermis is the innermost layer of cells in

the root cortex
b)It surrounds the vascular cylinder and is the last
checkpoint for selective passage of minerals from
the cortex into the vascular tissue

a)Water can cross the cortex via the symplast or
apoplast
b)The waxy Casparian strip of the endodermal
wall blocks apoplastic transfer of minerals from
the cortex to the vascular cylinder
c) Water and minerals in the apoplast must cross
the plasma membrane of an endodermal cell to
enter the vascular cylinder

Figure 36.8
Casparian strip
Endodermal cell
Pathway along
apoplast
4 Water
Pathway moves
through 5
upward
symplast in vascular
cylinder
Plasmodesmata
1 Apoplastic Casparian strip
route Plasma
membrane
1 Apoplastic
route
2 Symplastic
3
route 2 4 5 Vessels
(xylem)
Symplastic Root
route hair
3 Transmembrane
Epidermis Endodermis Vascular
route
cylinder
Cortex (stele)
4 The endodermis: controlled entry
to the vascular cylinder (stele) 5 Transport in the xylem
a) The endodermis regulates and transports needed
minerals from the soil into the xylem
b) Water and minerals move from the protoplasts of
endodermal cells into their own cell walls
c) Diffusion and active transport are involved in this
movement from symplast to apoplast
d) Water and minerals now enter the tracheids and
vessel elements

Bulk Flow Transport via the Xylem
a)Xylem sap, water and dissolved minerals, is

transported from roots to leaves by bulk flow
b)The transport of xylem sap involves
transpiration, the evaporation of water from a
plant’s surface
c) Transpired water is replaced as water travels up
from the roots
d)Is sap pushed up from the roots, or pulled up by
the leaves?
Pushing Xylem Sap: Root Pressure
a) At night root cells continue pumping mineral ions into

the xylem of the vascular cylinder, lowering the water
potential
b) Water flows in from the root cortex, generating root
pressure, a push of xylem sap
c) Root pressure sometimes results in guttation, the
exudation of water droplets on tips or edges of leaves

Figure 36.9

a) Positive root pressure is relatively weak and is a
minor mechanism of xylem bulk flow

Pulling Xylem Sap: The Cohesion-Tension Hypothesis
a)According to the cohesion-tension hypothesis,

transpiration and water cohesion pull water from
shoots to roots
b)Xylem sap is normally under negative pressure,
or tension

Transpirational Pull
a) Water vapor in the airspaces of a leaf diffuses down
its water potential gradient and exits the leaf via
stomata
b) As water evaporates, the air-water interface retreats
further into the mesophyll cell walls
c) The surface tension of water creates a negative
pressure potential

a) This negative pressure pulls water in the xylem into
the leaf
b) The transpirational pull on xylem sap is transmitted
from leaves to roots

Figure 36.10
4 Increased surface
5 Water from xylem pulled tension pulls
into cells and air spaces. water from cells
and air spaces.
Cuticle Xylem
Upper
epidermis
3 Air-water
interface
Microfibrils retreats.
in cell wall of
Mesophyll
Air mesophyll cell
space
2 Water vapor
Lower replaced
epidermis from water
Cuticle film.
Stoma
1 Water vapor diffuses outside Microfibril Water Air-water

via stomata. (cross section) film interface

Figure 36.11
Xylem sap
Outside air ψ Mesophyll cells
= −100.0 MPa Stoma
Water molecule
Leaf ψ (air spaces)
Atmosphere
= −7.0 MPa
Transpiration
Leaf ψ (cell walls)
= −1.0 MPa Xylem Adhesion by
Water potential gradient

cells hydrogen bonding
Cell
wall
Trunk xylem ψ Cohesion

= −0.8 MPa by hydrogen
Cohesion and bonding
adhesion in
the xylem
Water molecule
Root hair
Trunk xylem ψ
= −0.6 MPa Soil particle
Water
Soil ψ
Water uptake from soil
= −0.3 MPa
Adhesion and Cohesion in the Ascent of
Xylem Sap
a) Water molecules are attracted to cellulose in xylem
cell walls through adhesion
b) Adhesion of water molecules to xylem cell walls helps
offset the force of gravity

a) Water molecules are attracted to each other through
cohesion
b) Cohesion makes it possible to pull a column of xylem
sap
c) Thick secondary walls prevent vessel elements and
tracheids from collapsing under negative pressure
d) Drought stress or freezing can cause a break in the
chain of water molecules through cavitation, the
formation of a water vapor pocket

