THE INCLINED PLANE OF ANIMAL BEHAVIOUR
Before passing to a connected story of the gradual emergence of higher and higher forms of life in the course of the successive ages—the procession of life, as it may be called—it will be useful to consider the evolution of animal behaviour.
Evolution of Mind
A human being begins as a microscopic fertilised egg-cell, within which there is condensed the long result of time—Man's inheritance. The long period of nine months before birth, with its intimate partnership between mother and offspring, is passed as it were in sleep, and no one can make any statement in regard to the mind of the unborn child. Even after birth the dawn of mind is as slow as it is wonderful. To begin with, there is in the ovum and early embryo no nervous system at all, and it develops very gradually from simple beginnings. Yet as mentality cannot come in from outside, we seem bound to conclude that the potentiality of it—whatever that means—resides in the individual from the very first. The particular kind of activity known to us as thinking, feeling, and willing is the most intimate part of our experience, known to us directly apart from our senses, and the possibility of that must be implicit in the germ-cell just as the genius of Newton was implicit in a very miserable specimen of an infant. Now what is true of the individual is true also of the race—there is a gradual evolution of that aspect of the living creature's activity which we call mind. We cannot put our finger on any point and say: Before this stage there was no mind. Indeed, many facts suggest the conclusion that wherever there is life there is some degree of mind—even in the plants. Or it might be more accurate to put the conclusion in another way, that the activity we call life has always in some degree an inner or mental aspect.
In another part of this book there is an account of the dawn of mind in backboned animals; what we aim at here is an outline of what may be called the inclined plane of animal behaviour.
A very simple animal accumulates a little store of potential energy, and it proceeds to expend this, like an explosive, by acting on its environment. It does so in a very characteristic self-preservative fashion, so that it burns without being consumed and explodes without being blown to bits. It is characteristic of the organism that it remains a going concern for a longer or shorter period—its length of life. Living creatures that expended their energy ineffectively or self-destructively would be eliminated in the struggle for existence. When a simple one-celled organism explores a corner of the field seen under a microscope, behaving to all appearance very like a dog scouring a field seen through a telescope, it seems permissible to think of something corresponding to mental endeavour associated with its activity. This impression is strengthened when an amœba pursues another amœba, overtakes it, engulfs it, loses it, pursues it again, recaptures it, and so on. What is quite certain is that the behaviour of the animalcule is not like that of a potassium pill fizzing about in a basin of water, nor like the lurching movements of a gun that has got loose and "taken charge" on board ship. Another feature is that the locomotor activity of an animalcule often shows a distinct individuality: it may swim, for instance, in a loose spiral.
But there is another side to vital activity besides acting upon the surrounding world; the living creature is acted on by influences from without. The organism acts on its environment; that is the one side of the shield: the environment acts upon the organism; that is the other side. If we are to see life whole we must recognise these two sides of what we call living, and it is missing an important part of the history of animal life if we fail to see that evolution implies becoming more advantageously sensitive to the environment, making more of its influences, shutting out profitless stimuli, and opening more gateways to knowledge. The bird's world is a larger and finer world than an earthworm's; the world means more to the bird than to the worm.
The Trial and Error Method
Simple creatures act with a certain degree of spontaneity on their environment, and they likewise react effectively to surrounding stimuli. Animals come to have definite "answers back," sometimes several, sometimes only one, as in the case of the Slipper Animalcule, which reverses its cilia when it comes within the sphere of some disturbing influence, retreats, and, turning upon itself tentatively, sets off again in the same general direction as before, but at an angle to the previous line. If it misses the disturbing influence, well and good; if it strikes it again, the tactics are repeated until a satisfactory way out is discovered or the stimulation proves fatal.
It may be said that the Slipper Animalcule has but one answer to every question, but there are many Protozoa which have several enregistered reactions. When there are alternative reactions which are tried one after another, the animal is pursuing what is called the trial-and-error method, and a higher note is struck.
There is an endeavour after satisfaction, and a trial of answers. When the creature profits by experience to the extent of giving the right answer first, there is the beginning of learning.
