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Neurohormones and Neurohumors pp 140–184Cite as

The Mammalian Pineal Organ

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Summary

This survey is based on selected data from the large literature as well as on own observations and discusses aspects of pineal phylogenetic and ontogenetic development, the fine structure of the mammalian pineal, its biochemistry and physiology. The author has been trying to integrate data, to stress the structural, biochemical and functional specificity of the organ, and to point to problems which should be further investigated. He has not been shy of one or two, perhaps fruitful, speculations.

The phylogenetic development of the organ supports the opinion that the mammalian pinealocyte is a non-nervous element derived from the neurosensory photoreceptor cell present in the submammalian pineal organ. In Sauropsida the photosensory function of the epiphysis is gradually lost while its secretory function develops. This functional change, already foreshadowed in anuran amphibians, is accompanied by a change in pineal innervation. The sensory pineal tract disappears and the organ becomes innervated by peripheral autonomic fibers. Primarily, these fibers reach the pineal exclusively along the pineal vessels, running in the perivascular spaces. Secondarily, an additional contribution of sympathetic fibers is realized by way of the nervi eonarii. The latter fibers penetrate directly into the pineal parenchyma their endings making simple appositional as well as specific synaptic contacts with the pinealocytes. In this paper, such synaptic contacts are demonstrated in the rat pineal.

By way of a working hypothesis the mode of innervation of the pineal organ of mammals is compared with that of smooth musculature. Stimulation of pinealocytes is thought to occur by (1) diffusion of neurotransmitter released from the varicosities and endings of sympathetic fibers present in the perivascular spaces, (2) simple appositional contact of sympathetic nerve terminals with pinealocytes in the parenchyma, (3) specialized synaptic contact between sympathetic nerve terminals and pinealocytes in the parenchyma, and, speculatively, (4) by transmission of impulses from one pinealocyte to another which can either be (a) of a neurohumoral or (b) of an electrotonic nature. Probably, (1) is the phylogenetic earliest mode of stimulation.

The topographical relationship between the terminal buds of the processes of pinealocytes and the perivascular space varies in mammals. Although the secretory function of the pinealocyte can scarcely be doubted, the problem of the production, storage and extrusion of secretory products is not quite solved at the ultra-structural level. The compound(s) produced by the pinealocytes most probably reach the blood via the perivascular spaces, a condition which is compared with the release of compounds produced by neurons and neurosecretory cells into the blood via pericapillary spaces in the region of the median eminence and in the neural part of the hypophysis.

In the chapter on pineal biochemistry the presence of catecholamines, indole-amines, lipids, proteins and some enzymes is discussed as is the influence of pineal denervation and of light and darkness on the pineal content of some of them. The neural pathway for the transmission of photic stimuli from the retina of the lateral eyes to the epiphysis is also mentioned. In the pineal some compounds are synthetized which are exclusively produced in this organ.

In the chapter on pineal physiology reference is made to experimental research pointing to the effects of light and darkness, of pinealectomy and of administration of pineal extracts on the size, weight and function of the reproductive organs. It appears that the pineal exerts an inhibiting influence on these organs. A tentative explanation is given of the functional significance of the intact pineal innervation by sympathetic fibers. It is assumed that light and darkness, via this innervation, condition the basically internal regulated function of the organ. External stimuli, such as photic, act in a regulation mechanism which is superimposed on an intrinsic pineal regulation mechanism which is probably of a hormonal nature. Experimentally produced changes in pineal function are, after some time, no longer of any effect on the normal function of the reproductive system. It would seem that the pineal organ is not indispensable for the normal function of this system as appears, on the long run, after pinealectomy.

Regarding the nature of the pineal compound(s) inhibiting the reproductive system different opinions exist. They would be either indole derivatives such as melatonin and 5-methoxytryptophol, or peptides. The finding that by one peptide fraction of a pineal extract hypophyseal FSH secretion in vitro is increased whereas by another peptide fraction this secretion is decreased has now proven beyond doubt that (1) the pineal produces an antigonadotropic as well as a gonadotropic principle, (2) these principles act on the gonadotropic cells of the distal part of the hypophysis rather than at the level of the organs of reproduction. Other arguments are also in favor of the opinion that the direct target organ of the pineal compound(s) is the hypophysis. That this pinealo-hypophyseal action may be additionally regulated by hypothalamic centers is suggested by changes in activity of these centers after administration of pineal extracts. It also appears that, most probably, the distal part of the hypophysis rather than the gonads regulates the production of the pineal antigonadotropic principle.

In conclusion it can be stated that the mammalian pineal is a photo-neuroendocrine organ (E. Scharrer) on the understanding that the structure is a neuro-epithelial derivative, that the endocrine function is exerted by non-nervous but specific pineal cells, the pinealocytes, which are phylogenetically derived from neurosensory photoreceptor cells, and that the mammalian pineal is an indirect photosensory organ, photic stimuli reaching it via a neural pathway starting in the retina and ending in the organ by way of its sympathetic innervation. The mammalian pineal functions as a “regulator of regulators” and is not absolutely indispensable for the normal function of the organism.

The facts now known about this intriguing organ should stimulate further research to solve the many problems so far still unsolved.

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    Prof. Dr. J. Ariëns Kappers

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Kappers, J.A. (1969). The Mammalian Pineal Organ. In: Kappers, J.A. (eds) Neurohormones and Neurohumors. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-662-25519-3_7

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