Transport of Mineral Nutrients

Bhawna Madan

Contents

Transport of ions across cell membranes

Passive absorption
Electrochemical gradient
Donnans equilibrium
Facilitated diffusion
Accumulation against concentration gradient
Active absorption
Role of ATP, carrier systems
Role of cell membrane
Role of proton pump and ion flux.

Soil
Soil particle diameters
range over 6 orders of
magnitude
Coarse fragments > 2 mm
Sand < 2 mm to 0.05 mm
Silt < 0.05 mm to 0.002
mm
Clay < 0.002 m

Sand
Feels gritty
Considered non-cohesive does not stick
together in a mass unless it is very wet.
Sand has less nutrients for plants than smaller
particles
Voids between sand particles promote free
drainage and entry of air
Holds little water and prone to drought

Silt
<

0.05 mm to > 0.002 mm

Not visible without microscope
Quartz often dominant mineral in silt
since other minerals have weathered
away.
Does not feel gritty
Floury feel smooth like silly putty
Smaller particles retains more water
for plants and have slower drainage
than sand.
Easily washed away by flowing water
highly erosive.
Holds more plant nutrients than sand.

Clay
Pores spaces are very small
and convoluted
Movement of water and air
very slow

Water holding capacity

Tremendous capacity to
adsorb water- not all available
for plants.

Chemical adsorption is large

Clay
< 0.002 mm
Flat plates or tiny flakes
Small clay particles are colloids
If suspended in water will not settle
Wet clay is very sticky and is plastic or it can
be molded readily into a shape or rod.
Pores spaces are very small and convoluted
Movement of water and air very slow
Water holding capacity
Tremendous capacity to adsorb water- not
all available for plants.
Chemical adsorption is large

 What is Tyndall Effect?

Particles that are small enough to remain in suspension but too
large to go into solution are called colloids. A colloidal
suspension is a biphasic system. These suspended particles will
scatter the light, causing the path traversed by light to become
visible. This phenomenon is called Tyndall Effect. Clay and
humus form colloidal particles and exhibit this phenomenon.
Properties of Colloidal Particles
They have a large specific surface area.
They have a large no. of charges

 Soils have negative charge they are able to hold positively charged
cations.
The soil solution consists of a water layer around the soil particle and
water in the micropores.
There are cations attached to the colloids and in the soil solution.
Cation exchange takes place when one of the cations in the soil solution
replaces one of the cations an the soil colloid.
This exchange only takes place when the cations in the soil solution are
not in equilibrium to the cations on the soil colloid. This is almost
always the case

What is Cation Exchange Capacity of Soil?

the ability or capacity of a soil colloid to hold cations.
this is directly dependent on the amount of charge on the soil colloid.
More clay and organic matter content of soil, more the cation exchange
capacity of soil.
The amount of cations a soil can hold is dependent on:
how much clay is present.

type of clay that is present.

how much organic matter is present.
Hydrous Oxides of Fe and Al (Sesquioxides)
pH

Order of the strength of adsorption of cations

Al+3 > H+ > Ca+2 > Mg+ 2 > K+ >= NH4+> Na+
Anion Exchange Capacity of Soil Colloids is low
Soil Colloids are negatively charged so they have lower anion exchange
capacity

Lipid bilayers are selectively permeable

small,nonpolar
small
uncharged, polar
larger
uncharged, polar
molecules
ions

Size polarity - ions

Decreasing
permeability

The Permeability of the Lipid Bilayer

Hydrophobic molecules
Are lipid soluble and can pass through the membrane
rapidly
Polar molecules
Do not cross membrane rapidly
Ions
Do not cross the membrane at all

Transport processes
Solutes dissolved ions and small organic molecules
i.e., Na+,K+, H+, Ca++, Cl,- sugars, amino acids,
nucleotides

Three transport processes:

a. Simple diffusion directly thru membrane

b. Facilitated diffusion (passive transport)
c. Active transport requires energy
Req
Carrier
protein

Simple
Diffusion:

Tendancy of a material to spread out

Always moves toward equilibrium

Net diffusion
Figure 7.11 B
Net diffusion

Net diffusion
Net diffusion

Net diffusion

Equilibrium

Equilibrium
Net diffusion

Equilibrium

Ficks Law of Diffusion

According to Ficks law, the rate at which molecules in solution
diffuse from one region to another is a function of their
concentration difference,
F= PA (Co Ci)
Where F is the flux or amount of solute crossing the membrane
per unit time.
A= cross-sectional area of the diffusion path (Area of cell
membrane)
P= Permeability Coefficient (Velocity at which solute crosses the
membrane). It is specific for a particular membrane solute
combination. The permeability coefficient reflects the lipid
solubility of diffusing molecules.

