CAPE PLANT TRANSPORT - translocation

Ms. Seejoor

EVIDENCE: TRANSLOCATION OCCURS IN PHLOEM

Use of aphids:

o Aphids insert their stylets into phloem sieve tubes

o Researchers cut the stylets from the aphids’ bodies  , leaving a tiny tube with one end in
the phloem and the other in the atmosphere
o Phloem sap oozed out of the tubes (indicates that the contents of sieve tubes are under
positive pressure)
o A larger volume of phloem sap than can be contained in the tube was collected from a tube
(indicates that material is continually loaded into sieve tubes)
o Analysis of the exudate showed that it contained organic molecules such as sucrose
(indicates the organic molecules made in photosynthesis are transported in phloem
o Diurnal variations in the sucrose content of leaves are identical to the variations in the
sucrose content of the phloem, a little while later (indicates that sucrose is loaded into the
phloem)

Use of radioactive CO2 as a tracer:

o Leaves of a plant were enclosed in a transparent bag with 14CO2

o Radioactivity (radioactive sucrose) appeared in the leaves, leaf phloem, stem phloem, sinks
(young shoot tips, flowers, fruits, roots), in that order

Ringing experiments:

o Done by Mason and Maskel in Trinidad

o A collar (or ring) of tissue external to the xylem was removed (phloem was removed)
o Plant tissue bulged in the circumference of the tree above the ring (indicates the material
flowing in the phloem was flowing downwards and this movement was stopped)
o Concentration of sucrose was highest in the region above the ring and negligible in the
region below the ring. In the control (unringed plant), the concentration of sucrose was
approximately the same in the regions corresponding to the areas above and below the ring
EVIDENCE: PRESSURE FLOW HYPOTHESIS / MASS FLOW HYPOTHESIS

o Phloem sap has a high pH, approximately 8 (active transport of protons out of companion
cells)

o There is a higher negative charge within companion cells than outside (active transport of
protons out of companion cells)

o When sieve tubes are cut, sap exudes outside (the contents are under positive pressure)

o The concentration of sucrose is higher in leaves than in roots (loading occurs at source,
unloading at sink)

o Downward flow in phloem occurs in daylight, ceases when plants are shaded or at night
(sucrose no longer available for loading, to initiate creation of a pressure gradient)

o The distance travelled by sucrose in the phloem is very long, up to 100 metres (phloem
tubes are continuous throughout the length of a plant)

o The rate of flow of sucrose in phloem is usually between 0.05 and 0.25 m h -1, and may be as
high as 1m h-1 (flow of sucrose in phloem does not occur by diffusion, which would happen
at a much lower rate: less than 20mm h -1. The rates of flow in sieve tubes are similar to
those predicted according to the size of the pressure gradients found in phloem)

o The rate of flow in phloem decreases when temperature and oxygen concentrations
decrease. The presence of respiratory inhibitors stops the flow in phloem (active transport
is a part of the mechanism of flow of materials in the phloem)
o The hydrostatic pressures inside phloem sieve tubes are high: 1000 – 2000 kPa (mass flow
requires the existence of high pressure)

o The pressure gradient inside sieve tubes is approximately 20kPa m -1 (mass flow requires a
pressure gradient of approximately 13kPa m -1)

o Phloem sap may travel in different directions in adjacent sieve tubes. Even within a single
tube, the direction of flow changes during the life of the plant, e.g. sap is transported to
young non-photosynthetic leaves when they are actively growing, then later, sap is
transported away from them as they become sources (the direction of flow of materials
depends on the direction of the pressure gradient)

o The concentration of sucrose in sieve tubes is 10% - 30% and its concentration in mesophyll
cells is 0.5% (since the potential energy of the high proton concentration is used for the
diffusion of sucrose, then sucrose can be co-transported with protons even against a sucrose
concentration gradient)

o There are numerous mitochondria and a high concentration of ATP in companion cells
(needed for active transport of protons out of companion cells)

o Sucrose is loaded into sieve tubes but other sugars are not (the cotransporter protein in the
cell membrane of the companion cells is specific for protons and sucrose only)

o The cell surface membrane of companion cells in the leaves is highly folded and contain
numerous active proton pumps and passive proton-and-sucrose symporter proteins (protons
 are actively transported out of companion cells, and are passively cotransported back with
sucrose, into companion cells)

o The water potential of sieve tubes is approximately -2.5MPa and the water potential of
mesophyll cells is approximately -1.5MPa (a water potential gradient exists from
neighbouring cells and xylem vessels to sieve tubes, this allows water entry into sieve tubes
during loading)
o Increases in sucrose levels in leaves are followed by similar increases in sucrose levels in
phloem sieve tubes (sucrose is loaded into sieve tubes)

o Certain viruses move in the translocation stream (suggests mass flow rather than diffusion
since the virus is not in solution. Viruses are incapable of independent locomotion)

OBSERVATIONS WHICH ARE NOT EXPLAINED BY THE PRESSURE FLOW HYPOTHESIS

o Different solutes move at different speeds in the phloem: they should move at the same
speed if they are being transported by mass flow.

o Sucrose is delivered at the same rate to all sinks: it should be delivered at a faster rate to the
sinks with the lowest sucrose levels, if it is being transported by mass flow.

o There is no explanation for the role of sieve plates in the mass flow theory. It is possible that
the role of sieve plates is unrelated to the actual flow of materials. They may be present to
prevent bulging and explosion of the sieve tubes (due to high internal pressures), by
physically being connected to the inner circumference of the sieve tubes. It is also easier to
plug the sieve pores with phloem protein, then later with callose, in the event of damage to
the sieve tubes (than plugging the entire lumen of the sieve tubes). This reduces the
amount of material lost when a sieve tube is damaged.
o There is no reason for the sieve tubes to be alive, (as opposed to the dead tubes of xylem, in
the mass flow theory. It is possible that the reason is unrelated to the actual flow of material
in the tubes. Sieve tubes may be alive to maintain the presence of a cell membrane. Once
sucrose is loaded, the presence of a hydrophobic cell membrane would not allow the
diffusion of the polar molecule sucrose out of the tube. Leakage of the sucrose is possibly
prevented.