Among simple multicellular animals, such as sea-anemones, we find the beginnings of reflex actions, and a considerable part of the behaviour of the lower animals is reflex. That is to say, there are laid down in the animal in the course of its development certain pre-arrangements of nerve-cells and muscle-cells which secure that a fit and proper answer is given to a frequently recurrent stimulus. An earthworm half out of its burrow becomes aware of the light tread of a thrush's foot, and jerks itself back into its hole before anyone can say "reflex action." What is it that happens?
Certain sensory nerve-cells in the earthworm's skin are stimulated by vibrations in the earth; the message travels down a sensory nerve-fibre from each of the stimulated cells and enters the nerve-cord. The sensory fibres come into vital connection with branches of intermediary, associative, or communicating cells, which are likewise connected with motor nerve-cells. To these the message is thus shunted. From the motor nerve-cells an impulse or command travels by motor nerve-fibres, one from each cell, to the muscles, which contract. If this took as long to happen as it takes to describe, even in outline, it would not be of much use to the earthworm. But the motor answer follows the sensory stimulus almost instantaneously. The great advantage of establishing or enregistering these reflex chains is that the answers are practically ready-made or inborn, not requiring to be learned. It is not necessary that the brain should be stimulated if there is a brain; nor does the animal will to act, though in certain cases it may by means of higher controlling nerve-centres keep the natural reflex response from being given, as happens, for instance, when we control a cough or a sneeze on some solemn occasion. The evolutionary method, if we may use the expression, has been to enregister ready-made responses; and as we ascend the animal kingdom, we find reflex actions becoming complicated and often linked together, so that the occurrence of one pulls the trigger of another, and so on in a chain. The behaviour of the insectivorous plant called Venus's fly-trap when it shuts on an insect is like a reflex action in an animal, but plants have no definite nervous system.
What are Called Tropisms
A somewhat higher level on the inclined plane is illustrated by what are called "tropisms," obligatory movements which the animal makes, adjusting its whole body so that physiological equilibrium results in relation to gravity, pressure, currents, moisture, heat, light, electricity, and surfaces of contact. A moth is flying past a candle; the eye next the light is more illumined than the other; a physiological inequilibrium results, affecting nerve-cells and muscle-cells; the outcome is that the moth automatically adjusts its flight so that both eyes become equally illumined; in doing this it often flies into the candle.
It may seem bad business that the moth should fly into the candle, but the flame is an utterly artificial item in its environment to which no one can expect it to be adapted. These tropisms play an important rôle in animal behaviour.
On a higher level is instinctive behaviour, which reaches such remarkable perfection in ants, bees, and wasps. In its typical expression instinctive behaviour depends on inborn capacities; it does not require to be learned; it is independent of practice or experience, though it may be improved by both; it is shared equally by all members of the species of the same sex (for the female's instincts are often different from the male's); it refers to particular conditions of life that are of vital importance, though they may occur only once in a lifetime. The female Yucca Moth emerges from the cocoon when the Yucca flower puts forth its bell-like blossoms. She flies to a flower, collects some pollen from the stamens, kneads it into a pill-like ball, and stows this away under her chin. She flies to an older Yucca flower and lays her eggs in some of the ovules within the seed-box, but before she does so she has to deposit on the stigma the ball of pollen. From this the pollen-tubes grow down and the pollen-nucleus of a tube fertilises the egg-cell in an ovule, so that the possible seeds become real seeds, for it is only a fraction of them that the Yucca Moth has destroyed by using them as cradles for her eggs. Now it is plain that the Yucca Moth has no individual experience of Yucca flowers, yet she secures the continuance of her race by a concatenation of actions which form part of her instinctive repertory.
From a physiological point of view instinctive behaviour is like a chain of compound reflex actions, but in some cases, at least, there is reason to believe that the behaviour is suffused with awareness and backed by endeavour. This is suggested in exceptional cases where the stereotyped routine is departed from to meet exceptional conditions. It should also be noted that just as ants, hive bees, and wasps exhibit in most cases purely instinctive behaviour, but move on occasion on the main line of trial and error or of experimental initiative, so among birds and mammals the intelligent behaviour is sometimes replaced by instinctive routine. Perhaps there is no instinctive behaviour without a spice of intelligence, and no intelligent behaviour without an instinctive element. The old view that instinctive behaviour was originally intelligent, and that instinct is "lapsed intelligence," is a tempting one, and is suggested by the way in which habitual intelligent actions cease in the individual to require intelligent control, but it rests on the unproved hypothesis that the acquisitions of the individual can be entailed on the race. It is almost certain that instinct is on a line of evolution quite different from intelligence, and that it is nearer to the inborn inspirations of the calculating boy or the musical genius than to the plodding methods of intelligent learning.
