Innervation of the heart and aorta of Manduca sexta.

Innervation of the heart and aorta of Manduca sexta was studied by using anatomic, neuronal tracing and immunocytochemical techniques. The study was undertaken to provide a foundation for investigating the neural mechanisms controlling cardiac reversal in adults. Lateral cardiac nerves were not found in the larval or adult heart. The larval heart and aorta seem to lack innervation, but a neurohemal system for the release of a cardioactive peptide is associated with the larval alary muscles. At adult metamorphosis, this neurohemal system regresses, and, at the same time, processes grow onto the anterior aorta. These processes seem to be neurohemal and originate from two pairs of neurosecretory cells located in the subesophageal ganglion. This system is immunoreactive to cardioactive peptides and may function, therefore, in hormonal modulation of the activity of the adult heart. Also during metamorphosis, synaptic innervation develops on the terminal heart chamber, and this innervation is from axons extending through the seventh and eighth dorsal nerves of the terminal abdominal ganglion. These axons originate from cells that have been identified as serial homologs of motor neuron-1 of other abdominal ganglia. These neurons are immunoreactive to a cardioactive peptide, and this peptide probably modulates the synaptic innervation of the terminal heart chamber. During metamorphosis, the target of the motor neurons-1 of the seventh and eighth segments becomes respecified from larval skeletal muscles to the terminal chamber of the adult heart.

Neuronal control of heart reversal in the hawkmoth Manduca sexta.

Cardiograms demonstrate that heart activity of Manduca sexta changes from larva, to pupa, to adult. The larval heart has only anterograde contractions. During metamorphosis, heart activity becomes a cyclic alternation of anterograde and retrograde contractions. Thus, the adult heart has both an anterograde and a retrograde pacemaker. External stimuli also can initiate cardiac reversal. Cardiac reversal is blocked by tetrodotoxin, indicating that reversal is under neuronal control. A branch of each dorsal nerve 8 innervates the posterior chamber of the heart, the location of the anterograde pacemaker. Only retrograde contractions occur when dorsal nerves 8 are cut. Stimulation of ml(-1) 8 initiates anterograde contractions; when stimulation ceases, the heart reverses to retrograde contractions. These experiments indicate that the anterograde pacemaker receives neural input that makes it the dominant pacemaker. In the absence of neural input this pacemaker is inactive, and the retrograde pacemaker becomes active. Application of crustacean cardioactive peptide accelerates the heart but does not eliminate cardiac reversal. The terminal chamber of the heart is also innervated by a branch of each dorsal nerve 7; stimulation of this nerve increases the strength of contraction of the terminal chamber but has no effect on contractions of the remainder of the heart or on cardiac reversal.

Functional morphology of the dorsal vessel in the adult fly Protophormia terraenovae (diptera, calliphoridae).

The morphology and ultrastructure of the contractile tubular vessel acting as the cardiac pump in Protophormia terraenovae flies were analyzed by means of light microscopy, SEM and TEM. The results provide a novel anatomical picture of the two vessel portions, the abdominal heart and the aorta, and lay the foundations for an interpretation of the cardiocirculatory function in the fly. In the thorax, the thin and unchambered aorta is without apertures, while the abdominal heart presents a very small caudal aperture and pairs of lateral ostia, one in each of the five chambers of which it is composed. The ostia of the four more distal chambers are of the incurrent type, which is to say that they act as valves ensuring that hemolymph flows only into the heart. Conversely, the ostia in the most proximal chamber allow hemolymph to flow both into and out of the heart. The entire vessel is composed of a single layer of myofibers that are oriented circularly around the lumen in the abdominal heart and longitudinally in the thoracic aorta. The abdominal heart has a thicker wall, a far more diffused and thick distribution of tracheoles, and a far greater number of mitochondria with respect to the aorta. This arrangement ensures a greater availability of oxygen and energy in the abdominal heart compared to the aorta and leads one to suppose that the high- and low-frequency contractions of the cardiac cycle (Thon, [1982] J. Insect Physiol. 28:411-416) can be attributed to the abdominal heart and the aorta, respectively. J. Morphol. 240:15-31, 1999. © 1999 Wiley-Liss, Inc.