If bladder physiology did little, originally, to help theorizing the sacral parasympathetic outflow, blood vessels were the crux of the matter. In his 1899 Presidential Address to the Physiological Society [
2], credited by Langley himself [
21] as his first published statement on “the sacral and cranial pathways as one system”, he puts forward “one reason” to support this view: “if regions above and below [the middle portion of the spinal cord] were mere separated parts of the sympathetic region we should expect that, when some of these regions and the sympathetic region send nerves to the same spots, the effects produced by both sets of nerves would be the same in kind though it might differ in extent. But it is often not the case. Thus, certain blood vessels may receive nerve fibers from four spinal nerves in the sympathetic region and three spinal nerves in the sacral region, all the former cause contraction of the blood vessels, all the latter cause dilatation”. Prominent among the “certain blood vessels” are those to the external genitals, analyzed by Langley and Anderson [
27], that cause erection by vasodilation and detumescence by vasoconstriction. On this issue, however, Langley was and remained somewhat of an outsider. Twenty years earlier, Eckhard had found [
13] that “in the rabbit, the mechanism of erection is provided by nerves via a second pathway, that is by the nerve trunk that anatomically corresponds to the superior hypogastric plexus in man [i.e. the hypogastric nerve]”.
1 In 1895, the same year as Langley and Anderson’s paper, François-Franck announced that he “obtained a penile vasodilation from the descending branches of the inferior mesenteric ganglion with the same clarity as with the common sacral erector nerve”
2 [
46]. He also found an occasional vasoconstrictor effect of the anterior pelvic nerve and concluded that “with the exception of the posterior erector nerve of Eckhard one finds associated in all nerves vasoconstrictor and vasodilator fibers”, “whose combined effect depended on the “nature form, intensity and rhythm of the excitations”. Half a century later, a similar explanation (“a masking effect produced by concurrent stimulation of vasoconstrictor fibers”) was offered by Root and Bard [
47] for the failure of Langley to see “the suprasacral vasodilator pathway to the penis [that] runs through sympathetic channels”, that they deduced from a series of 154 tests on 21 cats with precise spinal lesions. Bessou and Laporte [
48] concluded their own study on cat by a similar observation: “The hypogastric nerve, belonging to the orthosympathetic system, contain, in the cat, cholinergic vasodilatating fibers whose prolonged and repetitive stimulation can provoke an erection, despite the presence of fibers with an opposing action”.
3 Human patients with complete destruction of the lower lumbar or sacral cord, but not the upper lumbar or thoracic one, experience psychogenic erections mediated by the lumbar pathway [
49,
50]. In 1979, Sjöstrand and Klinge found that the pelvic and hypogastric outflows are synergistically vasodilatory in rabbit [
51] and commented, with a hint of annoyance: “We feel that now, more than a hundred years after their original description (referring to [
13]), it is time to generally accept the existence of the sympathetic hypogastric erectile fibers.” However, 40 years later, the time for “general acceptance” has yet to come and, here again, we incriminate the conceptual sway of the sympatho-parasympathetic antagonism on the field: contemporary reviews span the range from more or less explicit acknowledgments that thoracolumbar sympathetic neurons are
“involved in erection” {[
45] (p 357), [
52] (p 29)}, to elaborate dismissals of this pathway as a plasticity phenomenon induced by lesions of the spinal cord [
50] or protests that “under normal conditions, stimulation of pelvic splanchnic nerves (PSN; nervi erigentes, parasympathetic) and of the hypogastric nerve (sympathetic) have different effects on erectile tissues” [
53]. Somehow, the lumbosacral synergy is less controversial concerning the emission phase of ejaculation (“nicely integrated” [
50] since “both sympathetic and parasympathetic tones act in a synergistic fashion to initiate seminal emission” [
54] and [
45] (p. 357) and references therein)—and the ejaculation center is lumbar [
55,
56], so that the command of the male sexual act has come to be understood as shifting seamlessly, and oddly, from start to finish, from “parasympathetic” to sympathetic neurons [
50].