Xylem Sap Ascent by Bulk Flow: A Review
a)The movement of xylem sap against gravity is

maintained by the cohesion-tension mechanism
b)Bulk flow is driven by a water potential difference
at opposite ends of xylem tissue
c) Bulk flow is driven by transpiration and does not
require energy from the plant; like photosynthesis
it is solar powered

a) Bulk flow differs from diffusion
a)It is driven by differences in pressure potential, not
solute potential
b)It occurs in hollow dead cells, not across the
membranes of living cells
c)It moves the entire solution, not just water or solutes
d)It is much faster

Concept 36.4: The rate of transpiration is regulated by
stomata
a)Leaves generally have large surface areas and

high surface-to-volume ratios
b)These characteristics increase photosynthesis,
but also increase water loss through stomata
c) Guard cells help balance water conservation with
gas exchange for photosynthesis

Figure 36.12

Stomata: Major Pathways for Water Loss
a) About 95% of the water a plant loses escapes

through stomata
b) Each stoma is flanked by a pair of guard cells, which
control the diameter of the stoma by changing shape
c) Stomatal density is under genetic and environmental
control

Mechanisms of Stomatal Opening and Closing
a) Changes in turgor pressure open and close stomata

a)When turgid, guard cells bow outward and the pore
between them opens
b)When flaccid, guard cells become less bowed and the
pore closes

Figure 36.13
Guard cells turgid/ Guard cells flaccid/
Stoma open Stoma closed
Radially oriented
cellulose microfibrils
Cell
wall
Vacuole
Guard cell
(a) Changes in guard cell shape and stomatal
opening and closing (surface view)
H2O H2O H2O H2O
K+ H2O
H2O
H2O
H2O
H2O H2O
(b) Role of potassium ions (K+) in stomatal

opening and closing
a)This results primarily from the reversible uptake
and loss of potassium ions (K) by the guard cells

Stimuli for Stomatal Opening and Closing
a) Generally, stomata open during the day and close at

night to minimize water loss
b) Stomatal opening at dawn is triggered by
a)Light
b)CO2 depletion
c)An internal “clock” in guard cells
c) All eukaryotic organisms have internal clocks;

circadian rhythms are 24-hour cycles

a)Drought, high temperature, and wind can cause
stomata to close during the daytime
b)The hormone abscisic acid (ABA) is produced in
response to water deficiency and causes the
closure of stomata

Effects of Transpiration on Wilting and Leaf
Temperature
a) Plants lose a large amount of water by transpiration

b) If the lost water is not replaced by sufficient transport
of water, the plant will lose water and wilt
c) Transpiration also results in evaporative cooling,
which can lower the temperature of a leaf and prevent
denaturation of various enzymes involved in
photosynthesis and other metabolic processes

Adaptations That Reduce Evaporative Water Loss
a)Xerophytes are plants adapted to arid climates

b)Some desert plants complete their life cycle
during the rainy season
c) Others have fleshy stems that store water or leaf
modifications that reduce the rate of transpiration
d)Some plants use a specialized form of
photosynthesis called crassulacean acid
metabolism (CAM) where stomatal gas exchange
occurs at night
Figure 36.14
▶ Ocotillo ▼ Oleander (Nerium oleander)
(Fouquieria
splendens)
▼ Oleander (Nerium oleander)

Upper
Thick cuticle epidermal tissue
100 µm
Trichomes Stoma Lower epidermal
(“hairs”) Crypt tissue
► Old man cactus

(Cephalocereus
senilis)
Concept 36.5: Sugars are transported from sources to
sinks via the phloem
a)The products of photosynthesis are transported

through phloem by the process of translocation

Movement from Sugar Sources to Sugar Sinks
a) In angiosperms, sieve-tube elements are the conduits

for translocation
b) Phloem sap is an aqueous solution that is high in
sucrose
c) It travels from a sugar source to a sugar sink
d) A sugar source is an organ that is a net producer of
sugar, such as mature leaves
e) A sugar sink is an organ that is a net consumer or
storer of sugar, such as a tuber or bulb
a) A storage organ can be both a sugar sink in summer
and sugar source in winter
b) Sugar must be loaded into sieve-tube elements
before being exported to sinks
c) Depending on the species, sugar may move by
symplastic or both symplastic and apoplastic
pathways
d) Companion cells enhance solute movement between
the apoplast and symplast

