Zoological Philosophy (1809)
J. B. Lamarck



WE are not here concerned with an argument, but with the examination of a positive factÑa fact which is of more general application than is supposed, and which has not received the attention that it deserves, no doubt because it is usually very difficult to recognise. This fact consists in the influence that is exerted by the environment on the various living bodies exposed to it.

It is indeed long since the influence of the various states of our organisation on our character, inclinations, activities and even ideas has been recognised; but I do not think that anyone has yet drawn attention to the influence of our activities and habits even on our organisation. Now since these activities and habits depend entirely on the environment in which we are habitually placed, I shall endeavour to show how great is the influence exerted by that environment on the general shape, state of the parts and even organisation of living bodies. It is, then, with this very positive fact that we have to do in the present chapter.

If we had not had many opportunities of clearly recognising the result of this influence on certain living bodies that we have transported into an environment altogether new and very different from that in which they were previously placed, and if we had not seen the resulting effects and alterations take place almost under our very eyes, the important fact in question would have remained for ever unknown to us.

The influence of the environment as a matter of fact is in all times and places operative on living bodies; but what makes this influence difficult to perceive is that its effects only become perceptible or recognisable (especially in animals) after a long period of time.

Before setting forth to examine the proofs of this fact, which deserves our attention and is so important for zoological philosophy, let us sum up the thread of the discussions that we have already begun.

In the preceding chapter we saw that it is now an unquestionable fact that on passing along the animal scale in the opposite direction from that of nature, we discover the existence, in the groups composing this scale, of a continuous but irregular degradation in the organisation of animals, an increasing simplification in their organisation, and, lastly, a corresponding diminution in the number of their faculties.

This well-ascertained fact may throw the strongest light over the actual order followed by nature in the production of all the animals that she has brought into existence, but it does not show us why the increasing complexity of the organisation of animals from the most imperfect to the most perfect exhibits only an irregular gradation, in the course of which there occur numerous anomalies or deviations with a variety in which no order is apparent.

Now on seeking the reason of this strange irregularity in the increasing complexity of animal organisation, if we consider the influence that is exerted by the infinitely varied environments of all parts of the world on the general shape, structure and even organisation of these animals, all will then be clearly explained.

It will in fact become clear that the state in which we find any animal, is, on the one hand, the result of the increasing complexity of organisation tending to form a regular gradation; and, on the other hand, of the influence of a multitude of very various conditions ever tending to destroy the regularity in the gradation of the increasing complexity of organisation.

I must now explain what I mean by this statement: the environment affects the shape and organisation of animals, that is to say that when the environment becomes very different, it produces in course of time corresponding modifications in the shape and organisation of animals.

It is true if this statement were to be taken literally, I should be convicted of an error; for, whatever the environment may do, it does not work any direct modification whatever in the shape and organisation of animals.

But great alterations in the environment of animals lead to great alterations in their needs, and these alterations in their needs necessarily lead to others in their activities. Now if the new needs become permanent, the animals then adopt new habits which last as long as the needs that evoked them. This is easy to demonstrate, and indeed requires no amplification.

It is then obvious that a great and permanent alteration in the environment of any race of animals induces new habits in these animals.

Now, if a new environment, which has become permanent for some race of animals, induces new habits in these animals, that is to say, leads them to new activities which become habitual, the result will be the use of some one part in preference to some other part, and in some cases the total disuse of some part no longer necessary.

Nothing of all this can be considered as hypothesis or private opinion; on the contrary, they are truths which, in order to be made clear, only require attention and the observation of facts.

We shall shortly see by the citation of known facts in evidence, in the first place, that new needs which establish a necessity for some part really bring about the existence of that part, as a result of efforts and that subsequently its continued use gradually strengthens, develops and finally greatly enlarges it; in the second place, we shall see that in some cases, when the new environment and the new needs have altogether destroyed the utility of some part, the total disuse of that part has resulted in its gradually ceasing to share in the development of the other parts of the animal; it shrinks and wastes little by little, and ultimately, when there has been total disuse for a long period, the part in question ends by disappearing. All this is positive; I propose to furnish the most convincing proofs of it.

