title: Ultrastructural evidence for a dual function of the phloem and programmed cell death in the floral nectary of Digitalis purpurea.
abstract: OBJECTIVE
  The floral nectary of Digitalis purpurea is a transitory organ with stomatal exudation of nectar. In this type of nectary, the nectar is thought to be transported to the exterior via intercellular ducts that traverse the nectariferous tissue. The latter is also traversed by a ramified system of phloem strands from which prenectar sugar is most probably unloaded. The aims of this study were to provide some of the basic information needed to evaluate the possible mechanism involved in nectar secretion and to discover the fate of the nectary.
  
  
  METHODS
  The ultrastructure of the nectary was investigated at different stages of development by analysis of a series of ultrathin (7 x 10(-8) m) sections 7 x 10(-7) m apart from one another. Proportions of the cells typical of the nectary were documented by 3D-reconstruction and morphometry.
  
  
  RESULTS
  The phloem consisted of variably shaped sieve elements and companion cells which, as a rule, were more voluminous than the sieve elements. Direct contact between the phloem strands and intercellular ducts was observed. In contrast to the phloem, which remained structurally intact beyond the secretory phase, the nectariferous tissue exhibited degenerative changes reminiscent of programmed cell death (PCD), which started as early as the onset of secretion and progressed in a cascade-like fashion until final cell death occurred in the exhausted nectary. Hallmarks of PCD were: increased vacuolation; increase in electron opacity of individual cells; progressive incorporation of plasmatic components into the vacuole reminiscent of autophagy; degradation of plastids starting with hydrolysis of starch; deformation of the nucleus and gradual disappearance of chromatin; loss of tonoplast integrity and subsequent autolysis of the rest of cellular debris. Degeneration of the cells occurred against a background of increasing cell size.
  
  
  CONCLUSIONS
  The cytological and anatomical evidence presented here, and calculations of the solute fluxes necessary for accumulation of starch and for the production of nectar support the view that: (a) in the foxgloves' nectary, apoplastic phloem unloading dominates, at least during exudation of nectar; (b) the obsolete nectary may be dismantled by PCD; and (c) at least the products of late nectary degradation are loaded via the apoplast into the unchanged phloem and exported to sinks elsewhere in the plant for reallocation.
authors.affiliation: Institut für Biologie, Lehrstuhl für Pharmazeutische Biologie der Friedrich-Alexander-Universität Erlangen-Nürnberg, Staudtstrasse 5, D-91058 Erlangen, Germany. kgaffal@biologie.uni-erlangen.de
authors.forename: Karl Peter
authors.initials: KP
authors.lastname: Gaffal
authors.forename: Gudrun Johanna
authors.initials: GJ
authors.lastname: Friedrichs
authors.forename: Stefan
authors.initials: S
authors.lastname: El-Gammal
mesh_terms: D017209:Apoptosis / Q000502:physiology*; D004070:Digitalis / Q000166:cytology / Q000502:physiology* / Q000648:ultrastructure; D035264:Flowers / Q000166:cytology / Q000502:physiology / Q000648:ultrastructure; D021621:Imaging, Three-Dimensional; D046529:Microscopy, Electron, Transmission; D052585:Phloem / Q000166:cytology / Q000502:physiology* / Q000648:ultrastructure