(a) Source processes

Fig_p_5.12.png

Figure 5.2 Time course of sucrose and phosphorus (P) net import and export from a leaf during its development. As a cucumber leaf expands, net sucrose export coincides with the rise in net leaf photosynthetic rate (O) to meet photoassimilate demands of young leaves. Once a leaf has reached some 30% of its final area, net photosynthesis by the whole leaf exceeds photoassimilate demand by growth and so excess sucrose can be exported. Thereafter, the rate of sucrose export closely follows photosynthetic rate, reaching a maximum when the leaf reaches its final size and gradually declining thereafter. Import of P (and other mineral nutrients) continues throughout leaf expansion and P export only starts once the leaf is fully expanded. Sucrose import and export were calculated from the difference between rates of whole-leaf photosynthesis and dry matter gain (Based on Hopkinson 1964; reproduced with permission of Journal of Experimental Botany)

Net export of photoassimilates occurs from fully expanded leaves (Figure 5.2) and long-term storage pools located along the axial transport pathway. Chloroplasts of C₃ plants (Chapters 1 and 2) partition photoassimilates between the photosynthetic oxidative cycle and starch biosynthesis or release them immediately to the cytosol as triose phosphate for sucrose synthesis. In non-starch-forming leaves, high concentrations of sugars can be accumulated in the vacuoles of mesophyll cells or made available for immediate loading into the phloem and export. Leaves also serve as secondary sources for nutrients and amino acids previously delivered in the transpiration stream. Nutrients and amino acids can be exported in the phloem immediately, or after accumulation in short-term storage pools.

An additional source of photoassimilates is located along the axial phloem path (petioles, stems, peduncles, pedicels and roots) as a result of leakage from the vascular tissues. Leaked photoassimilates accumulate in short- or long-term storage pools which serve as secondary sources to buffer photo-assimilate supplies to the sinks against shifts in export rates from the primary photoassimilate sources.

(b) Path processes

Assimilates including sucrose, amino acids are transferred into sieve elements of fully expanded leaves against significant concentration and electrochemical gradients. This process is referred to as phloem loading. The cellular pathways of phloem loading, and hence transport mechanisms and controls, vary between plant species. Longitudinal transport of assimilates through sieve elements is achieved by mass flow and is termed phloem translocation. Mass flow is driven by a pressure gradient generated osmotically at either end of the phloem pathway, with a high concentration of solutes at the source end and a lower concentration at the sink end. At the sink, assimilates exit the sieve elements and move into recipient sink cells where they are used for growth or storage. Movement from sieve elements to recipient sink cells is called phloem unloading. The cellular pathway of phloem unloading, and hence transport mechanisms and controls, vary depending upon sink function.

(c) Sink processes

Many sink organs are characterised by low rates of transpiration (an exception is a developing leaf) so that most assimilates are delivered by the phloem. Having reached the sink cell cytoplasm through the post-sieve-element transport pathway, assimilates are either metabolised to satisfy the energy, maintenance and growth requirements of sink cells or are compartmented into polymer or vacuolar storage. Collectively, metabolism and compartmentation create a demand for assimilates which is ultimately responsible for driving phloem import.

(a) Whole-plant growth

Sink and source strength must be in balance at a whole-plant level. Thus, an increase in whole-plant sink strength must be matched by an equal increase in source strength, either through increases in source activity or source size. Prior to canopy closure in a crop, much of the increase in source strength comes from increased source size, source activity remaining relatively constant. Significantly, until a leaf has reached some 30% of its final size, photoassimilates for leaf production are exclusively imported through the phloem from fully expanded leaves (Figure 5.2).

(b) Photoassimilate transport and crop yield

During domestication of crop plants, plant breeders selected for crop yield via maximum investment into harvested organs (mostly seeds). Total plant biomass production of advanced wheat is the same as its wild progenitors yet grain yield has increased some 30-fold through breeding. That is, whole-plant source and sink strength have not changed. Increases in wheat yield are associated with a diversion of photoassimilates from vegetative organs to the developing grain, as illustrated by the relative accumulation of ¹⁴C photoassimilates exported from the flag leaf.

Final grain yield is not only determined by partitioning of current photoassimilates, but also depends upon remobilisation of non-structural carbohydrates stored in stems, particularly under conditions where environmental stress impairs leaf photosynthesis (Wardlaw 1990). In fact, remobilisation of reserves affects yield in many food plants. For example, deciduous fruit trees depend entirely on remobilised photo-assimilates to support flowering and fruit set as do early stages of pasture regrowth following grazing.

Fig 5.14.jpg

Figure 5.3 Photoassimilate distribution in a rooted cutting of Washington Navel orange (mounted specimen shown on left; matching autoradiograph on right). ¹⁴CO₂ was supplied to source leaves (boxed area top left) for a day, and movement of ¹⁴C-labelled assimilate followed by autoradiography of harvested plant material. ¹⁴C photosynthates were distributed widely via vascular conduits to sinks including some roots and a fruit on an adjacent shoot (note stem labelling between sources and sinks). Nearby mature leaves failed to import; they were additional sources of photosynthate. Scale bar = 2 cm (Unpublished material courtesy P.E. Kriedemann)

Photoassimilate transport to harvestable organs plays a central role in crop yield brought about by greater harvest indices. This raises questions about transport and transfer processes that collectively influence photoassimilate partitioning between competing sinks.

Historically, these questions were elucidated by observing partitioning patterns of photoassimilates exported from specified source leaves labelled with ¹⁴C supplied as a pulse of ¹⁴CO₂. Following a chase period, in which ¹⁴C photoassimilates are transported to and accumulated by recipient sink organs, the plant is harvested. The pattern of photoassimilate partitioning operating during the pulse is deduced from ¹⁴C activity accumulated by sinks (Figure 5.3).

Photoassimilates are partitioned from source leaves to sinks in characteristic and reproducible patterns. For instance, in a vegetative plant, lower leaves are the principal suppliers of photoassimilate to roots, whereas upper leaves are the principal suppliers to the shoot apex. Leaves in an intermediate position export equal quantities of photoassimilates in either direction. However, the pattern of photoassimilate partitioning is not static, it changes with plant development. In vegetative plants, the direction of flow from a leaf changes as more leaves above it become net exporters. Furthermore, at the onset of reproductive development, growing fruits or seeds become dominant shoot sinks for photoassimilates at the expense of vegetative apices.

Photoassimilate partitioning patterns can be altered experimentally by removal of selected sources (e.g. leaves) or sinks (e.g. fruits). These manipulative experiments demonstrate that photoassimilate partitioning reflects the relative strengths of individual sources and sinks. Properties of the phloem pathway connecting sources with sinks are shown in the following Section 5.2.