In plants, where there are no activities and consequently no habits, properly so‑called, great changes of environment none the less lead to great differences in the development of their parts; so that these differences cause the origin and development of some, and the shrinkage and disappearance of others. But all this is here brought about by the changes sustained in the nutrition of the plant, in its absorption and transpiration, in the quantity of calorie, light, air and moisture that it habitually receives; lastly, in the dominance that some of the various vital movements acquire over others.

Among individuals of the same species, some of which are continually well fed and in an environment favourable to their development, while others are in an opposite environment, there arises a difference in the state of the individuals which gradually becomes very remarkable. How many examples I might cite both in animals and plants which bear out the truth of this principle. Now if the environment remains constant, so that the condition of the ill‑fed, suffering or sickly individuals becomes permanent, their internal organisation is ultimately modified, and these acquired modifications are preserved by reproduction among the individuals in question, and finally give rise to a race quite distinct from that in which the individuals have been continuously in an environment favourable to their development.

A very dry spring causes the grasses of a meadow to grow very little, and remain lean and puny; so that they flower and fruit after accomplishing very little growth.

A spring intermingled with warm and rainy days causes a strong growth in this same grass, and the crop is then excellent.

But if anything causes a continuance of the unfavourable environment, a corresponding variation takes place in the plants: first in their general appearance and condition, and then in some of their special characters.

Suppose, for instance, that a seed of one of the meadow grasses in question is transported to an elevated place on a dry, barren and stony plot much exposed to the winds, and is there left to germinate; if the plant can live in such a place, it will always be badly nourished, and if the individuals reproduced from it continue to exist in this bad environment, there will result a race fundamentally different from that which lives in the meadows and from which it originated. The individuals of this new race will have small and meagre parts; some of their organs will have developed more than others, and will then be of unusual proportions.

Those who have observed much and studied large collections, have acquired the conviction that according as changes occur in environment, situation, climate, food, habits of life, etc., corresponding changes in the animals likewise occur in size, shape, proportions of the parts, colour, consistency, swiftness and skill.

What nature does in the course of long periods we do every day when we suddenly change the environment in which some species of living plant is situated.

Every botanist knows that plants which are transported from their native places to gardens for purposes of cultivation, gradually undergo changes which ultimately make them unrecognisable. Many plants, by nature hairy, become glabrous or nearly so; a number of those which used to lie and creep on the ground, become erect; others lose their thorns or excrescences; others again whose stem was perennial and woody in their native hot climates, become herbaceous in our own climates and some of them become annuals; lastly, the size of their parts itself undergoes very considerable changes. These effects of alterations of environment are so widely recognised, that botanists do not like to describe garden plants unless they have been recently brought into cultivation.

Is it not the case that cultivated wheat (Triticum sativum) is a plant which man has brought to the state in which we now see it? I should like to know in what country such a plant lives in nature, otherwise than as the result of cultivation.

Where in nature do we find our cabbages, lettuces, etc., in the same state as in our kitchen gardens? and is not the case the same with regard to many animals which have been altered or greatly modified by domestication?

How many different races of our domestic fowls and pigeons have we obtained by rearing them in various environments and different countries; birds which we should now vainly seek in nature?

Those which have changed the least, doubtless because their domestication is of shorter standing and because they do not live in a foreign climate, none the less display great differences in some of their parts, as a result of the habits which we have made them contract. Thus our domestic ducks and geese are of the same type as wild ducks and geese; but ours have lost the power of rising into high regions of the air and flying across large tracts of country; moreover, a real change has come about in the state of their parts, as compared with those of the animals of the race from which they come.

Who does not know that if we rear some bird of our own climate in a cage and it lives there for five or six years, and if we then return it to nature by setting it at liberty, it is no longer able to fly like its fellows, which have always been free? The slight change of environment for this individual has indeed only diminished its power of flight, and doubtless has worked no change in its structure; but if a long succession of generations of individuals of the same race had been kept in captivity for a considerable period, there is no doubt that even the structure of these individuals would gradually have undergone notable changes. Still more, if instead of a mere continuous captivity, this environmental factor had been further accompanied by a change to a very different climate; and if these individuals had by degrees been habituated to other kinds of food and other activities for seizing it, these factors when combined together and become permanent would have unquestionably given rise imperceptibly to a new race with quite special characters.

