8. Trans Location of Photo Assimilates

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    Translocation of photoassimilates

    By

    Fredeslinda C. Evangelista, PhD.

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    Photoassimilate- product ofphotosynthesis

    Translocation- long distancetransport of photoassimilates

    Translocation occurs in thephloem

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    Evidences for phloemtranslocation of photoassimilates

    By girdling

    Analysis of phloem exudateusing the aphid technique

    Use of radioactive tracersusually on the leaf

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    Sievetubemember

    stylet Phloem sapdroplet

    Honeydewdroplet

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    The composition of phloem

    exudate Principal constituent is sugar i.e. sucrose

    Why sucrose?

    It is nonreducing so less likely to reactwith other substances along the way.

    It may also contain minerals, amino acids,proteins and hormones.

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    Direction of long distance

    translocation in the phloem From sourceto sink

    Source

    net exporter or producer of

    photoassimilate

    a plant organ in which sugar is beingproduced by either photosynthesisor the breakdown of starch.

    Examplemature leaf

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    From sourceto sink

    Sink

    net importer or consumer of

    photoassimilate

    Or a storer of sugar

    Example- growing fruit

    Direction of long distance

    translocation in the phloem

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    Partitioning of assimilate

    among sinks Sinks are competitive and are partitioned to

    all active sinks

    Partitioning depends on 3 factors

    Nature of the vascular connectionsbetween source and sinks

    Proximity of the sink to the source

    Sink strength

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    sinks-photoassimilate will move preferentiallytoward the sink leaf above and in line (same

    rank) with the source leaf

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    Sink strength- is a measure of the capacity of a sink toaccumulate metabolites

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    Mechanism of translocationin the phloem

    By pressure flow/mass flow

    Mass transfer of solute from source to sinkalong a hydrostatic (turgor) pressure gradient

    Translocation is fundamentally a passive

    process Transport is bidirectional

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    Pressure flow is the mechanismof translocation in angiosperms

    Phloem sap flows from source to sink atrates as great as 1 m/hr, faster than can be

    accounted for by either diffusion orcytoplasmic streaming.

    Higher levels of sugar at the source

    lowers the water potential and causeswater to flow into the tube.

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    Removal of sugar at the sinkincreases the water potential and

    causes water to flow out of the tube.

    The difference in hydrostatic pressuredrives phloem sap from the source tothe sink

    Pressure flow is the mechanismof translocation in angiosperms

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    (1) Loading of sugar into the sievetube at the source reduces the

    water potential inside the sieve-tube members and causes theuptake of water.

    (2) This absorption of watergenerates hydrostatic pressure thatforces the sap to flow along the

    tube.

    Pressure flow is the mechanismof translocation in angiosperms

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    Pressure flow is the mechanism

    of translocation in angiosperms

    (3) The pressure gradient isreinforced by unloading of sugarand loss of water from the tube atthe sink.

    (4) For leaf-to-root translocation,xylem recycles water from sink tosource.

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    Phloem loading

    Initial transfer ofassimilate frommesophyllcells intosieve elements at thesource end

    Movement of

    sucrose frommesophyll tophloem(parenchyma) is by

    the symplast

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    From the phloem parenchyma, thereare 2 possible routes:

    May continue to companion cell-sieve-tube member complex via thesymplast.

    Sucrose reaches sieve-tube membersby a combination of symplastic andapoplastic pathways.

    Phloem loading

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    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 thatcouples sucrose transport with thediffusion of H+ back into the cell.

    Phloem loading

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    symplastic pathway

    Sucrose-H+ cotransport exists as a

    mechanism for retrieving sucrose thatleaked from photosynthetic cells

    To prevent unneccessary loss of sugarfrom the transport stream

    Phloem loading

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    Cc-sc complex symplastically connectedwith adjacent cells- exhibit symplastic

    loading Cc-sc complex symplastically isolated

    exhibit apoplastic loading

    Phloem loading

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    Phloem unloading

    Final transfer of assimilate of the sieveelements into target cells

    Mechanisms

    By symplastic pathway

    Flow maintained by hydrolyzingsucrose into glucose and fructose

    By apoplastic pathway

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    Mechanisms

    By apoplastic pathway

    Passive unloading into apoplast(e.g. maize seeds)

    Active unloading into apoplast (e.g.legume seeds)

    Phloem unloading

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