Example: Glucose transporter GluT1 :

carrier-mediated facilitated diffusion
Glucoseout (HIGH)->glucose

(low)

outside cell
2. Conformational
3. Glucose
change
T2
ReleasedConformational
shift
2.

1. Glucose binds

T1

in

1.

inside cell

3.

Glucose + ATP glucose-6-phosphate + ADP

hexokinase

T1

Transmembrane transport proteins

allow selective transport of hydrophilic molecules & ions

aqueous channel
hydrophilic pore

2. channel protein

very rapid

EXTRACELLULAR
selective size/charge
FLUID

trap door

Channel protein

Solute
CYTOPLASM

(a) A channel protein (purple) has a channel through which

water molecules or a specific solute can pass.
Figure 7.15

Transport Rate
Carrier Proteins- 105- 106 molecules per second
Channel Proteins - 108 molecules per second

Kinetics of simple vs facilitated

Diffusion

(solute concentration gradient) ->

Gets
saturated
Maximum
rate

Does
Not
Get
saturated

For CHARGED solutes (ions): net driving force

is the electrochemical gradient
has both a concentration + charge component;
Ion gradients can create an electrical voltage
gradient across the membrane (membrane potential)

+ + +
+
+
+
+

-60 mVolts +

++ +
+ +
+
+++
+++

---

---

---

+++

+++

---

+ +
+ +

Carrier proteins: three types

(a) Uniport

(b) Co-transport

Uniport one solute transported

[Antiport two solutes in opposite directions

Symport two solutes in the same direction

Carrier Proteins can mediate either:

1. Passive transport
driving force ->

concentration/electrochemical gradient
OR

2. Active transport
against a gradient; unfavorable
requires energy input
Note: channel proteins mediate only passive transport

Active transport
Carrier protein moves solute AGAINST its concentration
gradient

Requires energy, usually in the form of ATP hydorlysis

Or a favorable gradient established by use of ATP
Active transport is unidirectional whereas passive transport is
bidirectional.
Active transport systems are also referred as pumps.

Active
transport:
Na+K+ Pump
(Na+K+ATPase)

3 Na+ out
2 K+ in
ATP!

Selective Accumulation of ions by roots

What is accumulation ratio?

The concentration of some ions inside the cell may reach levels
higher than their concentration in the surrounding medium. This
difference in ion concentration is called accumulation ratio. This is
also defined as ratio of concentration inside the cell to the
concentration outside the cell.
E.g., Internal concentration of K inside the cell is 1000 times greater
than outside the cell.
Accumultion ratio less than one implies that the solute is actively
extruded from the cell. E.g., low concentration of Na inside the
plant cells suggest that Na is actively extruded from the plant cell.

Chemical gradient and Electrochemical gradient

The transport of solutes is because of the chemical potential
gradient represented by following equation.
= i - o
Uncharged Solutes: Concentration gradient alone determines the
chemical potential gradient. In this case chemical potential is
represented by
= RT ln[Ui]/[Uo]
=59log[Ui]/[Uo]
Where Ui is the concentration of U inside the cytosol
Uo is the concentration of U outside the cell.
If Ui > Uo , then is positive, transport must occur by active
means
If Ui < Uo , then is negative, transport must occur by passive
means

Transport of Charged Solutes: Movement of solutes is in response

to electrochemical gradient (requires both electrical and chemical
component).
Donnans Equilibrium: According to this theory there are certain
pre-existing ions inside the cell which can not diffuse outside through
membrance. Such ions are called as in diffusible or fixed ions.
However, the membrane is permeable to both anions and cations of
the outer solution. For e.g., if K+ is moved outside the cell against the
concentration gradient, it leaves behind the non diffusible anion
charges, thus creating a charge imbalance and potential difference
across the membrane. This potential is referred to as Donnans
potential. The negative charges tend to pull K+ inside the cell. The
equilibrium is achieved when potential difference reaches a value
such that the concentration gradient that pulls K+ out of the cell is
balanced by the force of electrical gradient that pulls K+ inside the
cell.

