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MOLOISE T.

(201404014)
BIO 316 ASSIGNMENT.
21-09-2016
THE PHLOEM Loading AND
UNLOADING

INTRODUCTION
Plants are in need of sugars for energy and other cell activities, therefore they photosynthesize. In
vascular plants, phloem is the living tissue that carries organic nutrients (known as
photosynthate), in particular, sucrose (a sugar), to all parts of the plant where needed. Phloem
tissue consists of: conducting cells, generally called sieve elements; parenchyma cells, including
both specialized companion cells or albuminous cells and unspecialized cells; and supportive
cells, such as fibers and sclereids (Sharkey, T.D., 2005).
Sieve elements are the type of cell that are responsible for transporting sugars throughout the
plant. At maturity they lack a nucleus and have very few organelles, so they rely on companion
cells or albuminous cells for most of their metabolic needs. Sieve tube cells do contain vacuoles
and other organelles, such as ribosomes, before they mature, but these generally migrate to the
cell wall and dissolve at maturity; this ensures there is little to impede the movement of fluids.
One of the few organelles they do contain at maturity is the smooth endoplasmic reticulum,
which can be found at the plasma membrane, often nearby the plasmodesmata that connect them
to their companion or albuminous cells. All sieve cells have groups of pores at their ends that
grow from modified and enlarged plasmodesmata, called sieve areas. The pores are reinforced by
platelets of a polysaccharide called callose (Cousson, A. & Vavasseur, A., 1998).
Since photosynthesis takes place primarily on the leaves, the carbohydrates products are
transported to other nearer veins via diffusion and to distant parts of the plants via active
transport into the sieve tube elements of the phloem. The movement of photosynthates on the
phloem is called translocation, and contents on the phloem is called phloem sap. The
photosynthates are moved from the source to the sink. A source is an organ (such as mature leaf
or a storage root) that produces, by photosynthesis or digestion of stored reserves, more sugars
than they require. A sink is an organ (such as a root, flower, developing fruit, or immature leaf)
that consumes sugars from its own growth and storage needs of the plant.
A given organ can be a sink at some times and a source at other times. An example is a root of
male trees, which stores sugars sent down from the leaves during one growing season, then send
the sugars back upward to support the emergence of new leaves the following spring.

DISCUSSION
The phloem as flows under positive pressure through the sieve tubes ( A. M. & Atkinson, C. J.
1990). It moves by bulk flow (i.e,from a region of higher pressure potential to a region of a lower
pressure potential) from one sieve tube to the next. Two steps in translocation require metabolic
energy:

Transport of sucrose and other solutes from the into the sieve tubes; called loading
Removal of the solutes from the sieve tubes into the sink; called unloading

According to the pressure flow model of translocation in the phloem, sucrose is actively
transported into sieve tube elements at a source, giving those cells a greater sucrose
concentration than the surrounding cells. Therefore the cells develop a lower solute potential-a
more negative s at which point water enters the sieve tube elements from the xylem vessels by
osmosis. The entry of this water causes a greater pressure potential (turgor pressure) at the source
end of the sieve tube, so that the entire fluid content of the sieve tube is pushed toward the sink
end of the tube-in other words, the sap moves by bulk flow in response to a pressure gradient. In
the sink, the sucrose is unloaded both passively and by active transport, and water moves back to
the xylem vessels. In this way the gradients of solute potential and pressure potential, which are
needed for the movement of the phloem sap, are maintained (Sharkey, T.D., 2005).

Sugars and other solutes move from the mesophyll cells in the leaf to the phloem by two
general routes; apoplastic and symplastic. The exact detail of these routes vary widely
among plants species. In many plants, sugars and others solutes follow apoplastic
pathway: solutes move out of the mesophyll cells and then diffuse through the apoplast to
the sieve tubes. Specific sugars and amino acids are then actively transported into the
sieve tube elements. Active transport across membranes allows plants to regulate which
specific substances enter the phloem. In contrast, in the symplastic pathway, solutes
remain within the symplast and pass through plasmodemata all the way from the
mesophyll cells to the sieve tubes cells. Because no membranes crossed in the symplastic
pathway, the solutes are loaded into the phloem by mechanisms other than active
membrane transport. Upon arriving in the sink regions, solutes are actively transported
out of the sieve tube elements and into the surrounding tissues. This unloading serves
several purposes: it helps maintain the gradient of solute potential and hence the pressure
potential on the sieve tubes; it supplies carbohydrates and amino acids to developing
organs , where they are used for growth; and it helps build up high concentrations of the
proteins and carbohydrates in the storage organs such as storage roots, fruits, and seeds.
The demand for fixed carbon can be very large in some tissues, such as rapidly growing

stem or a developing potato tuber. These tissues receive a lot of phloem sap, while a
slowly growing organ receive less (Spreitzer, R.J., M.E. Salvucci, 2002).

Evidence has been outlined by different plant biologists to show that the phloem carries
photosynthates from the source to the sink using many ways. The evidence was first
obtained by an Italian scientist Marcello Malpighi in 1600s. He simply removed a ring
bark from a tree trunk, and the upper part swell towards the groove, and he concluded
that solutes were trapped as the phloem was cut. Other scientists used aphids to precisely
analyze the phloem sap contents of the phloem. Aphids are insects that feed on plants by
drilling into sieve tube elements with a specialized organ, the stylet. The pressure
potential in the sieve tubes is higher than that outside the plant, so the phloem contents
are forced through the stylet into the aphids digestive tract. So great is the pressure that
some of the liquid is forced through the insects body and out of its anus, and can be
collected for analysis (Sharkey, T.D., 2005).
CONCLUSION
Translocation is the movement of photosynthates from the source to the sink through a phloem
tissue. Transpiration pulls water up xylem vessels; Sucrose cells load sucrose into phloem sieve
tubes via the companion cells, reducing water potential, so water is taken up from xylem vessels
by osmosis, raising the pressure potential in the sieve tubes. Internal pressure differences drive
the sap along the sieve tube to sink cells. Sucrose is unloaded into the sink cells, increasing water
potential in the sieve and hence the water moves back into the xylem.

REFERENCES
Sharkey, T.D., 2005. Effects of moderate heat stress on photosynthesis: importance of thylakoid
reactions, rubisco deactivation, reactive oxygen species, and thermotolerance provided by
isoprene. Plant Cell Environ., 28, 269-277.
Spreitzer, R.J., M.E. Salvucci, 2002. Rubisco: structure, regulatory interactions, and
possibilitiesfor a better enzyme, Annu. Rev. Plant Biol., 53, 449-475.
Cousson, A. & Vavasseur, A. (1998) Planta 206, 308314. 36. Mansfield, T. A., Hetherington,
A. M. & Atkinson, C. J. (1990) Annu. Rev. Plant Physiol. Plant Mol. Biol. 41, 5575

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