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3.

8 THE RADIAL PATH OF ION


MOVEMENT THROUGH
ROOTS
3.8.1 IONS ENTERING THE STELE
MUST FIRST BE TRANSPORTED
FROM THE APPARENT FREE
SPACE INTO THE SYMPLAST
Rapid distribution of nutrient ions throughout the plant
is accomplished in the xylem vessels. In order to reach
these conducting tissues, which are located in the central
core, or stele, of the root, the ions must move in
a radial path through the root. The path these ions
must follow is diagrammed in Figure 3.12. For these
purposes we may consider the root as consisting of
three principal regions. The outermost region consists
of the root epidermis (often referred to as the rhizodermis)
and the cortical cells. The innermost region
consists of vascular tissues—the vessel elements and
associated parenchyma cells—which are of particular

interest for our discussion. Separating the two is the


endodermis with its suberized Casparian band.
Ion uptake begins with free diffusion into the apparent
free space. As noted in the previous section, the
apparent free space is equivalent to the apoplast outside
the endodermis and the Casparian band effectively
prevents further apoplastic diffusion through the endodermis
into the stele. Hence the only possible route for
ions to pass through the endodermis is to enter the symplast
by some carrier- or channel-mediated transport at
the cell membrane. This may occur either on the outer
tangential wall of the endodermal cell itself or through
any of the epidermal or cortical cells. Regardless of
which cell takes up the ions, symplastic connections
(i.e., plasmodesmata) facilitate their passive movement
from cell to cell until they arrive at a xylem parenchyma
cell in the stele. At this point the ions may be unloaded
into the xylem vessels for long distance transport to the
leaves and other organs.
3.8.2 IONS ARE ACTIVELY SECRETED
INTO THE XYLEM APOPLAST
With the exception of the very tip of the root where the
young xylem vessel elements are still maturing, functional
xylem is part of the apoplast. The interconnected
vessel elements are devoid of cytoplasm and consist
only of nonliving tubes filled with an aqueous solution.
Release of ions into the xylem thus requires a transfer
from the symplast into the apoplast. At one time, it was
thought that this transfer was simply a passive leakage,
but it is now clear that ions are actively secreted from
xylem parenchyma cells. Although there is some conflicting
evidence, ion concentration in the apoplast of the
stele is generally much higher than in the surrounding
cortex. This suggests that ions are being accumulated
in the xylem against a concentration gradient, presumably
by an energy-dependent, carrier-mediated process.
It is also interesting to speculate, in this regard, that
the Casparian band also functions to prevent loss of
ions from the stele by blocking their diffusion down a
concentration gradient.
In addition to working uphill against a concentration
gradient, delivery of ions into the xylem
vessels is sensitive to metabolic inhibitors such as
carbonyl-cyanide-m-chlorophenylhydrazone (CCCP),
which uncouples ATP formation. It is interesting
that ion transport into the xylem is also sensitive to
cycloheximide, an inhibitor of protein synthesis, but
uptake into the root, at least initially, is not affected.
Two plant hormones (abscisic acid and cytokinin) have
a similar effect. Whether inhibitors of protein synthesis
and hormones are affecting symplastic transport
through the endodermis or unloading of ions from the
endodermis into the xylem is not certain, but these
results at least raise the possibility that ion release into the vessels is a different kind of process
than ion uptake
by the roots.

FIGURE 3.12 The radial paths of ion movement through a root. Arrows indicate the
alternative paths that may be taken by nutrient ions as they move from the soil
solution into the vascular elements in the stele. Arrows with circles indicate active
transport of ions across plasma membranes.