Talk 7 - Translocation in The Phloem
Talk 7 - Translocation in The Phloem
A highly specialized process for redistributing: Photosynthesis products Other organic compounds (metabolites, hormones) some mineral nutrients
Phloem transport
SINK
How the growing parts of the plant are provided with sugar to synthesize new cells
A system of vascular tissue runs through all higher plants. It evolved as a response to the increase in the size of plants, which caused an progressing separation of roots and leaves in space. The phloem is the tissue that translocates assimilates from mature leaves to growing or storage organs and roots.
Photosynthesis
Translocation
New growth
sugar source
Direction of transport through phloem is determined by relative locations of areas of supply, sources and areas where utilization of photosynthate takes place, sinks.
Source:
Translocation
any transporting organ capable of mobilizing organic compounds or producing photosynthate in excess of its own needs, e.g., mature leaf, storage organ during exporting phase of development.
New growth is a
sugar sink
Sink:
non photosynthetic organs and organs that do not produce enough photoassimilate to meet their own requiements, e.g., roots, tubers, develpoping fruits, immature leaves.
less reactive
Reducing sugars, such as Glucose, Mannose and Fructose contain an exposed aldehyde or ketone group
be transported in the phloem
Fructose
is combined with a similar group on another sugar Or the ketone or aldehyde group is reduced to an alcohol
D-Mannitol
Most of the other mobile sugars transported contain Sucrose bound to varying numbers of Galactose units
Remember Sucrose?
Sucrose
The osmotic effect of a substance is tied to the number of particles in solution, so a millilitre of
sucrose solution with the same osmolarity as glucose will be have twice the number carbon atoms and therefore about twice the energy.
Thus, for the same osmolarity,
As a non-reducing sugar, sucrose is less reactive and more likely to survive the journey in the phloem. Invertase (sucrase) is the only
enzyme that will touch it and this is unlikely to be present in the phloem sieve tubes.
Other compounds
Water!!!!!!!!! Nitrogen is found in the phloem mainly in:
amino acids (Glutamic acid) Amides (Glutamine)
Phloem Structure
The main components of phloem are
sieve elements companion cells.
Sieve elements have no nucleus and only a sparse collection of other organelles . Companion cell
provides energy
so-named because end walls are perforated - allows cytoplasmic connections between vertically-
stacked cells
conducts sugars and amino acids - from the leaves, to the rest of the plant
Sieve tubes elements join to form continuous tube Pores in sieve plate between sieve tube elements are open channels for transport Each sieve tube element is associated with one or more companion cells.
Many plasmodesmata penetrate walls between sieve tube elements and companion cells Close relationship, have a
The flow is driven by an osmotically generated pressure gradient between the source and the sink. Source
speed
Sugars (red dots) is actively loaded into the sieve elementcompanion cell complex
Called phloem loading
Sink
yw = ys + yp + yg
In source tissue, energy driven phloem loading leads to a buildup of sugars
Makes low (-ve) solute potential Causes a steep drop in water potential In response to this new water potential gradient, water enters sieve elements from xylem
Thus phlem turgor pressure
increases
Makes a higher (+ve) solute potential Water potential increases Water leaves phloem and enters sink sieve elements and xylem
Thus phloem turgor pressure
decreases
Triose phosphate formed from photosynthesis during the day is moved from chloroplast to cytosol At night, this compound, together with glucose from stored starch, is converted to sucrose
Both these steps occur in a
Sucrose then moves from the mesophyll cell via the smallest veins in the leaf to near the sieve elements
mesophyll cell
Involves a sucrose-H+
The energy dissipated by protons
Requires the presence of open plasmodesmata between different cells in the pathway Dependant on plant species with intermediary companion cells
Sucrose, synthesized in mesophyll, diffuses into intermediary cells Here Raffinose is synthesized. Due to larger size, can NOT diffuse back into the mesophyll Raffinose and sucrose are able to diffuse into sieve element
Phloem unloading
Three steps (1) Sieve element unloading: (2) Short distance transport:
Transported sugars leave the sieve elements of sink tissue After sieve element unloading, sugars transported to cells in the sink by means of a short distance pathway Sugars are stored or metabolized in sink cells
Also can occur by symplastic or apoplatic pathways Varies greatly from growing vegetative organs (root tips
Phloem unloading
and young leaves) to storage tissue (roots and stems) to reproductive organs Symplastic:
Appears to be a completely symplastic pathway in young dicot leaves Again, moves through open plasmodesmata
Apoplastic: three types (1) [B] One step, transport from the sieve elementcompanion cell complex to successive sink cells, occurs in the apoplast. Once sugars are taken back into the symplast of adjoining cells transport is symplastic
Phloem unloading
Apoplastic: three types (2) [A] involves an apoplastic step close to the sieve element companion cell. (3) [B] involves an apoplastic step father from the sieve element companion cell Both involve movement through the plant cell wall
Phloem unloading
Summary
Pathway of translocation:
Sugars and other organic materials are conducted throughout the plant in the phloem by means of sieve elements
Sieve elements display a variety of structural
Patterns of translocation:
Materials are translocated in the phloem from sources (usually mature leaves) to sinks (roots, immature leaves)
Summary
Materials translocated in phloem:
Translocated solutes are mainly carbohydrates Sucrose is the most common translocated sugar Phloem also contains:
Amino acids, proteins, inorganic ions, and plant
Rate of translocation:
Movement in the phloem is rapid, well in excess of rates of diffusion
Average velocity is 1 meter per hour
hormones
Pressure-flow
Phloem and xylem are coupled in an osmotic system that transports sucrose and circulates water.
Physiological process of unloading sucrose from the phloem into the sink
Build-up of pressure at the source and release of pressure at the sink causes source-to-sink flow. At the source phloem loading causes high solute concentrations.
decreases, so water flows into the cells increasing hydrostatic pressure. At the sink y is lower outside the cell due to unloading of sucrose. Osmotic loss of water releases hydrostatic pressure. Xylem vessels recycle water from the sink to the source.