Where in natural conditions do we find that multitude of races of dogs which now actually exist, owing to the domestication to which we have reduced them? Where do we find those bull-dogs, greyhounds, water-spaniels, spaniels, lap-dogs, etc., etc.; races which show wider differences than those which we call specific when they occur among animals of one genus living in natural freedom?

No doubt a single, original race, closely resembling the wolf, if indeed it was not actually the wolf, was at some period reduced by man to domestication. That race, of which all the individuals were then alike, was gradually scattered with man into different countries and climates; and after they had been subjected for some time to the influences of their environment and of the various habits which had been forced upon them in each country, they underwent remarkable alterations and formed various special races. Now man travels about to very great distances, either for trade or any other purpose and thus brings into thickly populated places, such as a great capital, various races of dogs formed in very distant countries. The crossing of these races by reproduction then gave rise in turn to all those that we now know.

The following fact proves in the case of plants how the change of some important factor leads to alteration in the parts of these living bodies.

So long as Ranunculus aquatilis is submerged in the water, all its leaves are finely divided into minute segments; but when the stem of this plant reaches the surface of the water, the leaves which develop in the air are large, round and simply lobed. If several feet of the same plant succeed in growing in a soil that is merely damp without any immersion, their stems are then short, and none of their leaves are broken up into minute divisions, so that we get Ranunculus hederaceus, which botanists regard as a separate species.

There is no doubt that in the case of animals, extensive alterations in their customary environment produce corresponding alterations in their parts; but here the transformations take place much more slowly than in the case of plants; and for us therefore they are less perceptible and their cause less readily identified.

As to the conditions which have so much power in modifying the organs of living bodies, the most potent doubtless consist in the diversity of the places where they live, but there are many others as well which exercise considerable influence in producing the effects in question.

It is known that localities differ as to their character and quality, by reason of their position, construction and climate: as is readily perceived on passing through various localities distinguished by special qualities; this is one cause of variation for animals and plants living in these various places. But what is not known so well and indeed what is not generally believed, is that every locality itself changes in time as to exposure, climate, character and quality, although with such extreme slowness, according to our notions, that we ascribe to it complete stability.

Now in both cases these altered localities involve a corresponding alteration in the environment of the living bodies that dwell there, and this again brings a new influence to bear on these same bodies.

Hence it follows that if there are extremes in these alterations, there are also finer differences: that is to say, intermediate stages which fill up the interval. Consequently there are also fine distinctions between what we call species.

It is obvious then that as regards the character and situation of the substances which occupy the various parts of the earth's surface, there exists a variety of environmental factors which induces a corresponding variety in the shapes and structure of animals, independent of that special variety which necessarily results from the progress of the complexity of organisation in each animal.

In every locality where animals can live, the conditions constituting any one order of things remain the same for long periods: indeed they alter so slowly that man cannot directly observe it. It is only by an inspection of ancient monuments that he becomes convinced that in each of these localities the order of things which he now finds has not always been existent; he may thence infer that it will go on changing.

Races of animals living in any of these localities must then retain their habits equally long: hence the apparent constancy of the races that we call species,Ña constancy which has raised in us the belief that these races are as old as nature.

But in the various habitable parts of the earth's surface, the character and situation of places and climates constitute both for animals and plants environmental influences of extreme variability. The animals living in these various localities must therefore differ among themselves, not only by reason of the state of complexity of organisation attained in each race, but also by reason of the habits which each race is forced to acquire; thus when the observing naturalist travels over large portions of the earth's surface and sees conspicuous changes occurring in the environment, he invariably finds that the characters of species undergo a corresponding change.

Now the true principle to be noted in all this is as follows:

1. Every fairly considerable and permanent alteration in the environment of any race of animals works a real alteration in the needs of that race.

2. Every change in the needs of animals necessitates new activities on their part for the satisfaction of those needs, and hence new habits.

3. Every new need, necessitating new activities for its satisfaction, requires the animal, either to make more frequent use of some of its parts which it previously used less, and thus greatly to develop and enlarge them; or else to make use of entirely new parts, to which the needs have imperceptibly given birth by efforts of its inner feeling; this I shall shortly prove by means of known facts.