The higher reaches of the inclined plane of behaviour show intelligence in the strict sense. They include those kinds of behaviour which cannot be described without the suggestion that the animal makes some sort of perceptual inference, not only profiting by experience but learning by ideas. Such intelligent actions show great individual variability; they are plastic and adjustable in a manner rarely hinted at in connection with instincts where routine cannot be departed from without the creature being nonplussed; they are not bound up with particular circumstances as instinctive actions are, but imply an appreciative awareness of relations.
When there is an experimenting with general ideas, when there is conceptual as contrasted with perceptual inference, we speak of Reason, but there is no evidence of this below the level of man. It is not, indeed, always that we can credit man with rational conduct, but he has the possibility of it ever within his reach.
Animal instinct and intelligence will be illustrated in another part of this work. We are here concerned simply with the general question of the evolution of behaviour. There is a main line of tentative experimental behaviour both below and above the level of intelligence, and it has been part of the tactics of evolution to bring about the hereditary enregistration of capacities of effective response, the advantages being that the answers come more rapidly and that the creature is left free, if it chooses, for higher adventures.
There is no doubt as to the big fact that in the course of evolution animals have shown an increasing complexity and masterfulness of behaviour, that they have become at once more controlled and more definitely free agents, and that the inner aspect of the behaviour—experimenting, learning, thinking, feeling, and willing—has come to count for more and more.
Evolution of Parental Care
Mammals furnish a crowning instance of a trend of evolution which expresses itself at many levels—the tendency to bring forth the young at a well-advanced stage and to an increase of parental care associated with a decrease in the number of offspring. There is a British starfish called Luidia which has two hundred millions of eggs in a year, and there are said to be several millions of eggs in conger-eels and some other fishes. These illustrate the spawning method of solving the problem of survival. Some animals are naturally prolific, and the number of eggs which they sow broadcast in the waters allows for enormous infantile mortality and obviates any necessity for parental care.
But some other creatures, by nature less prolific, have found an entirely different solution of the problem. They practise parental care and they secure survival with greatly economised reproduction. This is a trend of evolution particularly characteristic of the higher animals. So much so that Herbert Spencer formulated the generalisation that the size and frequency of the animal family is inverse ratio to the degree of evolution to which the animal has attained.
Now there are many different methods of parental care which secure the safety of the young, and one of these is called viviparity. The young ones are not liberated from the parent until they are relatively well advanced and more or less able to look after themselves. This gives the young a good send-off in life, and their chances of death are greatly reduced. In other words, the animals that have varied in the direction of economised reproduction may keep their foothold in the struggle for existence if they have varied at the same time in the direction of parental care. In other cases it may have worked the other way round.
In the interesting archaic animal called Peripatus, which has to face a modern world too severe for it, one of the methods of meeting the environing difficulties is the retention of the offspring for many months within the mother, so that it is born a fully-formed creature. There are only a few offspring at a time, and, although there are exceptional cases like the summer green-flies, which are very prolific though viviparous, the general rule is that viviparity is associated with a very small family. The case of flowering plants stands by itself, for although they illustrate a kind of viviparity, the seed being embryos, an individual plant may have a large number of flowers and therefore a huge family.
Viviparity naturally finds its best illustrations among terrestrial animals, where the risks to the young life are many, and it finds its climax among mammals.
Now it is an interesting fact that the three lowest mammals, the Duckmole and two Spiny Ant-eaters, lay eggs, i.e. are oviparous; that the Marsupials, on the next grade, bring forth their young, as it were, prematurely, and in most cases stow them away in an external pouch; while all the others—the Placentals—show a more prolonged ante-natal life and an intimate partnership between the mother and the unborn young.
