In the same general way as light the heat-stimulus acts on organisms, and causes the sensations, sometimes pleasant and sometimes unpleasant, which we call the subjective feeling of heat, warmth, coolness, or cold. The sense-organ that receives these impressions of temperature is the surface of the unicellular plasmic body in the protists, and the skin (epidermis) that protects the surface from the outer world in the histona. In all living things the temperature of the surrounding medium (water or air) has a great influence in regulating the life-processes; in the stationary animals and plants it is the temperature of the ground to which they are attached. This temperature must always be between the freezing-point and boiling-point of water, as fluid water is indispensable for the imbibition of the living matter and the molecular movements within the plasm. At the same time, some of the lower protists (chromacea, bacteria) can endure very high and very low temperatures, but only for a short time. Some protists (monera and diatomes) can stand a temperature of 200° C. for several days, and others can be heated above boiling-point without being killed. Arctic and High-Alpine plants and animals may be in a frozen condition for several months, yet live again when they are thawed. However, the resistance to these extremes of cold lasts for only a limited time, and in the frozen state all vital functions are at a standstill.
In the great majority of living things the vital activity is confined within narrow limits of temperature. Many plants and animals in the tropics which have been accustomed for thousands of years to the constancy of the hot equatorial climate can endure only very restricted variations of temperature. On the other hand, many of the inhabitants of Central Siberia, where the climate is very hot in the short summer and very cold in the long winter, can stand great variations. Thus the living plasm has experienced considerable changes in its sense of warmth through adaptation to different environments; not only the maximum and the minimum, but the optimum (most agreeable point), is subject to very great variations. This can easily be observed and followed experimentally in the phenomena of thermotaxis or thermotropism—that is to say, the effect that follows from a one-sided action of the heat-stimulus. The organism that falls below the minimum of temperature is said to be stiff with cold, while the organism that rises above the maximum is stiff with heat.
The heat-stimulus acts on inorganic as well as organic bodies, like the light-stimulus. The law holds good in both cases that higher temperatures increase sensation, while lower ones paralyze it. There is a minimum, an optimum, and a maximum, for many chemical and physical processes in the inorganic world. As far as the melting effect of water is concerned, freezing is the minimum of the heat stimulus and boiling the maximum. As the various chemical compounds meet in water at very different temperatures, we have an optimum for many substances—that is to say, a degree of warmth which is most favorable to the solution of a given quantity of a solid body in water. On the whole, the law holds for chemical processes that they are accelerated by high temperatures and retarded by low ones (like the human passions!); the former have a stimulating and the latter a benumbing effect. As the action of the various chemical compounds on each other is determined by the nature of the elements and their affinities, we must trace the variations in their conduct towards thermic stimuli to a sensation of temperature in the constituent atoms; increase of temperature stimulates it, while decrease lessens or paralyzes it. Here, again, the simple inorganic processes have a general resemblance to the complicated vital phenomena in the organic body.
Since we regard the whole of organic life as, in the ultimate analysis, merely a very elaborate chemical process, we shall quite expect that chemical stimuli are the most important factors in sensation. And this is so in point of fact; from the simplest moneron up to the most highly differentiated cell and on to the flower in the plant and the mental life of man, the vital processes are dominated by chemical forces and conversions of energy, which are set in play by external or internal chemical stimuli. The excitation which they produce is called, in a general way, "sensation of matter" or chemæsthesis; the basis of it is the mutual relation of the chemical elements which we describe as chemical affinity. In this affinity we have the play of attractive forces which lie in the nature of the elements themselves, especially in the peculiar properties of their constituent atoms; and this cannot be explained unless we ascribe unconscious sensation (in the widest sense) to the atoms, an inherent feeling of pleasure and the reverse, which they experience in the contact of other atoms (the "loves and hatreds of the elements" of Empedocles).
