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Journal of Experimental Biology, Vol 199, Issue 4 749-758, Copyright © 1996 by Company of Biologists
JOURNAL ARTICLES |
J Truman, J Ewer and E Ball
A grasshopper hatches from its egg, which is laid in soil, as a vermiform larva. This larva continues the stereotyped hatching behaviour as it digs through the egg pod, which provides a passageway to the soil surface. Once at the surface, shedding, or ecdysis, of the vermiform cuticle is initiated. When this process is complete, the first-instar cuticle is expanded to assume the form of the first-instar hopper. We have demonstrated, using immunocytochemical techniques, that these behaviour patterns are associated with dramatic increases in intracellular levels of cyclic GMP in sets of identified neurones in the ventral central nervous system. The most prominent cyclic-GMP-expressing cells are 34 neurones that appear to contain crustacean cardioactive peptide (CCAP). These CCAP cells show no detectable cyclic GMP at hatching or while the vermiform larva digs through the soil. Upon reaching the surface and freeing itself, the larva initiates ecdysis and associated air-swallowing and tracheal filling within about 1 min. These changes are immediately preceded by the appearance of cyclic GMP in the CCAP cells. Cyclic GMP levels in these neurones peak by 5 min and then decline back to basal levels by 20-30 min. Conditions that cause ecdysing animals to resume digging prolong the elevation of cyclic GMP levels. Once animals have assumed their 'hopper' form, however, external stimuli can no longer affect the time course of the cyclic GMP response. The neurones containing elevated cyclic GMP levels probably influence the air-swallowing, tracheal filling and circulatory changes that are associated with ecdysis behaviour. Pairs of descending midline neurones in abdominal segments 2-4 also become cyclic-GMP-immunoreactive, but they show peak expression after cyclic GMP levels in the CCAP cells have declined. Also, neurones in the caudolateral region of the abdominal ganglia often become cyclic-GMP-immunoreactive when ecdysing animals are forced to resume digging for an extended period.
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