There is another way of looking at the sublime process of evolution. It has implied a mastery of all the possible haunts of life; it has been a progressive conquest of the environment.
1. It is highly probable that living organisms found their foothold in the stimulating conditions of the shore of the sea—the shallow water, brightly illumined, seaweed-growing shelf fringing the Continents. This littoral zone was a propitious environment where sea and fresh water, earth and air all meet, where there is stimulating change, abundant oxygenation and a copious supply of nutritive material in what the streams bring down and in the rich seaweed vegetation.
It is not an easy haunt of life, but none the worse for that, and it is tenanted to-day by representatives of practically every class of animals from infusorians to seashore birds and mammals.
The Cradle of the Open Sea
2. The open-sea or pelagic haunt includes all the brightly illumined surface waters beyond the shallow water of the shore area.
It is perhaps the easiest of all the haunts of life, for there is no crowding, there is considerable uniformity, and an abundance of food for animals is afforded by the inexhaustible floating "sea-meadows" of microscopic Algæ. These are reincarnated in minute animals like the open-sea crustaceans, which again are utilised by fishes, these in turn making life possible for higher forms like carnivorous turtles and toothed whales. It is quite possible that the open sea was the original cradle of life and perhaps Professor Church is right in picturing a long period of pelagic life before there was any sufficiently shallow water to allow the floating plants to anchor. It is rather in favour of this view that many shore animals such as crabs and starfishes, spend their youthful stages in the relatively safe cradle of the open sea, and only return to the more strenuous conditions of their birthplace after they have gained considerable strength of body. It is probably safe to say that the honour of being the original cradle of life lies between the shore of the sea and the open sea.
The Great Deeps
3. A third haunt of life is the floor of the Deep Sea, the abyssal area, which occupies more than a half of the surface of the globe. It is a region of extreme cold—an eternal winter; of utter darkness—an eternal night—relieved only by the fitful gleams of "phosphorescent" animals; of enormous pressure—2½ tons on the square inch at a depth of 2,500 fathoms; of profound calm, unbroken silence, immense monotony. And as there are no plants in the great abysses, the animals must live on one another, and, in the long run, on the rain of moribund animalcules which sink from the surface through the miles of water. It seems a very unpromising haunt of life, but it is abundantly tenanted, and it gives us a glimpse of the insurgent nature of the living creature that the difficulties of the Deep Sea should have been so effectively conquered. It is probable that the colonising of the great abysses took place in relatively recent times, for the fauna does not include many very antique types. It is practically certain that the colonisation was due to littoral animals which followed the food-débris, millennium after millennium, further and further down the long slope from the shore.
4. A fourth haunt of life is that of the freshwaters, including river and lake, pond and pool, swamp and marsh. It may have been colonised by gradual migration up estuaries and rivers, or by more direct passage from the seashore into the brackish swamp. Or it may have been in some cases that partially landlocked corners of ancient seas became gradually turned into freshwater basins. The animal population of the freshwaters is very representative, and is diversely adapted to meet the characteristic contingencies—the risk of being dried up, the risk of being frozen hard in winter, and the risk of being left high and dry after floods or of being swept down to the sea.
Conquest of the Dry Land
5. The terrestrial haunt has been invaded age after age by contingents from the sea or from the freshwaters. We must recognise the worm invasion, which led eventually to the making of the fertile soil, the invasion due to air-breathing Arthropods, which led eventually to the important linkage between flowers and their insect visitors, and the invasion due to air-breathing Amphibians, which led eventually to the higher terrestrial animals and to the development of intelligence and family affection. Besides these three great invasions, there were minor ones such as that leading to land-snails, for there has been a widespread and persistent tendency among aquatic animals to try to possess the dry land.
Getting on to dry land had a manifold significance.
It implied getting into a medium with a much larger supply of oxygen than there is dissolved in the water. But the oxygen of the air is more difficult to capture, especially when the skin becomes hard or well protected, as it is almost bound to become in animals living on dry ground. Thus this leads to the development of internal surfaces, such as those of lungs, where the oxygen taken into the body may be absorbed by the blood. In most animals the blood goes to the surface of oxygen-capture; but in insects and their relatives there is a different idea—of taking the air to the blood or in greater part to the area of oxygen-combustion, the living tissues. A system of branching air-tubes takes air into every hole and corner of the insect's body, and this thorough aeration is doubtless in part the secret of the insect's intense activity. The blood never becomes impure.
