It is becoming increasingly apparent that the longâdistance signalling associated with many developmental processes is complex and that novel hormoneâlike signals may play substantial roles. The past decades have seen several substances (e.g. brassinosteroids, systemin and other polypeptides, mevalonic and jasmonic acids, polyamines, oligosaccharides, flavonoids, and quinones) vie for a place among the classical plant hormones (e.g. Spaink, 1996). Recent microinjection and grafting studies have also shown that RNA may act as a longâdistance signal (Jorgensen et
al., 1998; XoconostleâCĂĄzares et al., 1999). In this issue,
Hannah et al. describe longâdistance signalling and the
regulation of rootâshoot partitioning in dwarf lethal or
dosageâdependent lethal (DL) mutants of common bean
(Shii et al., 1980, 1981), and present evidence indicating that substances in addition to classical plant hormones (e.g. cytokinins) may be involved.
As in the report by Hannah et al., much of the evidence for roles of unidentified longâdistance signals in the control of plant development is indirect. The possibility that a small number of longâdistance signals might control a multitude of developmental processes arises through the potential for differences in tissue sensitivity, fluctuations in hormone levels and differences in the nature of responses of different tissues to the same hormone. Consequently, particular hormones may influence numerous processes seemingly simultaneously, yet independently. Even so, longâdistance signalling is involved in processes as diverse as rootâshoot balance, senescence, branching, flowering, nodulation, stress responses and nutrient uptake. Through comparison of even a few different developmental processes, progress can be made to reveal the true complexity of plant development. Using this approach it is also clear that many unknown signals may be involved.