Figure 37.
2 The uptake of nutrients by a plant: an overview
Xylem and Phloem Flow in Plants
1. Nutrient requirements of plants
2. Overview of flow in plants
3. Lateral (short-distance flow)
4. Long-distance flow in the xylem
5. Phloem flow
Table 37.1 Essential Nutrients in Plants
Diffusion vs. Bulk Flow
• Diffusion: Net movement down a
concentration gradient due to the random
motion of individual molecules. (Note:
solutes may move independently of
water.)
• Bulk flow: Movement of water and solutes
together due to a pressure gradient.
• Osmosis: movement of water in or out
What makes up the mass of
plants?
1. Water makes up most of the
weight of a living plant
2. Carbon (organic)
compounds make up most of
the dry weight of plants
Thus, plants get most of their
mass from water and air, not
from soil mineral nutrients
(inorganic ions).
Nutrients and Xylem Flow in Plants
1. Nutrient requirements of plants
2. Overview of flow in plants
3. Lateral (short-distance flow)
4. Long-distance flow in the xylem
• Physical forces drive the transport of
materials in plants over a range of distances
• Transport in vascular plants occurs on three
scales
– Transport of water and solutes by individual
cells, such as root hairs
– Short-distance transport of substances from
cell to cell at the levels of tissues and organs
– Long-distance transport within xylem and
phloem at the level of the whole plant
1
 Figure 36.2 An overview of transport in whole plants (Layer 4)
Selective Permeability of
Note:
Short-distance, or Membranes: A Review
“lateral,” flow of fluids
in plants (e.g. from cell
to cell, or into the root • The selective permeability of a plant cell’s
form the soil) happens
by diffusion plasma membrane
– Controls the movement of solutes into and out
Long-distance flow of the cell
(i.e. through the xylem
and phloem) can only • Specific transport proteins
happen by “bulk flow,” – Enable plant cells to maintain an internal
i.e. movement that is environment different from their surroundings
driven by pressure

The Central Role of Proton Pumps • Plant cells use energy stored in the proton
gradient and membrane potential to drive the
• Proton pumps in plant cells transport of many different solutes
– Create a hydrogen ion gradient that is a form
+
of potential energy that can be harnessed to CYTOPLASM
–
EXTRACELLULAR FLUID

do work –
+
K+
– Contribute to a voltage known as a membrane K+ Cations ( , for
example) are driven
+
potential
–
K+ K+ into the cell by the
membrane potential.

CYTOPLASM EXTRACELLULAR FLUID

– + K+

+ H+
ATP – K+
H+ K+
– + –
+
H+ K+
Proton pump generates
membrane potential
H+ H+ and H+ gradient. – Transport protein
+
H+ – + H+
H+ (a) Membrane potential and cation uptake
– +

• The “coattail” effect of cotransport

• In a mechanism called cotransport: a – Is also responsible for the uptake of the sugar
transport protein couples the passage of sucrose by plant cells
one solute to the passage of another – +
H+ H+ H+
S +
– + –
H+ H+ –
– – + NO 3 H+ Plant cells can
O3 – +
N H+ also accumulate a
– + H+
H+ S neutral solute,
Cell accumulates
NO3 –
H+ anions ( , for such as sucrose
H+
example) by – H+ ( S ), by
coupling their
H+
H+ transport to the H+ cotransporting
H+ inward diffusion down the
H+ H+
of H through a
+ S steep proton
– – S
S

NO 3 cotransporter. + H+ gradient.
–
– NO –
+ – +
NO 3 3

– + H+ S H+
NO – H+
3
H+ – +
+ H+ –
H+
(c) Contransport of a neutral solute

(b) Cotransport of anions

2
 Effects of Differences in Water
Potential • Water potential
– Is a measurement that combines the effects of
• To survive solute concentration and pressure
– Plants must balance water uptake and loss – Determines the direction of movement of water
• Osmosis • Water
– Determines the net uptake or water loss by a – Flows from regions of high water potential to
cell regions of low water potential
– Is affected by solute concentration and pressure

Water Potential Water Potential

• Water moves from areas of higher (more
positive) potential to lower (more negative)
• Pressure potential is created by physical
pressure on water (positive) or a
vacuum/sucking (negative)
• Solute potential is created by a higher
concentration of solutes (= lower
concentration of water)
• Water moves from areas of higher (more
positive) potential to lower (more negative)
• Pressure potential is created by physical
pressure on water (positive) or a
vacuum/sucking (negative)
• Solute potential is created by a higher
concentration of solutes (= lower
concentration of water)

Ψ= Ψs + Ψp
Water pot. = solute pot. + pressure pot.

