A common observation in studies of neuronal structure is that axons differ in the size of their synaptic boutons. The significance of this size variation is unclear, in part because we do not know how the size of synaptic boutons is related to their internal organization. The present study has addressed this issue by using three-dimensional reconstruction of serial thin sections to examine the ultrastructure of synaptic boutons that vary in size. Our observations are based on complete or near-complete reconstructions of 53 synaptic boutons contacting large neurons in the ventromedial gray matter of the upper cervical spinal cord (probable neck motor neurons). We characterized bouton size in terms of volume and total area of membrane apposed to the motor neuron surface (apposition area). Boutons vary in apposition area by a factor of 40, and there is a significant positive correlation between our two measures of bouton size. In addition, bouton size is systematically related to four ultrastructural variables: 1) total active zone area, 2) number of active zones, 3) individual active zone area, and 4) number of synaptic vesicles. The correlations between these variables and both of our measures of bouton size are positive and significant. These data suggest that bouton size may be an index of ultrastructural features that are thought to influence transmitter storage and release.

In this and the accompanying paper (Yeow and Peterson, '86) we characterize motor nuclei of the cervical spinal cord in Pseudemys scripta and the motor pools of eight cervical muscles. We have identified three motor nuclei that supply the cervical musculature by using serial reconstructions of Nissl-stained spinal cords cut in three cardinal planes, and in experimental cases in which horseradish peroxidase (HRP) was applied to individual neck muscles. These nuclei are named according to their position as visualized in the transverse plane: dorsal, ventral, and medial. A fourth (ventrolateral) motor nucleus was never labelled following application of HRP to the cervical musculature and presumably innervates the forelimbs. The dorsal motor nucleus occupies the mid-dorsal to dorsolateral ventral horn of C1 and C2. It is composed of at least two morphological groups of motor neurons; one of these is a population of very large, fusiform profiles with transversely oriented dendrites that is found primarily in C1. The ventral motor nucleus occupies the tip of the ventral horn from C1 to C8. Its cells are significantly smaller and more numerous in rostral than in caudal cervical segments. In Nissl material, ventral nuclear profiles show little tendency to cluster into subgroups, but experimental cases suggest that there is some spatial dissociation of different motor pools within the ventral nucleus. The medial motor nucleus is described in the accompanying paper together with the motor pools of three cervical muscles that it supplies. Having identified the cervical motor nuclei we then used retrograde transport of HRP to characterize the motor pools of individual cervical muscles. Two superficial ventral muscles (mm. coracohyoideus and plastro-squamosus) are supplied by dorsal nuclear cells. M. coracohyoideus motor neurons are significantly larger than those of m. plastrosquamosus and the very large, fusiform cell type is relatively more numerous in the m. coracohyoideus motor pool. Dorsal and lateral muscles (mm. cervicocapitis, testocapitis, and transversalis cervicis) are supplied by ventral nuclear motor neurons. These cells are smaller, on average, than motor neurons supplying ventral musculature. The m. cervicocapitis motor pool lies dorsomedially in the tip of the ventral horn of C1 and C2; motor neurons supplying the more laterally placed mm. testocapitis and transversalis cervicis occur more ventrolaterally, in C2-C3 and C3-C5, respectively. Thus each of the five cervical muscles is supplied by a single motor nucleus, and their motor pools are organized into a musculotopic pattern.

In this paper we describe the medial motor nucleus of Pseudemys cervical spinal cord and the motor pools of three neck muscles that exhibit an unusual pattern of innervation. Cells of the medial motor nucleus form a longitudinal column at the dorsomedial gray/white border of the ventral horn from C1 through C8. In Nissl-stained transverse sections they appear fusiform with prominent medially projecting dendrites; in HRP material these dendrites are seen to cross into the contralateral ventral funiculus. Medial nuclear cells vary in size (12-31 micron in diameter) and are often relatively large (greater than 21 micron in diameter). They are significantly larger and more numerous in caudal than in rostral cervical segments. Medial nuclear cells supply three of the cervical muscles examined in this study: mm. retrahens capitis collique (RCCQ), testocervicis, and longus colli. These three muscles differ from other cervical muscles in Pseudemys and from vertebrate limb muscles in that they are supplied in parallel by two populations of motor neurons: the medial and ventral motor nuclei (cf. Yeow and Peterson, '86). Ventral nuclear cells supplying these three muscles are organized into a musculotopic pattern with m. testocervicis motor neurons most medial and m. RCCQ motor neurons lateral; in contrast, the location of medial nuclear motor neurons is invariant with respect to muscle position. HRP-positive medial nuclear cells are sometimes smaller (m. testocervicis) but more often are as large or larger (mm. RCCQ and longus colli) than ventral nuclear cells supplying the same muscles, thus suggesting that they supply extrafusal muscle fibers, perhaps different muscle unit types in the three muscles. Based on the morphology of medial nuclear cells and the probable actions of the muscles they innervate, we hypothesize that the medial motor nucleus may represent a discrete functional system for producing bilaterally synchronous muscle activation, and that this system is accessed by a subset of muscles in the cervical complex.