Thus to obtain a knowledge of the true causes of that great diversity of shapes and habits found in the various known animals, we must reflect that the infinitely diversified but slowly changing environment in which the animals of each race have successively been placed, has involved each of them in new needs and corresponding alterations in their habits. This is a truth which, once recognised, cannot be disputed. Now we shall easily discern how the new needs may have been satisfied, and the new habits acquired, if we pay attention to the two following laws of nature, which are always verified by observation.


In every animal which has not passed the limit of its development, a more frequent and continuous use of any organ gradually strengthens, develops and enlarges that organ, and gives it a power proportional to the length of time it has been so used; while the permanent disuse of any organ imperceptibly weakens and deteriorates it, and progressively diminishes its functional capacity, until it finally disappears.


All the acquisitions or losses wrought by nature on individuals, through the influence of the environment in which their race has long been placed, and hence through the influence of the predominant use or permanent disuse of any organ; all these are preserved by reproduction to the new individuals which arise, provided that the acquired modifications are common to both sexes, or at least to the individuals which produce the young.

Here we have two permanent truths, which can only be doubted by those who have never observed or followed the operations of nature, or by those who have allowed themselves to be drawn into the error which I shall now proceed to combat.

Naturalists have remarked that the structure of animals is always in perfect adaptation to their functions, and have inferred that the shape and condition of their parts have determined the use of them. Now this is a mistake: for it may be easily proved by observation that it is on the contrary the needs and uses of the parts which have caused the development of these same parts, which have even given birth to them when they did not exist, and which consequently have given rise to the condition that we find in each animal.

If this were not so, nature would have had to create as many different kinds of structure in animals, as there are different kinds of environment in which they have to live; and neither structure nor environment would ever have varied.

This is indeed far from the true order of things. If things were really so, we should not have race-horses shaped like those in England; we should not have big draught-horses so heavy and so different from the former, for none such are produced in nature; in the same way we should not have basset-hounds with crooked legs, nor grey-hounds so fleet of foot, nor water-spaniels, etc.; we should not have fowls without tails, fantail pigeons, etc.; finally, we should be able to cultivate wild plants as long as we liked in the rich and fertile soil of our gardens, without the fear of seeing them change under long cultivation.

A feeling of the truth in this respect has long existed; since the following maxim has passed into a proverb and is known by all, Habits form a second nature.

Assuredly if the habits and nature of each animal could never vary, the proverb would have been false and would not have come into existence, nor been preserved in the event of any one suggesting it.

If we seriously reflect upon all that I have just set forth, it will be seen that I was entirely justified when in my work entitled Recherche sur les corps vivants (p. 50), I established the following proposition:

"It is not the organs, that is to say, the nature and shape of the parts of an animal's body, that have given rise to its special habits and faculties; but it is, on the contrary, its habits, mode of life and environment that have in course of time controlled the shape of its body, the number and state of its organs and, lastly, the faculties which it possesses."

If this proposition is carefully weighed and compared with all the observations that nature and circumstances are incessantly throwing in our way, we shall see that its importance and accuracy are substantiated in the highest degree.

Time and a favourable environment are as I have already said nature's two chief methods of bringing all her productions into existence: for her, time has no limits and can be drawn upon to any extent.

As to the various factors which she has required and still constantly uses for introducing variations in everything that she produces, they may be described as practically inexhaustible.

The principal factors consist in the influence of climate, of the varying temperatures of the atmosphere and the whole environment, of the variety of localities and their situation, of habits, the commonest movements, the most frequent activities, and, lastly, of the means of self-preservation, the mode of life and the methods of defence and multiplication.

Now as a result of these various influences, the faculties become extended and strengthened by use, and diversified by new habits that are long kept up. The conformation, consistency and, in short, the character and state of the parts, as well as of the organs, are imperceptibly affected by these influences and are preserved and propagated by reproduction.

These truths, which are merely effects of the two natural laws stated above, receive in every instance striking confirmation from facts; for the facts afford a clear indication of nature's procedure in the diversity of her productions.

But instead of being contented with generalities which might be considered hypothetical, let us investigate the facts directly, and consider the effects in animals of the use or disuse of their organs on these same organs, in accordance with the habits that each race has been forced to contract.

Now I am going to prove that the permanent disuse of any organ first decreases its functional capacity, and then gradually reduces the organ and causes it to disappear or even become extinct, if this disuse lasts for a very long period throughout successive generations of animals of the same race.