Transport of Charged Solutes:

Nernst Equation

The membrane electrical potential at which the concentration

and electrical influences on a solutes movement are exactly
balanced, so there is no net movement.
It is used to predict the distribution of charged ion across the
membrane.
-z Ec = 59 log[Ci]/[Co]
Where z = valency or electric charge on the ion.
for an anion (z negative), Nernst predicts Co/Ci > 1
and for a cation (z positive), Nernst predicts Co/Ci <1
Ci and Co are the concentrations of ions inside and outside the cell
respectively.
Ec = Electrical potential difference (inside relative to outside),
and is negative for the plant plasmamembrane.

Nernst Equation can be used to predict difference between active

and passive transport
Compare the concentration gradient predicted by the Nernst
Equation to that actually observed.
If they differ, then active transport must be involved.

Electrogenic pumps are critical for cellular active transport

F-type ATPases: It has both a proton pump activity and ATP-synthase in
which it can generate ATP (the only ATPase.
V-type ATPases have similar structure to F-type ATP-ases. They are also
proton pumps and allow protons to enter organelles (2protons / 1 ATP
molecule). the point of this pump is to acidify organelles, like lyzosomes for
protein degradation. It is insensitive to vanadate ion VO3- but strongly
inhibited by NO3 P-type ATPases are broader. more simple in structure, they're ubiquitously
expressed. they are cation pumps (Na+, K+, Ca+, H+ etc). these are really
important since the exchange of cations is vital for cells to survive and
function. These are inhibited by vanadate ion VO3- but are insensitive to
NO3-. Vanadate competes with phosphate for binding sites which indicate
that ATP transfer phosphate group to the ATPase protein. The resulting
phosphoenzyme then goes a conformational change that expose protein
binding site to outside
Ca2+ type ATPase couples the hydrolysis of ATP with translocation of Ca2+
across the membrane. This serves to keep the cytosolic Ca2+ concentration
low which is necessary to avoid precipitating phosphates and Ca2+ signaling
pathways operate properly

 So, the proton pumping ATPases create both the proton

concentration gradient (pH gradient) and the membrane
electrical potential.
The electrical gradient explains why some ions are distributed
unequally across the membrane, even without the direct input
of energy.
The proton concentration gradient provides the free energy for
many kinds of secondary active transport.
The proton gradient pH together with membrane potential
contribute to proton motive force that tends to move protons
back across the membrane
pmf = 59 pH

 Couple movement of one substrate down the gradient to the

movement of another substrate against the gradient
Symporter - both go in the same direction
Antiporter - molecules go in opposite directions
Cotransporters require the existence of a gradient of one
molecule (created by pumps)
Energy of the transport down the gradient allows for the
transport of second substrate against the gradient

K+ exchange is mediated by two classes of transport proteins

Classified into two types

High affinity Transport System (HAT): active at low K+
concentrations (< 200 M). They have low capacity for ion
uptake. However, their affinity is very high because of their
specificity with that ion. It exhibit non-linear saturation kinetics.
E.g., H+ ATPase linked K+ H+ symporter
Low Affinity Transport System (LAT). Their capacity is high
and affinity is low. Ion uptake is linear concentration dependent.
E.g., K+ channels

Root architecture is important for ion uptake

Q1.What is Apparent Free Space (AFS)? How is it important?
Ans. It is the volume of the root that is accessible by free diffusion. It consists
of cell wall and intercellular spaces (equivalent to apoplastic space) of the
epidermis and cortex. These are the regions of the root which can be entered
without crossing the space. The cation exchange capacity of the AFS shows
direct correlation with the carboxyl groups associated with the galacturonic
acid residues.
Q2 Why the ions cannot enter the steel directly from cortex?
Ans. Endodermis has specialized thickenings at radial and transverse walls
called casparian stripes. The casparian stripes are principally composed of
complex mixture of hydrophobic long chain fatty acids and alcohols called
suberin. These are strongly attached to the plasma membrane of the cell wall.
The hydrophobic space filling nature of casparian stripe reduces the
possibility of ions or small hydrophillic molecules without first entering the
symplast.

Transport of mineral nutrients within plants

Outermost region - root epidermis/ Rhizodermis and Cortical Cells
Middle Region- Endodermis with its suberized casparian band
Innermost region consists of vascular tissues-the vessel elements and
associated parenchyma cells
Apparent Free Space- Apoplast outside the endodermis and the casparian
stripe.
Symplastic connections facilitate passive movement from cell to cell
until they arrive at xylem parenchyma in the stele.