This paper originates from a presentation at the International Conference on Assimilate Transport and Partitioning, Newcastle, NSW, August 1999
Carbohydrates are synthesised in photosynthetically active source tissues and
exported, in most species in the form of sucrose, to photosynthetically less
active or inactive sink tissues. Sucrose hydrolysis at the site of utilisation
contributes to phloem unloading. This phenomenon links sink metabolism with
phloem transport to, and partitioning between, sinks. Invertases catalyse the
irreversible hydrolysis of sucrose and thus are expected to contribute to
carbohydrate partitioning. Different invertase isoenzymes may be distinguished
based on their intracellular location, their isoelectric points and pH optima.
Extracellular, cell-wall-bound invertase is uniquely positioned to supply
carbohydrates to sink tissues via an apoplasmic pathway, and links the
transport sugar sucrose to hexose transporters. A number of studies
demonstrate an essential function of this invertase isoenzyme for phloem
unloading, carbohydrate partitioning and growth of sink tissues. Extracellular
invertases were shown to be specifically expressed under conditions that
require a high carbohydrate supply to sink tissues. Further, their expression
is upregulated by a number of stimuli that affect source–sink relations.
Substrate and reaction products of invertases are not only nutri-ents, but
also signal molecules. Like hormones and in combination with hormones and
other stimuli, they can regu-late many aspects of plant development from gene
expression to long-distance nutrient allocation. Based on studies in
Chenopodium rubrum, tomato
(Lycopersicon esculentum) and tobacco
(Nicotiana tabacum), the regulation of extracellular
invertase and its function in assimilate partitioning, defence reactions and
sugar signal transduction pathways are discussed.