I shall then show that the habit of using any organ, on the contrary, in any animal which has not reached the limit of the decline of its functions, not only perfects and increases the functions of that organ, but causes it in addition to take on a size and development which imperceptibly alter it; so that in course of time it becomes very different from the same organ in some other animal which uses it far less.

The permanent disuse of an organ, arising from a change of habits, causes a gradual shrinkage and ultimately the disappearance and even extinction of that organ.

Since such a proposition could only be accepted on proof, and not on mere authority, let us endeavour to make it clear by citing the chief known facts which substantiate it.

The vertebrates, whose plan of organisation is almost the same throughout, though with much variety in their parts, have their jaws armed with teeth; some of them, however, whose environment has induced the habit of swallowing the objects they feed on without any preliminary mastication, are so affected that their teeth do not develop. The teeth then remain hidden in the bony framework of the jaws, without being able to appear outside; or indeed they actually become extinct down to their last rudiments.

In the right-whale, which was supposed to be completely destitute of teeth, M. Geoffroy has nevertheless discovered teeth concealed in the jaws of the foetus of this animal. The professor has moreover discovered in birds the groove in which the teeth should be placed, though they are no longer to be found there.

Even in the class of mammals, comprising the most perfect animals, where the vertebrate plan of organisation is carried to its highest completion, not only is the right-whale devoid of teeth, but the ant-eater (Myrmecophaga) is also found to be in the same condition, since it has acquired a habit of carrying out no mastication, and has long preserved this habit in its race.

Eyes in the head are characteristic of a great number of different animals, and essentially constitute a part of the plan of organisation of the vertebrates.

Yet the mole, whose habits require a very small use of sight, has only minute and hardly visible eyes, because it uses that organ so little.

Olivier's Spalax (Voyage en Égypte et en Perse), which lives underground like the mole, and is apparently exposed to daylight even less than the mole, has altogether lost the use of sight: so that it shows nothing more than vestiges of this organ. Even these vestiges are entirely hidden under the skin and other parts, which cover them up and do not leave the slightest access to light.

The Proteus, an aquatic reptile allied to the salamanders, and living in deep dark caves under the water, has, like the Spalax, only vestiges of the organ of sight, vestiges which are covered up and hidden in the same way.

The following consideration is decisive on the question which I am now discussing,

Light does not penetrate everywhere; consequently animals which habitually live in places where it does not penetrate, have no opportunity of exercising their organ of sight, if nature has endowed them with one. Now animals belonging to a plan of organisation of which eyes were a necessary part, must have originally had them. Since, however, there are found among them some which have lost the use of this organ and which show nothing more than hidden and covered up vestiges of them, it becomes clear that the shrinkage and even disappearance of the organ in question are the results of a permanent disuse of that organ.

This is proved by the fact that the organ of hearing is never in this condition, but is always found in animals whose organisation is of the kind that includes it: and for the following reason.

1Physicists believe and even affirm that the atmospheric air is the actual substance of sound, that is to say, that it is the substance which, when set in motion by the shocks or vibrations of bodies, transmits to the organ of hearing the impression of the concussions received.
   That this is an error is attested by many known facts, showing that it is impossible that the air should penetrate to all places to which the substance producing sound actually does penetrate.
   See my memoir On the Substance of Sound, printed at the end of my HydrogŽologie, p. 225, in which I furnished the proofs of this mistake.
   Since the publication of my memoir, which by the way is seldom cited, great efforts have been made to make the known velocity of the propagation of sound in air tally with the elasticity of the air, which would cause the propagation of its oscillations to be too slow for the theory. Now, since the air during oscillation necessarily undergoes alternate compressions and dilatations in its parts, recourse has been had to the effects of the caloric squeezed out during the sudden compressions of the air and of the caloric absorbed during the rarefactions of that fluid. By means of these effects, quantitatively determined by convenient hypotheses, geometricians now account for the velocity with which sound is propagated through air. But this is no answer to the fact that sound is also propagated through bodies which air can neither traverse nor set in motion.
   These physicists assume forsooth a vibration in the smallest particles of solid bodies; a vibration of very dubious existence, since it can only be propagated through homogeneous bodies of equal density, and cannot spread from a dense body to a rarefied one or vice versâ. Such a hypothesis offers no explanation of the well-known fact that sound is propagated through heterogeneous bodies of very different densities and kinds.

