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Sap Translocation

Transpiration

Plants can manipulate the solute potential and pressure potential within tissues in order to establish water potentials that facilitate movement

  • Transpiration involves the generation of negative hydrostatic pressures within the leaves (via evaporation) and positive hydrostatic pressures within the roots

  • This pressure differential creates tension within the xylem vessels that allow water to be transported against gravity via capillary action

Root Pressure

Root pressure is a positive pressure potential within the root tissue that can generate water movement up the xylem

  • Root pressure can allow water to move up the stem when transpiration is insufficient (e.g. when humidity is high or before bud burst in spring)

Plants control the uptake of water from the soil by regulating the movement of mineral ions via indirect active transport

  • Root cells contain proton pumps that actively expel H+ ions (stored in the vacuole of root cells) into the surrounding soil

  • The H+ ions displace the positively charged mineral ions from the clay, allowing them to diffuse into the root along a gradient

  • Water will follow the mineral ions into the root via osmosis – with the rate of water uptake regulated by water channels (aquaporins) on the root cell membrane

Mineral Uptake in Roots

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Protons are pumped into the soil
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Ions displaced to diffuse into root
Regulating Water Uptake

Once inside the root, water will move towards the xylem via one of two pathways (symplastic or apoplastic)

  • In the symplastic pathway, water moves continuously through the cytoplasm of root cells (connected via plasmodesmata)

  • In the apoplastic pathway, the water is water moved through the cell walls of the intervening root cells (more rapid)

The endodermal cells that separate the root from the xylem contain a waterproof band of suberin within the cell walls

  • This is called the Casparian strip and prevents further water movement via the apoplastic pathway

  • The water is forced into the symplastic pathway and must therefore cross cell membranes to access the xylem

  • This allows the plant to regulate the uptake of water into the xylem vessels and control root pressure

Water Movement in Root Tissue

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Translocation

Translocation is the movement of organic compounds (e.g. sugars, amino acids) from sources to sinks

  • The source is where the organic compounds are synthesised – this is typically the photosynthetic tissues (leaves)

  • The sink is where the compounds are delivered to for use or storage – this includes the roots, fruits and seeds

  • Organic compounds are transported via a vascular tube system called the phloem as part of a viscous fluid called sap

Phloem Structure

Phloem sieve tubes are primarily composed of two main types of cells – sieve element cells and companion cells

  • Plasmodesmata connect the sieve elements and companion cells to mediate symplastic exchange (continuous cytoplasm)

Sieve elements are the long and narrow cells that are connected together to form the phloem sieve tubes

  • Sieve elements are connected by sieve plates at their transverse ends, which are porous to enable flow between cells

  • Sieve elements have no nuclei and reduced cytoplasm and organelles to maximise space for the translocation of materials

  • The sieve elements also have thick and rigid cell walls to withstand the hydrostatic pressures which facilitate flow

Companion cells provide metabolic support for sieve elements and facilitate loading and unloading at sources and sinks

  • They possess an infolding plasma membrane which increases SA:Vol ratio to allow for more material exchange

  • Have many mitochondria to fuel the active transport of materials between the sieve tube and the source or sink

  • Contain appropriate transport proteins within the plasma membrane to move materials into or out of the sieve tube

Sieve Tube Adaptations

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Sap Transport

At the source (e.g. leaves), the active transport of sugars into the phloem by companion cells makes the sap solution hypertonic

  • This causes water to be drawn from the xylem via osmosis (water moves towards higher solute concentrations)

  • Due to the incompressibility of water, this build up of water in the phloem causes the hydrostatic pressure to increase

  • This increase in hydrostatic pressure forces the phloem sap to move towards areas of lower pressure (mass flow)

  • Hence, the phloem transports solutes away from the source (and consequently towards the sink)

At the sink (e.g. roots, fruits, seeds), the solutes within the phloem are unloaded by the companion cells

  • This causes the sap solution at the sink to become increasingly hypotonic (lower solute concentration)

  • Consequently, water is drawn out of the phloem and back into the xylem by osmosis

  • This ensures that the hydrostatic pressure at the sink is always lower than the hydrostatic pressure at the source

  • When organic molecules are transported into the sink, they are either metabolised or stored within the tonoplast of vacuoles

Translocation in Plants

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