Spinal distribution of ascending lamina I axons anterogradely labeled with Phaseolus vulgaris leucoagglutinin (PHA‐L) in the cat

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Abstract
The location of the ascending axons of spinal lamina I cells was studied in cats that received injections of Phaseolus vulgaris leucoagglutinin (PHA-L) in the superficial dorsal horn of the cervical or lumbosacral enlargement. Lamina I axons that could be ascribed to the spinothalamic tract (STT) were of particular interest. The cases were divided into three sets: in seven optimal cases the injections were restricted to lamina I; in ten nominal cases the injections involved laminae I–II or laminae I–III and occasionally lamina IV; and in eight mixed cases laminae I–V were injected. Since ipsilateral propriospinal and bilateral supraspinal axons originate from laminae I and V, but only ipsilateral propriospinal axons from laminae II–IV, this categorization facilitated a comparative analysis. Ascending axons labeled immunohistochemically with avidin/Texas Red were observed in oblique transverse sections from the C1, C3/4, T6, T12, and L3/4 levels. Incidental axonal labeling occurred in the ipsilateral dorsal columns because of passing primary afferent fiber uptake and, in nominal and mixed cases with involvement of laminae III–IV, in the superficial dorsolateral funiculus at the location of the spinocervical tract. Ipsilateral ascending lamina I axons in optimal cases were located in Lissauer's tract and in the white matter adjacent to the dorsal horn. Since these appeared to terminate in lamina I, and few remained at C1, they were ascribed to propriospinal projections. Contralateral ascending lamina I axons in optimal and nominal cases were distributed throughout the dorsal and ventral portions of the lateral funiculus (LF), but, despite considerable variability between animals in their location and dispersion, they were consistently concentrated in the middle of the LF (i.e., at the level of the central canal). This concentration was observed in a slightly more ventral location at C1, and a similar but weaker concentration of lamina I axons was located slightly more dorsally in C1 on the ipsilateral side. These supraspinal lamina I projections were ascribed to the spinomesencephalic tract (SMT) and-to the STT. In mixed cases, additional ascending axons ascribed to lamina V cells were labeled in the ventrolateral and ventral funiculi. Many labeled axons were found in this region following a large injection of biocytin into lumbosacral laminae V–VIII in a supplementary case. These results thus together support previous descriptions of a dorsoventral distribution of STT axons according to laminar origin, but they contradict recent reports that lamina I axons ascend in the dorsolateral funiculus. These observations indicate that lamina I STT (and SMT) axons are concentrated in the middle of the contralateral LF of the cat, that they are nonetheless present throughout the LF, that there is a weaker ipsilateral LF projection, and that there is individual variability. These attributes are all consistent with clinical and behavioral findings regarding the spinal location of ascending fibers critical for pain and temperature sensation. These results thus directly support the concept that the lamina I STT projection is an integral component of the central representation of specific pain and temperature sensibility.

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