The substance of sound,1 that namely which, when set in motion by the shock or the vibration of bodies, transmits to the organ of hearing the impression received, penetrates everywhere and passes through any medium, including even the densest bodies: it follows that every animal, belonging to a plan of organisation of which hearing is an essential part, always has some opportunity for the exercise of this organ wherever it may live. Hence among the vertebrates we do not find any that are destitute of the organ of hearing; and after them, when this same organ has come to an end, it does not subsequently recur in any animal of the posterior classes.

It is not so with the organ of sight; for this organ is found to disappear, re-appear and disappear again according to the use that the animal makes of it.

In the acephalic molluscs, the great development of the mantle would make their eyes and even their head altogether useless. The permanent disuse of these organs has thus brought about their disappearance and extinction, although molluscs belong to a plan of organisation which should comprise them.

Lastly, it was part of the plan of organisation of the reptiles, as of other vertebrates, to have four legs in dependence on their skeleton. Snakes ought consequently to have four legs, especially since they are by no means the last order of the reptiles and are farther from the fishes than are the batrachians (frogs, salamanders, etc.).

Snakes, however, have adopted the habit of crawling on the ground and hiding in the grass; so that their body, as a result of continually repeated efforts at elongation for the purpose of passing through narrow spaces, has acquired a considerable length, quite out of proportion to its size. Now, legs would have been quite useless to these animals and consequently unused. Long legs would have interfered with their need of crawling, and very short legs would have been incapable of moving their body, since they could only have had four. The disuse of these parts thus became permanent in the various races of these animals, and resulted in the complete disappearance of these same parts, although legs really belong to the plan of organisation of the animals of this class.

Many insects, which should have wings according to the natural characteristics of their order and even of their genus, are more or less completely devoid of them through disuse. Instances are furnished by many Coleoptera, Orthoptera, Hymenoptera and Hemiptera, etc., where the habits of these animals never involve them in the necessity of using their wings.

But it is not enough to give an explanation of the cause which has brought about the present condition of the organs of the various animals,Ña condition that is always found to be the same in animals of the same species; we have in addition to cite instances of changes wrought in the organs of a single individual during its life, as the exclusive result of a great mutation in the habits of the individuals of its species. The following very remarkable fact will complete the proof of the influence of habits on the condition of the organs, and of the way in which permanent changes in the habits of an individual lead to others in the condition of the organs, which come into action during the exercise of these habits.

M. Tenon, a member of the Institute, has notified to the class of sciences, that he had examined the intestinal canal of several men who had been great drinkers for a large part of their lives, and in every case he had found it shortened to an extraordinary degree, as compared with the same organ in all those who had not adopted the like habit.

It is known that great drinkers, or those who are addicted to drunkenness, take very little solid food, and eat hardly anything; since the drink which they consume so copiously and frequently is sufficient to feed them.

Now since fluid foods, especially spirits, do not long remain either in the stomach or intestine, the stomach and the rest of the intestinal canal lose among drinkers the habit of being distended, just as among sedentary persons, who are continually engaged on mental work and are accustomed to take very little food; for in their case also the stomach slowly shrinks and the intestine shortens.

This has nothing to do with any shrinkage or shortening due to a binding of the parts which would permit of the ordinary extension, if instead of remaining empty these viscera were again filled; we have to do with a real shrinkage and shortening of considerable extent, and such that these organs would burst rather than yield at once to any demand for the ordinary extension.

Compare two men of equal ages, one of whom has contracted the habit of eating very little, since his habitual studies and mental work have made digestion difficult, while the other habitually takes much exercise, is often out-of-doors, and eats well; the stomach of the first will have very little capacity left and will be filled up by a very small quantity of food, while that of the second will have preserved and even increased its capacity.

Here then is an organ which undergoes profound modification in size and capacity, purely on account of a change of habits during the life of the individual.

The frequent use of any organ, when confirmed by habit, increases the functions of that organ, leads to its development and endows it with a size and power that it does not possess in animals which exercise it less.