Quantitative Analysis of Water • Application of physical pressure

Potential (a)
– Increases water potential

• The addition of 0.1 M

solution (b) (c)

solutes
– Reduces water
potential
Pure
water

H 2O

H 2O
ψP = 0 H 2O
ψS = −0.23
ψ = 0 MPa ψ = −0.23 MPa
ψP = 0.23 ψP = 0.30
ψS = −0.23 ψS = −0.23
ψ = 0 MPa ψ = 0 MPa ψ = 0 MPa ψ = 0.07 MPa

3
 • Negative (d)
• Water potential
pressure – Affects uptake and loss of water by plant cells
(suction) • If a flaccid cell is placed in an environment
with a higher solute concentration
– Decreases – The cell will lose water and become plasmolyzed
water potential – Cellular potential is greater than environmental
potential Initial flaccid cell:
ψ = 0 P
ψS = −0.7

Water flow in plants is 0.4 M sucrose solution:

ψP = 0
ψ = −0.7 MPa

controlled by both of ψS = −0.9

these forces. For Plasmolyzed cell

ψ = −0.9 MPa
H2 O
example: potassium at osmotic equilibrium
with its surroundings
concentration and
ψP = 0
pressure from the cell ψP = −0.30 ψP = 0 ψS = −0.9
ψS = 0 ψS = −0.23
wall ψ = −0.30 MPa ψ = −0.23 MPa ψ = −0.9 MPa

• If the same flaccid cell is placed in a Transport is also regulated by the

solution with a lower solute concentration compartmental structure of plant cells
– The cell will gain water and become turgid

• The plasma membrane

Initial flaccid cell:
ψP = 0 – Directly controls the traffic of molecules into
ψS = −0.7

ψ = −0.7 MPa Distilled water: and out of the protoplast

ψP = 0
ψS = 0 – Is a barrier between two major compartments,
ψ = 0 MPa
Turgid cell the cell wall and the cytosol
at osmotic equilibrium
with its surroundings
– Aquaporins are transport proteins that
facilitate water movement across membranes
ψP = 0.7
ψS = −0.7

ψ = −0 MPa

• A major compartment in most mature plant cells is the

vacuole, a large organelle that can occupy as much as Nutrients and Xylem Flow in Plants
90% of more of the protoplast’s volume
• The vacuolar membrane
– Regulates transport between the cytosol and the
1. Nutrient requirements of plants
vacuole
Cell wall

Transport proteins in
the plasma membrane Cytosol Transport proteins in
2. Overview of flow in plants
regulate traffic of the vacuolar
molecules between membrane regulate
the cytosol and the Vacuole traffic of molecules
cell wall. between the cytosol
and the vacuole.
3. Lateral (short-distance flow)

4. Long-distance flow in the xylem

Vacuolar membrane
(tonoplast)
Plasmodesma Plasma membrane

(a)
Cell compartments. The cell wall, cytosol, and vacuole are the three main
compartments of most mature plant cells.

4
 Short distance or lateral transport: movement within Key
• In most plant tissues and organs radially:
3 routes Symplast
tissues Apoplast
– The cell walls and 1. Transmembrane route
cytosol are
continuous from
cell to cell The symplast is the
Apoplast

• The cytoplasmic continuum of

cytosol connected Symplast The apoplast is
continuum by plasmodesmata. the continuum
of cell walls and
– Is called the extracellular
spaces.
symplast
• The apoplast 2. Symplastic route
– Is the continuum 3. Apoplastic route
of cell walls plus
extracellular (b) Transport routes between cells. At the tissue level, there are three passages:

spaces the transmembrane, symplastic, and apoplastic routes. Substances may transfer
from one route to another.

Bulk Flow in Long-Distance

• Water and minerals can travel through a
plant by one of three routes
Transport
• In bulk flow
– Out of one cell, across a cell wall, and into
– Movement of fluid in the xylem and phloem is driven
another cell (transmembrane route) by pressure differences at opposite ends of the xylem
– Via the symplast vessels and sieve tubes
– Diffusion does not work well over long distances, such
– Along the apoplast as from roots to leaves
– In xylem it is negative pressure that drives flow
(transpiration)
– In phloem it is hydrostatic pressure in one end of the
sieve tube that forces sap to the opposite end

The Roles of Root Hairs,

Xylem transport: Root uptake
• Roots absorb water and minerals from the
soil
Mycorrhizae, and Cortical Cells
• Much of the absorption of water and minerals occurs near root
tips, where the epidermis is permeable to water and where root
hairs are located

• These enter the plant through the • Root hairs account for much of the surface area of roots, and
epidermis of roots and ultimately flow to greatly enhance absorption
the shoot system through the xylerm
• Root hairs are extensions of epidermal cells

• The soil solution flows into the hydrophilic cell walls, along the
apoplast and into the root cortex

