We examined the organization of somatosensory projections to the parabrachial (PB) and Kölliker-Fuse (KF) nuclei by employing the retrograde and anterograde axonal transport of Fluorogold and Phaseolus vulgaris-leucoagglutinin (PHA-L), respectively. Small PHA-L injections were made into different parts of the spinal trigeminal complex, including the paratrigeminal nucleus, and into different segments and laminae of the spinal dorsal horn. The subnuclear distribution of axonal labeling in the PB and KF was mapped with a camera lucida. Our results show that the somatosensory input to the PB and KF is highly organized. Neurons in the spinal trigeminal nuclei project predominantly to the KF and to the ventral portion of the external lateral PB. Neurons in the paratrigeminal nucleus project to the ventral lateral PB, the external medial PB, and to caudal aspects of the medial PB. These findings were supported by retrograde tracing experiments with Fluorogold. Spinal cord neurons located in the superficial dorsal horn (laminae I-II) of upper cervical segments project specifically to the ventral portion of the external lateral PB and, although more sparsely, to various other lateral PB nuclei. In contrast, neurons in the superficial dorsal horn of thoracic and lumbar spinal segments project mainly to the dorsal lateral and the central lateral PB. Finally, neurons in the lateral reticulated area and the lateral spinal nucleus of all spinal segments project almost exclusively to the internal lateral PB, whereas neurons in the respective nuclei of upper cervical segments also project to the KF. From our data we conclude that the somatosensory projections to the PB and KF are topographically organized. It is assumed that these pathways, which run from trigeminal and spinal neurons through the PB and KF to various forebrain, medullary, and spinal nuclei, form functionally different neural circuits that are involved in somatoautonomic processing.

The segmental and laminar organization of spinal projections to the functionally distinct ventrolateral (vlPAG) and lateral periaqueductal gray (lPAG) columns was examined by using retrograde and anterograde tracing techniques. It was found 1) that spinal input to both vlPAG and lPAG columns arose predominantly from neurons in the upper cervical (C1-4) and sacral spinal cord; 2) that there was a topographical separation of vl-PAG projecting and lPAG-projecting neurons within the upper cervical spinal cord; but 3) that below spinal segment C4, vlPAG-projecting and lPAG-projecting spinal neurons were similarly distributed, predominantly within contralateral lamina I, the nucleus of the dorsolateral fasciculus (the lateral spinal nucleus) and the lateral (reticular) part of lamina V. Consistent with the retrograde results, the greatest density of anterograde label, within both the vlPAG and lPAG, was found after tracer injections made either in the superficial or deep dorsal horn of the upper cervical spinal cord. Tracer injections made within the thoraco-lumbar spinal cord revealed that the vlPAG column received a convergent input from both the superficial and deep dorsal horn. However, thoraco-lumbar input to the lPAG was found to arise uniquely from the superficial dorsal horn; whereas the deep dorsal horn was found to innervate the "juxta-aqueductal" PAG region rather than projecting to the lPAG. These findings suggest that similar to spino-parabrachial projections, spinal projections to the lPAG (and juxta-aqueductal PAG) are topographically organised, with distinct subgroups of spinal neurons projecting to specific lPAG or juxta-aqueductal PAG subregions. In contrast, the vlPAG receives a convergent spinal input which arises from the superficial and deep dorsal horn of cervical, thoracic, lumbar, and sacral spinal segments.