Roots, Soils, and Nutrient

Uptake
Chapter 3
 Learning objectives
Understand the properties of soils and
how these properties dictate the
availability of nutrients to plants

Understand the mechanisms by which

solutes are transported across
membranes, including the role of
membrane transport proteins
 Learning objectives
Understand how solute ions move through
root tissues and cell walls

Understand how soil microorganisms

assist plants in acquiring nutrients for
uptake.
 Nutrients in the soil
Soil colloids, clay particles suspended in
but not dissolved in the soil solution, are
the main reservoir of soil nutrients.

Soil colloids are distinguished from other

soil constituents (e.g., silt and sand), by
the Tyndall effect.

Colloidal suspensions form micelles.

 Nutrients in the soil
Some soils also contain colloidal organic
material called humus.

The role of the colloidal fraction as a

nutrient reservoir depends upon:
Specific surface area.
The density of fixed negative charges.
 Nutrients in the soil
Figure 3.1
 Nutrients in the soil
The ability of colloidal surfaces to bind soil
cations follows the lyotropic series.

Al3+ > H+ > Ca2+ > Mg2+ > K+ = NH4+ > Na+

Cation binding to colloidal surfaces is

reversible, so nutrient ions in the soil show
ion exchange, or exchangability.
 Nutrients in the soil
Colloidal binding contributes to the
dynamic equilibrium of cationic nutrients in
the soil.

Soils do not generally bind anionic

nutrients.

Anionic nutrients may have to be supplied

in order to meet plant growth requirements.
 Nutrients in the soil
Figure 3.2
 Nutrient uptake
In order for nutrients to enter plants, they
must cross cell membranes.

The mechanisms by which nutrients cross

membranes include:
Simple diffusion
Facilitated diffusion
Active transport
 Nutrient uptake
Figure 3.3
 Simple diffusion
Simple diffusion is governed by Ficks law.
o i
J = PA(C -C )
J is the flux.
P is the permeability coefficient of the
membrane.
A is the cross-sectional area.
(Co-Ci) represents the concentration
gradient.
 Simple diffusion
Since membranes are made of lipids,
nonpolar substances can diffuse more
readily into cells than nutrient ions.

The permeability coefficient is a measure of

how readily a given solute can diffuse
through a membrane.

Diffusion is a passive process in response

to a concentration gradient.
 Membrane transport proteins
Facilitated diffusion describes the rapid,
passive, protein-assisted diffusion of
solutes across membranes.

Facilitated diffusion can be mediated by:

Carrier proteins
Channel proteins
 Membrane transport proteins
Pumps are membrane transport proteins
that move solutes against their gradient.

Pumps require the input of energy to

achieve transport, so this is referred to as
active transport.
 Selective accumulation of ions
Active transport allows for the
accumulation of ions inside cells to
concentrations higher than those outside
the cells.

This corresponds to an accumulation

ratio (ratio of plant concentration to soil
concentration) >1.0.
 Electrochemical gradients dictate
ion movement
For uncharged solutes, the concentration
gradient can be expressed in terms of a
gradient of chemical potential ().
[U i ] [U i ]
u = RT ln = 59 log
[U o ] [U o ]

The charge of ions creates an electrical

potential that must also be considered.
 Electrochemical gradients dictate
ion movement
Hence, ion transport is dictated by the
electrochemical gradient across the cell
membrane.

Because the electrochemical gradient

influences charge as well as concentration
gradients of ions, cells have a
transmembrane potential (a net negative
charge) across their membranes.
 Electrochemical gradients dictate
ion movement
Figure 3.8
 The Nernst equation
When the charge and concentration
gradients are taken into account, it is
possible to estimate whether an ion moves
by passive or active transport.

The equation used is the Nernst equation.

[Ci ]
- z E c = 59 log
[Co ]
 The Nernst equation
The equation predicts the Nernst
potential (Ec) that would be established if
an ion with charge z came to equilibrium
due to simple diffusion.

