Translocation of photoassimilates

By Fredeslinda C. Evangelista, PhD.

Photoassimilate- product of photosynthesis Translocation- long distance transport of photoassimilates Translocation occurs in the phloem

Evidences for phloem translocation of photoassimilates

By girdling Analysis of phloem exudate using the aphid technique Use of radioactive tracers usually on the leaf

Sieve tube member

Honeydew droplet

stylet

Phloem sap droplet

The composition of phloem exudate

Principal constituent is sugar i.e. sucrose Why sucrose? It is nonreducing so less likely to react with other substances along the way. It may also contain minerals, amino acids, proteins and hormones.

Direction of long distance translocation in the phloem

From source to sink Source net exporter or producer of photoassimilate a plant organ in which sugar is being produced by either photosynthesis or the breakdown of starch. Example mature leaf

Direction of long distance translocation in the phloem

From source to sink Sink net importer or consumer of photoassimilate Or a storer of sugar Example- growing fruit

Partitioning of assimilate among sinks

Sinks are competitive and are partitioned to all active sinks Partitioning depends on 3 factors Nature of the vascular connections between source and sinks

Proximity of the sink to the source Sink strength

sinks-photoassimilate will move preferentially toward the sink leaf above and in line (same rank) with the source leaf

Sink strength- is a measure of the capacity of a sink to accumulate metabolites

Mechanism of translocation in the phloem

By pressure flow/mass flow

Mass transfer of solute from source to sink along a hydrostatic (turgor) pressure gradient Translocation is fundamentally a passive process Transport is bidirectional

Pressure flow is the mechanism of translocation in angiosperms

Phloem sap flows from source to sink at rates as great as 1 m/hr, faster than can be accounted for by either diffusion or cytoplasmic streaming.

Higher levels of sugar at the source lowers the water potential and causes water to flow into the tube.

Pressure flow is the mechanism of translocation in angiosperms

Removal of sugar at the sink increases the water potential and causes water to flow out of the tube.

The difference in hydrostatic pressure drives phloem sap from the source to the sink

Pressure flow is the mechanism of translocation in angiosperms

(1) Loading of sugar into the sieve tube at the source reduces the water potential inside the sievetube members and causes the uptake of water.
(2) This absorption of water generates hydrostatic pressure that forces the sap to flow along the tube.

Pressure flow is the mechanism of translocation in angiosperms

(3) The pressure gradient is reinforced by unloading of sugar and loss of water from the tube at the sink.
(4) For leaf-to-root translocation, xylem recycles water from sink to source.

Phloem loading

Initial transfer of assimilate from mesophyll cells into sieve elements at the source end Movement of sucrose from mesophyll to phloem (parenchyma) is by the symplast

Phloem loading

From the phloem parenchyma, there are 2 possible routes: May continue to companion cellsieve-tube member complex via the symplast.

Sucrose reaches sieve-tube members by a combination of symplastic and apoplastic pathways.

Phloem loading

Evidence indicates that a sucrose-H+ cotransport mediates sucrose transport between the apoplast and symplast

Proton pumps generate an H+ gradient, which drives sucrose across the membrane via a cotransport protein that couples sucrose transport with the diffusion of H+ back into the cell.

Phloem loading

symplastic pathway + Sucrose-H cotransport exists as a mechanism for retrieving sucrose that leaked from photosynthetic cells

To prevent unneccessary loss of sugar from the transport stream

Phloem loading

Cc-sc complex symplastically connected with adjacent cells- exhibit symplastic loading Cc-sc complex symplastically isolated exhibit apoplastic loading

Phloem unloading

Final transfer of assimilate of the sieve elements into target cells

Mechanisms

By symplastic pathway Flow maintained by hydrolyzing sucrose into glucose and fructose By apoplastic pathway

Phloem unloading

Mechanisms By apoplastic pathway Passive unloading into apoplast (e.g. maize seeds) Active unloading into apoplast (e.g. legume seeds)