The numbers of different stimuli that act chemically on the plasm and excite its "sensation of matter" may be divided into two groups—external and internal stimuli. The latter lie within the organism itself, and cause the internal "organic sensations"; the former are in the outer world, and are felt as taste, smell, sex-impulse, etc. In the higher animals special chemical sense-organs have been developed for these chemical stimuli. As these are well known to us from our own human experience, and comparative physiology shows us the same structures in the higher animals, we will deal first with them. In general the same law holds for these external chemical stimuli as for optical and thermic stimuli; we can recognize a maximum limit of their action, a minimum below which they fail to stimulate, and an optimum or stage in which their influence is strongest.
The important part played in human life by taste and the pleasure associated with it is well known. The careful choice and preparation of savory food—which has become an art in gastronomy and a branch of practical philosophy in gastrosophy—was just as important two thousand years ago with the Greeks and Romans as it is to-day in royal banquets or the Lucullic dinners of millionaires. The excitement that we see associated with this refined combination of rich foods and drinks, and that finds expression in so many speeches and toasts, has its philosophic root in the harmony of gustatory sensations and the varying play of stimuli that the delicate dishes and wines exercise on the organs of taste, the tongue and palate. The microscopic organs of these parts of the mouth are the gustatory papillæ—cup-shaped structures, covered with spindle-shaped "taste-cells," and having a narrow opening into the cavity of the mouth. When sapid matters, drinks and fluid or loose particles of food, touch the taste-cells, they excite the fine terminal branchlets of the gustatory nerve which enters the cells. As we find that there are similar structures in most of the higher animals, and that they also choose their food with some care, we may confidently assume that they have sensations of taste like man. However, no trace of this is found in many of the lower animals; in these cases it is impossible to lay down a line of demarcation between taste and smell.
In man and the higher air-breathing vertebrates the seat of the sense of smell is in the nostrils; in man it is especially that part of the mucous lining of the nasal cavity which we call the "olfactory region" (the uppermost part of the nasal dividing wall, the superior and middle meatus). It is necessary for a sensation of smell that the odorous matter, or olfactory stimuli, be brought in a finely divided condition over the moist olfactory membranes. When they touch the olfactory cells—slender, rod-shaped cells with very fine hairs at the free end—they excite the ends of the olfactory nerve which are connected with the cells.
In many animals, especially mammals, the sense of smell has a much more important part in life than it has in man, in whom it is relatively feeble. It is well known that dogs and other carnivora, and even ungulates, have a much keener smell. In these cases the nasal cavity, which is the seat of the sense, is much larger, and the muscles in it are much stronger. The nostrils of the air-breathing vertebrates have been developed from a pair of open nasal depressions in the skin of the fish's head. But in these aquatic vertebrates the chemical action of the olfactory stimuli must be of a different character, like the sensation of taste. The odorous matter is, in these cases, brought into contact with the olfactory membrane in a liquid form (in which condition it is not perceptible to man). In fact, the division between the senses of smell and taste disappears altogether in the lower animals. These two "chemical senses" are closely related, and have a common feature in the direct chemical action of the stimulus on the sensitive part of the skin.
A chemical sensation of matter that corresponds completely to the real taste-sensation in the higher animals is found in some of the higher carnivorous plants. The leaves of the sun-dew (drosera rotundifolia) are very sensitive insect-traps, and are armed at the edge with knob-like tentacles, sticky hairs that secrete an acid, flesh-digesting juice. When a solid body (but not a raindrop) touches the surface of the leaf the stimulus acts in such a way on the tentacle heads as to contract the leaf. But the acid fluid which serves for digestion, and corresponds to the gastric juice in the animal, is only secreted by the corpuscles if the solid foreign body is nitrogenous (flesh or cheese). Hence the leaves of these insectivorous plants taste their meat diet, and distinguish it from other solids, to which they are indifferent. In the broader sense, in fact, we may describe the points of the roots of plants as organs of taste; they plunge into the richer parts of the earth which yield more nourishment, and avoid the poor parts. In unicellular plants and animals the action of chemical stimuli is especially conspicuous when it is one-sided, and provokes definite movements in one particular direction (chemotaxis).