Functional analyses of the leech swim oscillator.

The oscillations that underlie swimming movements in the leech arise from a series of identified concatenated circuits within the ventral nerve cord. In the intact nerve cord, ascending and descending intersegmental interactions via axons within the lateral connectives function both to generate robust oscillations throughout the cord and to establish an anterior-to-posterior phase delay among segmental oscillators. We addressed two questions about this system. First, do the intrasegmental swim circuits in each ganglion function as a single oscillator or do they comprise a pair of coupled oscillators? Second, what are the relative strengths of the ascending and descending intersegmental interactions between the segmental oscillators? Experiments were carried out on semi-intact leeches (Hirudo medicinalis) and on isolated leech nerve cords in which "Z-cut" ganglia were generated by cutting one lateral connective nerve anterior and the contralateral connective nerve posterior to the target ganglion. In these Z-cut ganglia, all rhythmic ascending intersegmental input is conveyed via one lateral connective while rhythmic descending input is conveyed via the contralateral connective. We found that rhythmic bursting recorded from the left and right sides of Z-cut ganglia had identical cycle periods with no phase difference, despite strong intersegmental inputs with differing periods from the two swimming ends of the preparations. We conclude that the swim circuits within individual leech ganglia act as single units. Moreover, we determined through correlation and Fourier spectral analyses, that the functional strengths of ascending and descending intersegmental inputs to Z-cut ganglia located in the middle of the nerve cord are approximately equal.