Figure 36.15
Apoplast
Symplast
Companion
Mesophyll cell (transfer) cell High H+ concentration Cotransporter
Cell walls (apoplast) Proton H+
Plasma Sieve-tube
pump S
membrane element
Plasmodesmata
ATP Sucrose
Phloem H+ H+
Bundle- S
parenchyma cell Low H+ concentration
Mesophyll cell sheath cell
(a) Sucrose manufactured in mesophyll cells (b) A chemiosmotic mechanism is
can travel via the symplast (blue arrows) responsible for the active transport
to sieve-tube elements. of sucrose.

a)In many plants, phloem loading requires active
transport
b)Proton pumping and cotransport of sucrose and
H enable the cells to accumulate sucrose
c) At the sink, sugar molecules diffuse from the
phloem to sink tissues and are followed by water

Bulk Flow by Positive Pressure: The Mechanism of
Translocation in Angiosperms
a) Phloem sap moves through a sieve tube by bulk flow

driven by positive pressure called pressure flow

Figure 36.16
Sieve Source cell
Vessel tube (leaf)
(xylem) (phloem) 1 Loading of sugar
H2O 1 Sucrose
H2O
Bulk flow by negative pressure 2
Bulk flow by positive pressure

2 Uptake of water
3 Unloading of sugar
Sink cell
(storage
root)
4 Recycling of water
4 3
Sucrose
H2O

a) The pressure flow hypothesis explains why phloem
sap always flows from source to sink
b) Experiments have built a strong case for pressure
flow as the mechanism of translocation in
angiosperms
c) Self-thinning, the dropping of sugar sinks such as
flowers, seeds, or fruits, occurs when there are more
sugar sinks than the sources can support

Figure 36.17
25 µm
Sieve-
tube
element
Sap
droplet Stylet Sap droplet
Aphid feeding Stylet in sieve- Separated stylet

tube element exuding sap

Concept 36.6: The symplast is highly dynamic
a) The symplast is a living tissue and is responsible for

dynamic changes in plant transport processes

Changes in Plasmodesmatal Number and Pore Size
a) Plasmodesmata can open or close in response to

turgor pressure, cytoplasmic calcium levels, or
cytoplasmic pH
b) Plant viruses can cause plasmodesmata to dilate,
allowing viral RNA to pass between cells

Figure 36.18
Plasmodesma Cytoplasm
of cell 2
Virus
particles
Cytoplasm Cell walls

of cell 1
100 nm

Phloem: An Information Superhighway
a) Phloem is a “superhighway” for systemic transport of

macromolecules and viruses
b) Systemic communication through the phloem helps
integrate functions of the whole plant

Electrical Signaling in the Phloem
a) The phloem allows for rapid electrical communication

between widely separated organs
a)For example, rapid leaf movements in the sensitive
plant (Mimosa pudica)

Biology: Transport in Vascular Plants

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BIOLOGY TENTH EDITION

Campbell • Reece • Urry • Cain • Wasserman • Minorsky • Jackson

© 2015 Pearson Education Ltd

a) Plants have various adaptations that aid in the

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)The algal ancestors of land plants absorbed water,

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) Stems serve as conduits for water and nutrients and

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) Soil contains resources mined by the root system

c) Roots are less competitive with other roots from the

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) There are two major pathways through plants

© 2015 Pearson Education Ltd

a)The apoplast consists of everything external to

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)Plasma membrane permeability controls short-

© 2015 Pearson Education Ltd

(c) H+ and cotransport of ions (d) Ion channels

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)To survive, plants must balance water uptake and

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) Both solute concentration and pressure affect water

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) Water potential affects uptake and loss of water by

© 2015 Pearson Education Ltd

Final plasmolyzed cell at osmotic

(a) Initial conditions: cellular ψ > environmental ψ

Final turgid cell at osmotic

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)Aquaporins are transport proteins in the cell

© 2015 Pearson Education Ltd

a)Efficient long-distance transport of fluid requires

a) Plants can move a large volume of water from their

© 2015 Pearson Education Ltd

a) Most water and mineral absorption occurs near root

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)The endodermis is the innermost layer of cells in

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)Xylem sap, water and dissolved minerals, is

a) At night root cells continue pumping mineral ions into