Plant systems - Phloem Flashcards
Phloem
Living tissue made up of several cell types including sieve tube elements and companion cells
Sieve tube elements
- Cells without nuclei and other organelles that join end to end to form sieve tubes
- Their end walls are perforated with pores and are known as sieve plates
- Strands of cytoplasm containing phloem proteins extend from one sieve tube to the next through the sieve plates
Companion cell
- Very metabolically active
- Large nucleus and dense cytoplasm with high numbers of RER and mitochondria
- Connected to neighbouring sieve tube elements via plasmodesmata
- ->Allowing exchange of materials to take place
Translocation
The active movement of the soluble products of photosynthesis, like sucrose and amino acids, through the phloem from sources to sinks
Source
Any region of a plant where sucrose is formed
- Usually leaves
- In deciduous plants it is the roots during winter as leaves fall off and stored starch in roots is hydrolysed to sucrose
Sink
Any region of the plant where sucrose is stored as starch
- Growing regions of plants that need a source of sucrose
- Roots, flowers and buds etc
Ringing experiment
- A ring of bark is removed from just below a section of a tree trunk or woody stem
- Phloem lies just beneath so it is removed in that section too
- After time the tissue above the ring swells, and below the ring shrinks/withers due to lack of nutrients
- Liquid in swollen tissue shown to be amino acids and sucrose
- Suggesting that removal of the phloem interrupts the downward movement of these substances
- Proves translocation of sucrose and amino acids occurs in the phloem
Aphids experiment
It is difficult to extract sap and to measure the speed of translocation as when the phloem is cut phloem proteins clog pores in sieve plates to prevent loss of sap.
- Aphids feed on sap within phloem sieve tubes using a very narrow stylet mouthpiece
- The stylet cannot be blocked by phloem proteins, so if cut off from Aphid sap flows out freely
- The experiment in conjunction with radioactive source of CO2 have shown the rate of translocation is much faster than rate of diffusion alone
- Suggesting an additional mechanism is involved in the process of translocation
Radioactive CO2
- If plants photosynthesise in the presence of radioactive CO2 its sucrose will incorporate it when it is formed
- Allows transport and location of sucrose to be visualised
- Transverse section (vertical) cut through the stem and placed on photographic film
- Positions of exposure and therefore radioactivity coincide with position of the phloem
- Indicating it is the phloem that translocates the sucrose made by photosynthesis
Theories of translocation
- The Mass flow hypothesis
- Active processes
- Cytoplasmic streaming
Mass flow hypothesis / Translocation as a passive process
Explains how sucrose synthesised at the source (e.g. leaves) is translocated via the phloem to the sink (e.g. roots)
LOADING SUCROSE AT SOURCE:
- Excess glucose synthesised during photosynthesis is converted to the disaccharide sucrose in mesophyll cells
- The sucrose enters companion cells via the plasmodesmata
- Sucrose is then loaded into the sieve tube elements by facilitated diffusion via plasmodesmata
- Water potential in sieve tubes is lowered
- Water from adjacent xylem vessels is drawn in by osmosis, increasing the hydrostatic pressure
- Water moves down a hydrostatic pressure gradient to sink
UNLOADING SUCROSE AT SINK:
- Sucrose moves down a concentration gradient into tissues of the sink
- This increases water potential in sieve tubes so water moves into xylem by osmosis
- This lowers the hydrostatic pressure in sieve tubes near the sink, creating a hydrostatic pressure gradient
- Sucrose is hydrolysed by enzymes inside sink cell into glucose and fructose to maintain a concentration gradient between sieve tube and sink
The mass flow model
X represents source
Y represents sink
T represents phloem
S represents xylem
- X contains concentrated sucrose solution surrounded by partially permeable membrane
- Y contains dilute sucrose solution surrounded by partially permeable membrane
- Tendency for loss of water from both X and Y into S by osmosis. But tendency is greater for X
- Water moves down a water potential gradient into X, increasing hydrostatic pressure in X
- The solution is forced out of X and into Y via tube T
- Hydrostatic pressure in Y will increase, which forces water out into S, down a water potential gradient
Limitations of mass flow hypothesis
- Rate of translocation is 10,000 times faster than if moving just by diffusion, suggesting translocation is an active process and not a passive one
- Companion cells are very biochemically active, but mass flow does not suggest a role for them
- Phloem have high oxygen consumption, suggesting translocation is an active process
- Does not take into account the presence of the sieve plates in the sieve tubes
Translocation as an active process
- H+ ions are actively transported out of companion cells into the mesophyll cells
- Creates excess H+ ions in the mesophyll
- Hydrogen ions move back into the companion cells down their concentration gradient via co-transporter proteins
- Sucrose is carried at the same time into companion cells
- Sucrose then loaded into into the sieve tubes down a concentration gradient via apoplast or symplast pathway
Cytoplasmic streaming theory
- Strands of cytoplasm containing tiny microtubules exist between phloem sieve tube elements, running through sieve pores in sieve plates
- The theory suggests solutes also move within this stream
- This accounts for movement of solutes in both upwards and downwards directions