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Explanation of the mineral uptake into the cells
of plants
Uptake of Mineral Nutrients into Cells
Nutrients cannot be used by cells while they are in the apoplasm.
For metabolism they need to be transferred to the inside of
the cell. This process occurs in several ways, all of which
are dependent in some way upon the electrochemical gradient
across the plasma membrane separating the cell cytoplasm and
the cell wall. The same is true of transport across the tonoplast
into the vacuole.
Transport into the cell (or vacuole) can be divided into two
main types: passive which doesn’t require any energy input,
and active that does. An example of passive transport would
be when an ion is present in a higher concentration outside
of a cell than inside. This would lead to the ion diffusing
across the membrane along the chemical gradient set up by the
difference in concentration. This passage would be facilitated
either by carrier proteins or through aqueous pores in the membrane.
Further accumulation of such elements beyond the equilibrium
point can be achieved by de-activating them e.g. by incorporating
them into organic molecules or by attaching them to charged
molecules.

All of the other known methods of transport across the plasma
membrane are active processes although some of these use the
energy to set up an appropriate electro-chemical gradient rather
than to directly transport the ion in question.
One of the most significant processes involved in the active
transport of ions is the acidification of the apoplasm by pumping
H+ ions into the cell wall from the cytoplasm. This sets up
both a pH and a charge gradient across the plasma membrane.
Ions can then use this potential difference in various ways
to aid their crossing of the membrane.
This gradient is set up by a membrane bound, ATP consuming (ATPase)
H+ pump. The importance of this process is highlighted by the
proportion of ATP used in the roots to import ions using such
processes which can be as high as 36% (Werf et al., 1988). Another
gradient across the tonoplast is also set up. Here H+ ions are
pumped from the cytoplasm into the vacuole. This leaves the
cytoplasm with a pH of 7.3-7.6, with the apoplasm and vacuole
having pHs of approx 5.5 and 4.5-5.9 respectively.
The power of the potential differences set up across membranes
in higher plants is demonstrated by an equation called the Nerst
equation:
This shows that at a potential difference of -59mv (real differences
are often in excess of this), monovalent ions have to be present
in the cell at concentrations ten times higher(cations) or ten
times lower(anions) in order for electrochemical equilibrium
to be reached. The factor of ten is increased to a factor of
a hundred in divalent ions. This means that cations that are
present in the external solution in reasonable amounts are able
to use the electrochemical gradient set up across the membrane
to travel across it.

This passage is mediated by proteins within the membrane. These
are either carrier proteins which allow relatively small numbers
of ions across, or channel proteins which allow large number
of ions to pass through there central pore. Channel proteins
spend most of their time in the closed position, opening in
response to environmental stimuli.
Carrier proteins have three different methods of transport:
- uniport, symport, and antiport. Uniport is when an ion is
transferred across the membrane on its own. This method is likely
to be used in the case of ions passing down an electrochemical
gradient i.e. cations whose external concentration is sufficiently
large.
Symport is when an ion which may not be able to travel along
an electro-chemical gradient is attached to an ion that is able
to travel along the gradient, and so travels through the membrane
with it. K+ ions which sometimes have a prohibitively low external
concentration for crossing the plasma membrane can pass across
in this way by attaching to H+ ions, which are able to travel
along the electrochemical gradient set up by the ATPase pump.
It also believed that anions such as nitrate are transported
across the membrane using this method.

Counter transport is transporting an ion against a concentration
gradient by attaching it to a transporter which is transporting
an ion in the opposite direction along a concentration gradient.
This method is believed to be used to transport ions from the
cytoplasm into the vacuole. All three methods can be viewed
in the diagram below.
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