Phloem loading and transport

How do the products of photosynthesis travel from sites of carbon fixation

to sites of carbon and energy utilization and storage?

Phloem
Phloem anatomy
Phloem
Phloem sieve elements conduct material
• Sieve tube elements (angiosperms)
• Sieve cells (gymnosperms)
Companion cells (support sieve element functions)
Parenchyma (uptake, storage, release of food)
Sclereids/sclerenchyma (strength, support, protection)
Laticifers (latex conduits)
Bundle sheath surrounding vascular tissue (not just in C4 plants)
1º and 2º phloem
10.1 Transverse section of a vascular bundle of trefoil, a clover ( Trifolium)

10.2 Transverse section of a 3-year-old stem of an ash (Fraxinus excelsior)

tree

Phloem anatomy
Phloem sieve tube elements are living at maturity
Relatively few organelles (lack nuclei, vacuoles)
Few ribosomes, cytoskeletal elements
Thickened primary cell wall, but no lignified secondary cell wall
Sieve plate (angiosperms) and lateral sieve areas
10.3 Schematic drawings of mature sieve elements (sieve tube elements)
Transverse section of ordinary companion cells and
mature sieve tube elements
10.5 Sieve elements and open sieve plate pores
Sieve plate pores between sieve tube elements can be
very large
(1-15 μm)
10.6 Electron micrograph of a sieve area (sa) linking two sieve cells of a
conifer (Pinus resinosa)
Gymnosperm sieve areas have much smaller pores; no
sieve plates

Pores are not open, but occluded and traversed by

smooth ER

No P-proteins
Phloem protection
Angiosperm P-proteins are diverse proteins that clog pores when turgor
released rapidly
Wound callose (β-1,3-glucan) is produced to seal pores of functional sieve
tube elements from damaged cells

Companion cells
Tightly linked through numerous complex plasmodesmata to adjacent sieve
tube element
Perform many of the metabolic functions of the organelle-deficient sieve
tube elements
Three basic types:
Ordinary companion cells

• Plasmodesmata almost
exclusively to sieve tube
elements
• Chloroplasts
10.7 Electron micrographs of companion cells in minor veins of mature
leaves (Part 1)

Companion cells

Transfer cells
• Similar to ordinary companion
cells
• Cell wall invaginations yielding
increased PM surface area
• Transfer from apoplast (cell wall
space) to symplast/phloem
10.7 Electron micrographs of companion cells in minor veins of mature
leaves (Part 2)

Companion cells

Intermediary cells
• Abundant plasmodesmata
connections to bundle sheath
cells
• Poorly developed chloroplasts
The companion cell types may be emphasized differently in different
tissues or species
10.7 Electron micrographs of companion cells in minor veins of mature
leaves (Part 3)

Phloem links sources to sinks

Photosynthate sources

Sites of net export

• Mature leaves
• Storage tissues during some
phases of development
Photosynthate sinks

Sites of import
• Rapidly growing regions
(meristems, young leaves)
• Non-photosynthetic tissues
(roots, floral organs, fruits)
• Storage tissues during some
phases of development
10.8 Source-to-sink patterns of phloem translocation
--Proximity may determine which sources feed which
sinks
--Development may shift source-sink relationships
--Source-sink connections may follow direct
connections along the plant body
--Vascular interconnections (anastomoses) can
circumvent blockages or wounds
Phloem sap
Collected through aphid stylets and analyzed:
Sugars (especially sucrose and non-reducing di-, tri- and small oligo-
saccharides)
Amino acids (especially Glu, Gln, Asp, Asn) and other nitrogenous
compounds
Organic acids, mineral nutrients and ions
Hormones
Proteins & RNA (regulatory, defense, stress responses)

10.9 Structures of compounds both not normally and commonly

translocated in the phloem (Part 1)

10.9 Structures of compounds both not normally and commonly

translocated in the phloem (Part 2)

10.9 Structures of compounds both not normally and commonly

translocated in the phloem (Part 3)

Phloem movement
Much too fast for diffusion—bulk flow required
Pressure flow model
Driven by differences in osmotic pressure between sources and sinks
Phloem loading and accompanying water uptake at source
Phloem unloading and reduced osmotic potential at sink
10.10 Pressure-flow model of translocation in the phloem

10.11 Translocation in living, functional sieve elements of a leaf (Part 1)

10.11 Translocation in living, functional sieve elements of a leaf (Part 2)

Leaf vein architecture

Phloem loading
Transport of photosynthates from mesophyll to sieve tube elements:
Chloroplasts to cytosol as triose phosphates
Conversion to hexose phosphates and then to sucrose (fructose +
glucose) in the cytosol
Short-distance transport of sucrose to small veins
Concentration in the companion cells and sieve elements
Export and long-distance transport to sinks
Symplastic vs. Apoplastic loading is species-specific
10.14 Schematic diagram of pathways of phloem loading in source leaves
(Part 1)

10.14 Schematic diagram of pathways of phloem loading in source leaves

(Part 2)

10.15 Labeled sugar moves from the apoplast into sieve elements and
companion cells
Sucrose is actively transported into the phloem
Phloem loading is product specific

10.16 ATP-dependent sucrose transport in sieve element loading

Active sucrose transport involves a H - +

ATPase and a H+-sucrose symporter

(AtSUC2) for uptake from the apoplast to
the symplast
Phloem loading
In species utilizing the symplast pathway, the polymer-trapping model
explains net movement into the phloem
Sucrose from mesophyll diffuses into intermediate cells
Addition of galactoses (e.g. raffinose, stachyose) prevent them from
diffusing back
These larger sugars (as well as sucrose) can diffuse into the phloem
and move by bulk flow down the sieve tubes
10.17 Polymer-trapping model of phloem loading
Phloem unloading
Sugars move from phloem to sink tissues
Short-distance transport into and through local cells may be symplastic or
apoplastic
Storage/metabolism in the sink tissue cells
Requires energy
10.18 Pathways for phloem unloading and short-distance transport

Source-sink
relationships change
Sinks may gradually transition to sources
Young leaves to mature leaves
Storage organs
10.19 Autoradiographs of a leaf of summer squash ( Cucurbita pepo)
Developmental transition from sink to
source
10.20 Division of labor in the veins of a tobacco leaf
Different sets of veins are used for unloading (as a
sink) versus loading (as a source)
Immature (sink) minor veins not used for
unloading
Minor veins mature (source) and can be used for
export.
Figure 10.21 Export from source tissue depends on active sucrose
transporters
Sugar allocation
In sources, fixed carbon may be
allocated for
Storage
Metabolism
Export
Sources regulate allocation to storage
or export via triose phosphate/Pi
chloroplast antiporter and
regulation of starch- and sucrose-
synthesizing enzymes
In sinks, imported sugars may be
allocated for metabolism or storage
Determination of which sink tissues
receive photosynthates is
partitioning
10.21 A simplified scheme for starch and sucrose synthesis during the day

Photosynthate partitioning
Regulated dynamically via complex mechanisms
Sink strength (size and activity)
Turgor
Overall sucrose levels
 Hormones
Gene expression (sucrose metabolism, transporters, etc.)
Other molecules move in the phloem stream
Proteins (e.g. pathogenesis-related proteins)
RNAs (mRNAs, small gene-silencing miRNAs, pathogen RNA,
ribonuclear proteins)
Hormones, remobilized minerals, etc.
10.22 GFP fluorescence in source and sink leaves from transgenic
Arabidopsis plants
SUC2 promoter-driven expression of GFP in source
and sink leaves shows plasmodesmatal
movement of protein from source to sink tissues