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Professor Eimer. He began, like myself, by what in the next chapter I call the "fundamental observation on the effects of excising the nervecentres, and from this basis he worked both at the physiology and the morphology of the neuromuscular tissues. In point of time, I was the first to make the fundamental observation, and he was the first to publish it. The sundry features in which our subsequent investigations agreed, and those in which they differed, I shall mention throughout the course of the following pages.
I shall now conclude this chapter by giving a brief account of those general principles of the physiology of nerve and muscle with which it is necessary to be fully acquainted, in order to understand the course of the following experiments.
Nerve-tissue, then, universally consists of two elementary structures, viz. very minute nerve-cells and very minute nerve-fibres. The fibres proceed to and from the cells, so in some cases serving to unite the cells with one another, and in other cases with distant parts of the animal body. Nerve-cells are usually found collected together in aggregates, which are called nerve-centres or ganglia, to and from which large bundles of nerve-fibres come and go.
To explain the function of nerve-tissue, it is necessary to begin by explaining what physiologists mean by the term "excitability." Suppose that a
* "Die Medusen physiologisch und morphologisch auf ihr Nervensystem untersucht" (Tübingen, 1878).
muscle has been cut from the body of a freshly killed animal; so long as it is not interfered with in any way, so long will it remain quite passive. But every time a stimulus is supplied to it, either by means of a pinch, a burn, an electrical shock, or a chemical irritant, the muscle will give a single contraction in response to every stimulation. And it is this readiness of organic tissues to respond to a suitable stimulus that physiologists designate by the term "excitability."
Nerves, no less than muscles, present the property of being excitable. If, together with the excised muscle, there had been removed from the animal's body an attached nerve, every time any part of this nerve is stimulated the attached muscle. will contract as before. But it must be carefully observed that there is this great difference between these two cases of response on the part of the muscle that while in the former case the muscle responded to a stimulus applied directly to its own substance, in the latter case the muscle responded to a stimulus applied at a distance from its own substance, which stimulus was then conducted to the muscle by the nerve. And in this we perceive the characteristic function of nerve-fibres, viz. that of conducting stimuli to a distance. The function of nerve-cells is different, viz. that of accumulating nervous energy, and, at fitting times, of discharging this energy into the attached nerve-fibres. The nervous energy, when thus discharged, acts as a stimulus to the nerve-fibre; so that if a muscle is attached to the end of a fibre, it contracts on receiv
ing this stimulus. I may add that when nerve-cells are collected into ganglia, they often appear to discharge their energy spontaneously; so that in all but the very lowest animals, whenever we see apparently spontaneous action, we infer that ganglia are probably present. Lastly, another important distinction must be borne in mind-the distinction, namely, which is to be drawn between muscle and nerve. A stimulus applied to a nerveless muscle can only course through the muscle by giving rise to a visible wave of contraction, which spreads in all directions from the seat of disturbance as from a centre. A nerve, on the other hand, conducts the stimulus without sensibly moving or undergoing any change of shape. Now, in order not to forget this distinction, I shall always speak of musclefibres as conveying a visible wave of contraction, and of nerve-fibres as conveying an invisible, or molecular, wave of stimulation. Nerve-fibres, then, are functionally distinguished from muscle-fibresand also from protoplasm-by displaying the property of conducting invisible, or molecular, waves of stimulation from one part of an organism to another, so establishing physiological continuity between such parts, without the necessary passage of waves of contraction.
THE naked-eyed Medusæ are very much smaller in size than the covered-eyed, and as we shall find that the distribution of their nervous elements is somewhat different, it will be convenient to use different names for the large umbrella-shaped part of a covered-eyed Medusa, and the much smaller though corresponding part of a naked-eyed Medusa. The former, therefore, I shall call the umbrella, and the latter the swimming-bell, or nectocalyx. In each case alike this portion of the animal performs the office of locomotion, and it does so in the same way. I have already said that this mushroom-like organ, which constitutes the main bulk of the animal, is itself mainly constituted of thick transparent and non-contractile jelly, but that the whole of its concave surface is lined with a thin sheet of muscular tissue. Such being the structure of the organ, the mechanism whereby it effects locomotion is very simple, consisting merely of an alternate contraction and relaxation of the entire muscular sheet which lines the cavity of the bell. At each contraction of this muscular sheet the gelatinous
walls of the bell are drawn together; the capacity of the bell being thus diminished, water is ejected from the open mouth of the bell backwards, and the consequent reaction propels the animal forwards. In these swimming movements, systole and diastole follow one another with as perfect a rhythm as they do in the beating of a heart.
Effects of excising the entire Margins of Nectocalyces.
Confining our attention under this heading to the naked-eyed Medusæ, I find that the following proposition applies to every species of the group which I have as yet had the opportunity of examining: Excision of the extreme margin of a nectocalyx causes immediate, total, and permanent paralysis of the entire organ. Nothing can possibly be more definite than in this highly remarkable effect. I have made hundreds of observations upon various species of the naked-eyed Medusæ, of all ages and conditions of freshness, vigour, etc.; and I have constantly found that if the experiment be made with ordinary care, so as to avoid certain sources of error presently to be named, the result is as striking and decided as it is possible to desire.* Indeed, I do not know of any case in the animal kingdom where the removal of a centre of spontaneity causes so sudden and so complete a paralysis
* I have only met with one individual exception. This occurred in a specimen of Staurophora laciniata, where, after removal of the entire margin, three centres of spontaneity were found to remain in the sheet of contractile tissue lining the nectocalyx.