The conquest of the dry land also implied a predominance of that kind of locomotion which may be compared to punting, when the body is pushed along by pressing a lever against a hard substratum. And it also followed that with few exceptions the body of the terrestrial animal tended to be compact, readily lifted off the ground by the limbs or adjusted in some other way so that there may not be too large a surface trailing on the ground. An animal like a jellyfish, easily supported in the water, would be impossible on land. Such apparent exceptions as earthworms, centipedes, and snakes are not difficult to explain, for the earthworm is a burrower which eats its way through the soil, the centipede's long body is supported by numerous hard legs, and the snake pushes itself along by means of the large ventral scales to which the lower ends of very numerous ribs are attached.
Methods of Mastering the Difficulties of Terrestrial Life
A great restriction attendant on the invasion of the dry land is that locomotion becomes limited to one plane, namely, the surface of the earth. This is in great contrast to what is true in the water, where the animal can move up or down, to right or to left, at any angle and in three dimensions. It surely follows from this that the movements of land animals must be rapid and precise, unless, indeed, safety is secured in some other way. Hence it is easy to understand why most land animals have very finely developed striped muscles, and why a beetle running on the ground has far more numerous muscles than a lobster swimming in the sea.
Land animals were also handicapped by the risks of drought and of frost, but these were met by defences of the most diverse description, from the hairs of woolly caterpillars to the fur of mammals, from the carapace of tortoises to the armour of armadillos. In other cases, it is hardly necessary to say, the difficulties may be met in other ways, as frogs meet the winter by falling into a lethargic state in some secluded retreat.
Another consequence of getting on to dry land is that the eggs or young can no longer be set free anyhow, as is possible when the animal is surrounded by water, which is in itself more or less of a cradle. If the eggs were laid or the young liberated on dry ground, the chances are many that they would be dried up or devoured. So there are numerous ways in which land animals secure the safety of their young, e.g. by burying them in the ground, or by hiding them in nests, or by carrying them about for a prolonged period either before or after birth. This may mean great safety for the young, this may make it possible to have only a small family, and this may tend to the evolution of parental care and the kindly emotions. Thus it may be understood that from the conquest of the land many far-reaching consequences have followed.
Finally, it is worth dwelling on the risks of terrestrial life, because they enable us better to understand why so many land animals have become burrowers and others climbers of trees, why some have returned to the water and others have taken to the air. It may be asked, perhaps, why the land should have been colonised at all when the risks and difficulties are so great. The answer must be that necessity and curiosity are the mother and father of invention. Animals left the water because the pools dried up, or because they were overcrowded, or because of inveterate enemies, but also because of that curiosity and spirit of adventure which, from first to last, has been one of the spurs of progress.
Conquering the Air
6. The last great haunt of life is the air, a mastery of which must be placed to the credit of insects, Pterodactyls, birds, and bats. These have been the successes, but it should be noted that there have been many brilliant failures, which have not attained to much more than parachuting. These include the Flying Fishes, which take leaps from the water and are carried for many yards and to considerable heights, holding their enlarged pectoral fins taut or with little more than a slight fluttering. There is a so-called Flying Frog (Rhacophorus) that skims from branch to branch, and the much more effective Flying Dragon (Draco volans) of the Far East, which has been mentioned already. Among mammals there are Flying Phalangers, Flying Lemurs, and more besides, all attaining to great skill as parachutists, and illustrating the endeavour to master the air which man has realised in a way of his own.
The power of flight brings obvious advantages. A bird feeding on the ground is able to evade the stalking carnivore by suddenly rising into the air; food and water can be followed rapidly and to great distances; the eggs or the young can be placed in safe situations; and birds in their migrations have made a brilliant conquest both of time and space. Many of them know no winter in their year, and the migratory flight of the Pacific Golden Plover from Hawaii to Alaska and back again does not stand alone.