Sectoriality and physiological organisation in herbaceous plants: an overview

The physiological organisation of plants is considered in relation to the carbon economy of plant parts. Although assimilate is partitioned according to the relative strength of sinks, in many species there is also a very close relationship between partitioning and shoot phyllotaxy, giving rise to sectorial patterns of allocation whereby only certain sinks are supported by any source leaf. Essentially these sinks are in the same orthostichy as the source leaf. This constraint of the vascular architecture on assimilate distribution to developing sinks such as leaves, flowers and fruits is not always absolute, as following the loss of their principal source leaves these sinks can in many cases be supplied with assimilate by other leaves via new inter-orthostichy pathways. The supply of assimilate to major sinks such as developing fruits becomes more and more localised with time so that a fruit in an axillary position becomes largely supported by its subtending leaf; the reproductive node—a metamer-can thus be regarded as a relatively autonomous unit of the plant (an IPU). Similary, once established after a developmental phase of assimilate import, tiller ramets and branches in unitary plants tend to become physiologically autonomous modules. However, the functional autonomy of tillers is reversed following defoliation or shading as they are then sustained by the import of assimilate, subject to its availability, from unaffected tillers. Consequently the plant becomes physiologically integrated by the flow of assimilate from one part to another. The mainly autonomous ramets of many stoloniferous and rhizomatous species display a similar pattern of physiological integration in response to source manipulation, but in some species the ramets appear to maintain their independent functioning as a normal feature of the carbon allocation within the clone. In other clonal species, as the clone develops and becomes more structurally complex, vascular constraints start to restrict the movement of resources, and the clone becomes composed of a number of semi-autonomous IPUs. In unitary plants branches appear to remain very physiologically isolated in terms of their carbon economy once they become established, irrespective of a range of source-sink manipulations.These different patterns of physiological integration and organisation are discussed in relation to different strategies of assimilate utilisation and conservation.