The neuromuscular transform in a single segment of a segmented heart tube.

Leech hearts are hybrids; they are myogenic but need entrainment by a heartbeat central pattern generator (CPG) to execute functional constriction patterns. Leech hearts are modular: two lateral segmented heart tubes running the length of the animal. Moving blood through the segmented heart tubes of leeches requires sequential constrictions, timed by a heartbeat CPG and relayed to each heart segment by likewise segmental motor neurons. The heartbeat CPG produces bilaterally asymmetric coordinations: rear-to-front peristaltic on one side and nearly synchronous on the other, periodically switching sides. We examined the neuromuscular transform of isolated heart segments in response to electrical nerve stimulation to identify the range of parameters (burst duration, intraburst pulse frequency, period) allowing the heart to constrict continuously and reliably. Constriction amplitudes increased with increasing intraburst frequencies and decreased with decreasing burst durations. Similar amplitudes were achieved with longer burst durations combined with lower frequencies or with shorter burst durations combined with higher frequencies. Long burst durations delayed relaxation, leading to summation and tetanus. The time, and its variability, between stimulus onset and time to constriction onset or to peak decreased with increasing frequency. Data previously obtained in vivo showed that the heart excitatory motor neurons fired longer bursts at lower frequencies at long periods moving to shorter bursts with higher intraburst frequencies as the period shortened. In this scenario, active constriction started earlier and the time to reach full systole shortened, allowing more time for relaxation. Relaxation time before the next motor neuron burst appears critical for maintaining constriction amplitude. Moving blood through the segmented heart tubes of leeches requires sequential constrictions driven by motor neurons controlled by a central pattern generator. In a single heart segment, we varied stimuli to explore the neuromuscular transform. Decreasing the cycle period, e.g., to increase volume pumped over time, without altering motor burst duration and intraburst spike frequency shortens relaxation time and decreases amplitude. The likely strategy to preserve constriction amplitude is to shorten burst duration while increasing spike frequency.