5

• Most plants form mutually beneficial relationships
with fungi, which facilitate the absorption of water
and minerals from the soil
• Roots and fungi form mycorrhizae, symbiotic
structures consisting of plant roots united with
fungal hyphae
2.5 mm
Lateral transport in roots
• Water can cross the cortex
– Via the symplast or apoplast
• The waxy Casparian strip of the endodermal
wall
– Blocks apoplastic transfer of minerals from the
cortex to the vascular cylinder
Nutrients and Xylem Flow in Plants
1. Nutrient requirements of plants
2. Overview of flow in plants
3. Lateral (short-distance flow)
4. Long-distance flow in the xylem
Lateral transport of minerals and water in roots
Note the important role of the
Casparian strip in the endodermis:
blocking the apoplastic pathway into
the stele. Why is this useful to the
plant?
Lateral transport in roots
• Once soil solution enters the roots
– The extensive surface area of cortical cell
membranes enhances uptake of water and selected
minerals
• The endodermis is a selective sentry
– It is the innermost layer of cells in the root cortex
– Surrounds the vascular cylinder and functions as the
last checkpoint for the selective passage of minerals
from the cortex into the vascular tissue (via the waxy
casparian strip)
– All material must pass via the symplast
Ascent of water in a tree
(long-distance flow). Water
is pulled upward by
extreme negative water
potential generated by
evaporation out of stomata
(transpiration).
6
 Figure 36.12 The generation of transpirational pull in a leaf
Cohesion and Adhesion in the
Ascent of Xylem Sap
• The transpirational pull on xylem sap
– Is transmitted all the way from the leaves to the root
tips and even into the soil solution
– Is facilitated by cohesion (water molecules to one
another via their polar bonds) and adhesion (to the
hydrophilic vessel walls)
– Small diameter of vessels and tracheids increases
adhesive surface
• The movement of xylem sap against gravity
– Is maintained by the transpiration-cohesion-
tension mechanism

At night, some xylem sap is also pushed up by root

pressure: roots pump ions in, and water follows,
creating high water potential in roots. Guttation (see
– Stomata increase below) is one consequence of this.
photosynthesis
– Stomata increase water loss However, root pressure cannot push very much xylem
sap. Most upward movement of xylem sap is due to
– Closing them reduces transpirational pull, not push by root pressure.
photosynthesis and can lead
to overheating in plants

20 µm

• Phloem sap
Phloem flow – Is an aqueous solution that is mostly sucrose
• Products of photosynthesis, organic nutrients – Travels from a sugar source to a sugar sink
(sugars), are translocated through the phloem – Also carries minerals, amino acids and
hormones
• In angiosperms the specialized cells are called
sieve tube members (with companion cells) • A sugar source
– Is a plant organ that is a net producer of sugar,
• In gymnosperms these are sieve cells (with such as mature leaves
albuminous cells) • A sugar sink
– Is an organ that is a net consumer or storer of
sugar, such as a tuber or bulb

7
 Loading of sucrose into phloem: Notice that it can follow either the symplastic
• Phloem sap moves from sugar or apoplastic pathway. Loading from the latter requires active transport.
source to sugar sink Loading occurs in sugar sources, such as mature leaves.

• Sugar must be loaded into

sieve-tube members before
being exposed to sinks
• In many plant species, sugar
moves by symplastic and
apoplastic pathways Mesophyll cell
Companion
(transfer) cell
Sieve-tube
member
Cell walls (apoplast)
Plasma membrane
Plasmodesmata

Sucrose manufactured in mesophyll cells

can travel via the symplast (blue arrows)
to sieve-tube members. In some species,
sucrose exits the symplast (red arrow)
near sieve tubes and is actively
accumulated from the apoplast by sieve- Bundle- Phloem
parenchyma cell
tube members and their Mesophyll cell sheath cell

companion cells.

Pressure flow in a sieve tube of

• In many plants the phloem is from sugar sources
to sugar sinks.
– Phloem loading requires active transport
Sugar loading at sources and
• Proton pumping and cotransport of unloading at sinks creates a
pressure gradient that drives bulk
sucrose and H+ flow of the phloem sap.

– Enable the cells to accumulate sucrose Sources: Mature leaves, other

photosynthetic organs, storage
organs (such as roots)
High H+ concentration Cotransporter

H+ Sinks: Growing shoots, flowers,

Proton
pump S
developing fruits, root tips

Key

ATP Sucrose
H+ H+ Apoplast
S
Low H+ concentration
Symplast

Tapping phloem sap with the help of an aphid, which is a phloem-feeder. An

aphid stylus, minus the aphid, functions as a phloem sap pressure probe
(right). Things to remember about
Sieve tube member water potential
• Water will diffuse from areas of low solute
concentration (high potential) to areas of
Stylet high solute concentration (low potential)
• Water will move from areas of high
pressure to areas of low pressure by bulk
flow
• Cell membranes are selective in the
Attach stylus near mature leaves, and another near base of plant. solutes allowed to pass through, some
Where would you predict sap pressure to be higher? solutes are actively pumped