Phloem is specialized for Carbohydrate Transport and is typically found
adjacent to Xylem. In order for plants to evolve in size and complexity it was essential
to have a rapid system for the movement of carbohydrates from one part of a plant to
another.
This is obvious if you consider a large
tree like
the Giant Sequioa. Photosynthesis occurs in the leaves of the
crown. These are several hundred feet above the soil. In order for sucrose to reach roots
which may be a hundred feet below the soil, a rapid physiological transport system must be
present. Diffusion operates efficiently over very short
distances. Carbohydrate diffusion could sustain microscopic plants or very small
macroscopic plants. Calculations
show that it would take 1 year for a molecule to diffuse 25 cm through cells (Niklas
1997).
The
conducting cells of Phloem are called Sieve Elements. There are two kinds of these,
namely: Sieve Cells and Sieve Tube Members (STM). This is analogous to the situation in Xylem.
Both types of Sieve Elements have Sieve Pores lined with Callose and both are Enucleate at maturity. Callose is a polysaccharide that is involved in the
regulation of the Sieve Pore diameter. It is a biochemical
hallmark of Sieve Elements.
STM are
rectangular and have many sieve pores on their end walls which are called Sieve Plates.
Sieve Cells
are elongate with tapered end walls. They lack Sieve Plates as their Sieve Pores are more
widely distributed along their length. Sieve Cells have a smaller diameter than STM, and
are less efficient at translocation.
Sieve Cells are ancestral and STM are advanced. The Magnoliophyta typically have STM while most other plants have Sieve Cells in their Phloem. A stack of STM
make a Sieve Tube!
This helps to explain why the largest
Bryophyta had "Vascular Tissues". The Leptoids of Bryophytes have most of the traits of Sieve Elements from typical
Vascular Plants, including Callose.