If the observed values deviate from those

predicted by the model, this suggests that
there is a protein-mediated active uptake
or efflux of the ion of interest.
 Electrogenic pumps
The energy for active transport comes
from two sources.
The direct use of ATP to transport a solute
across the membrane.
The use of an ATPase proton pump to
establish a proton motive force.
The ATPase is a uniport that is
electrogenic, in that it maintains a charge
gradient across the membrane.(+ outside)
 Electrogenic pumps
Proton motive force is responsible for
activities such as
active transport of solutes (cations,
anions, amino acids, and sugars),
regulation of cytoplasmic pH,
stomatal opening and closure,
sucrose transport during phloem loading,
hormone-mediated cell elongation.
 Electrogenic pumps
Figure 3.9
 Electrogenic pumps
Figure 3.10
 Electrogenic pumps
Solutes coupled to the proton motive force
(cotransport) can be moved against their
concentration gradient.
Symports move the proton and the ion of
interest in the same direction across the
membrane.
Antiports move the proton and the ion of
interest in opposite directions across the
membrane.
 Electrogenic pumps
There are electrogenic ATPases in both
the plasma membrane and the tonoplast.

The tonoplast ATPases (V-type) differ from

those in the plasma membrane.
This ATPase is not inhibited by vanadate as
the plasma membrane ATPase is.
This ATPase is inhibited by oligomycin.
The V-type ATPase is more similar to the
mitochondrial F-type ATPase.
 Electrogenic pumps
In addition to facilitating ion transport, the
V-type ATPase helps to maintain pH
balance between the vacuole and cytosol.

There are also Ca2+-ATPases that use ATP

to pump calcium against its concentration
gradient to store this ion in specific
subcellular compartments.
 K transport
+

Like other ions, K+ transport is mediated

by two classes of transport proteins.
Low affinity transporters, such as membrane
channels, are used when K+ concentrations
outside the cell are high.
High affinity transporters, such as carriers
energized by the proton motive force, are
used when K+ concentration outside the cell
are low.
 Ion transport interactions
The transport of various ions interact with
one another.
Deficiencies of one nutrient can increase the
uptake of other elements.
Some nutrients compete with one another for
uptake.

Ion transport interactions such as these

are examples of cross talk.
 Transport across the cell wall
Before ions can be transported across the
cell membrane, they must cross the cell
wall.

The cell wall has an apparent free space

(APS) that retains soil solution and ions.

The APS is an apoplastic compartment.

 Transport across the cell wall
Ions in the APS can move through the
APS to the cell membranes, where they
can then be taken up.

Ions in the APS can also diffuse back out

of the APS because they may be unbound
or on exchangeable sites in the cell wall.
 Transport across the cell wall
Figure 3.11
 Radial transport
Movement through the APS begins at the
rhizodermis (root epidermis).

Ions move radially through the cortex via:

The apoplastic pathway through the APS.
A transmembrane pathway across cortex cell
membranes.

Ion are transported in an energy-dependent

manner across the endodermis into the stele.
 Radial transport
Figure 3.12
 Radial transport
Passage cells in the endodermis also
allow for the transport of ions.

Lateral roots, emerging from the

meristematic pericycle layer inside the
endodermis, give rise to lateral roots
which also transport ions.
 Root-microbe interactions
The rhizosphere is the zone of soil
influenced by root biological processes.

The various exudates released by roots

provided a habitat for soil microorganisms.

The mixture of root exudates, living and

dead microorganisms, and soil colloids
forms the mucigel.
 Root-microbe interactions
These microorganisms aid the root in
obtaining nutrient ions.
Some bacteria convert nitrogen in the soil to
ammonium and nitrate.
Other bacteria can stimulate the formation of
proteoid roots specialized for the uptake of
nutrients such as phosphorus.
 Root-microbe interactions
Mycorrhizal fungi form mutualistic
interactions with the roots of most plants.

There are two types of mycorrhizae.

Ectotrophic or ectomycorrhizal
Endotropic or endomycorrhizal

The ecotmycorrhizae form a Hartig net

that penetrates into the apoplastic space.
 Root-microbe interactions
Endomycorrhizae such as the vesicular-
arbuscular mycorrhiza form branched
arbuscles inside cells of the root cortex.

Both types of mycorrhizae increase the

surface area of roots and create a larger
nutrient depletion zone to provide the
plant with nutrients.
 Root-microbe interactions
Figure 3.13