On the problem of lamination in the superficial dorsal horn of mammals: a reappraisal of the substantia gelatinosa in postnatal life

Although it is one of the most distinctive and earliest recognized features in the spinal cord, the substantia gelatinosa (SG) remains among the most enigmatic of central nervous system regions. The present neuroanatomical studies employed transganglionic transport of horseradish peroxidase conjugates of choleragenoid (B-HRP) and the B4 isolectin of Bandeiraea simplicifolia (IB4-HRP) on opposite sides to compare the projection patterns of myelinated and unmyelinated cutaneous primary afferents, respectively, within the superficial dorsal horn of the spinal cord in postnatal mice, from shortly after birth to adulthood. Putative unmyelinated afferents labeled with IB4-HRP gave rise to a dense sheet of terminal-like labeling restricted to the outer half of the SG. In contrast, myelinated inputs labeled with B-HRP gave rise to a similarly dense sheet of terminal-like labeling that occupied the inner half of the SG. This adult organization, with two dense sheets of terminal labeling in the superficial dorsal horn, was clearly evident shortly after birth using these markers, prior to the emergence of the SG. Furthermore, the location of the SG proper varied considerably within the dorsoventral plane of the dorsal horn according to mediolateral and segmental locations, a finding that was also seen in comparative studies in rat and cat. These findings caution against equating the SG in particular, and the superficial dorsal horn in general, with nociceptive processing; at minimum, the SG subserves a clear duality of function, with only a thin portion of its outermost aspect devoted to pain.