Gn. Thomopoulos et al., Ultrastructural histochemical studies of secretory processes in rat submandibular granular tubules during intermittent sympathetic nerve stimulation, EUR J MORPH, 38(2), 2000, pp. 69-79
Secretory changes in the cells of granular tubules in rat submandibular gla
nds have been studied sequentially during electrical stimulation of their s
ympathetic nerves. Results were assessed in a series of biopsied lobes from
the same gland, taken at different times during the sympathetic stimulatio
n. Changes were not synchronous between adjacent cells and it appeared that
the time for the onset of secretory events differed between cells but, onc
e set in action, a chain of similar events occurred. Nevertheless, some cel
ls appeared to remain refractory throughout. Initially, some alignment of g
ranules to the adjacent plasma membrane occurred and occasional evidence fo
r classical exocytosis was seen. However, from early on microvesicles appea
red in more luminally located granule membranes and were associated with gr
anule fusions, that became common and led to the formation of large irregul
ar aggregates. Most of the secretion of granule contents appeared to be thr
ough openings of aggregates into lumina. With granule fusions the intra-mem
brane microvesicles became internalised and tended to increase in size with
time; they were commonly expelled with the contents of the aggregates. Fra
gments of cytoplasm also became incorporated in aggregate formation. Cytopl
asm, often containing glycogen, also formed luminal blebs over some granula
r tubule cells and appeared to pass into the secretion by an apocrine proce
ss. At the end of stimulation multivesicular bodies were seen in associatio
n with redundant aggregates.