We have seen that the disuse of any organ modifies, reduces and finally extinguishes it. I shall now prove that the constant use of any organ, accompanied by efforts to get the most out of it, strengthens and enlarges that organ, or creates new ones to carry on functions that have become necessary.

The bird which is drawn to the water by its need of finding there the prey on which it lives, separates the digits of its feet in trying to strike the water and move about on the surface. The skin which unites these digits at their base acquires the habit of being stretched by these continually repeated separations of the digits; thus in course of time there are formed large webs which unite the digits of ducks, geese, etc., as we actually find them. In the same way efforts to swim, that is to push against the water so as to move about in it, have stretched the membranes between the digits of frogs, sea-tortoises, the otter, beaver, etc.

On the other hand, a bird which is accustomed to perch on trees and which springs from individuals all of whom had acquired this habit, necessarily has longer digits on its feet and differently shaped from those of the aquatic animals that I have just named. Its claws in time become lengthened, sharpened and curved into hooks, to clasp the branches on which the animal so often rests.

We find in the same way that the bird of the water-side which does not like swimming and yet is in need of going to the water's edge to secure its prey, is continually liable to sink in the mud. Now this bird tries to act in such a way that its body should not be immersed in the liquid, and hence makes its best efforts to stretch and lengthen its legs. The long-established habit acquired by this bird and all its race of continually stretching and lengthening its legs, results in the individuals of this race becoming raised as though on stilts, and gradually obtaining long, bare legs, denuded of feathers up to the thighs and often higher still. (Systmedes Animaux sans vertbres, p. 14.)

We note again that this same bird wants to fish without wetting its body, and is thus obliged to make continual efforts to lengthen its neck. Now these habitual efforts in this individual and its race must have resulted in course of time in a remarkable lengthening, as indeed we actually find in the long necks of all water-side birds.

If some swimming birds like the swan and goose have short legs and yet a very long neck, the reason is that these birds while moving about on the water acquire the habit of plunging their head as deeply as they can into it in order to get the aquatic larvae and various animals on which they feed; whereas they make no effort to lengthen their legs.

If an animal, for the satisfaction of its needs, makes repeated efforts to lengthen its tongue, it will acquire a considerable length (ant-eater, green-woodpecker); if it requires to seize anything with this same organ, its tongue will then divide and become forked. Proofs of my statement are found in the humming-birds which use their tongues for grasping things, and in lizards and snakes which use theirs to palpate and identify objects in front of them.

Needs which are always brought about by the environment, and the subsequent continued efforts to satisfy them, are not limited in their results to a mere modification, that is to say, an increase or decrease of the size and capacity of organs; but they may even go so far as to extinguish organs, when any of these needs make such a course necessary.

Fishes, which habitually swim in large masses of water, have need of lateral vision; and, as a matter of fact, their eyes are placed on the sides of their head. Their body, which is more or less flattened according to the species, has its edges perpendicular to the plane of the water; and their eyes are placed so that there is one on each flattened side. But such fishes as are forced by their habits to be constantly approaching the shore, and especially slightly inclined or gently sloping beaches, have been compelled to swim on their flattened surfaces in order to make a close approach to the water's edge. In this position, they receive more light from above than below and stand in special need of paying constant attention to what is passing above them; this requirement has forced one of their eyes to undergo a sort of displacement, and to assume the very remarkable position found in the soles, turbots, dabs, etc. (Pleuronectes and Achirus). The position of these eyes is not symmetrical, because it results from an incomplete mutation. Now this mutation is entirely completed in the skates, in which the transverse flattening of the body is altogether horizontal, like the head. Accordingly the eyes of skates are both situated on the upper surface and have become symmetrical.

Snakes, which crawl on the surface of the earth, chiefly need to see objects that are raised or above them. This need must have had its effect on the position of the organ of sight in these animals, and accordingly their eyes are situated in the lateral and upper parts of their head, so as easily to perceive what is above them or at their sides; but they scarcely see at all at a very short distance in front of them. They are, however, compelled to make good the deficiency of sight as regards objects in front of them which might injure them as they move forward. For this purpose they can only use their tongue, which they are obliged to thrust out with all their might. This habit has not only contributed to making their tongue slender and very long and contractile, but it has even forced it to undergo division in the greater number of species, so as to feel several objects at the same time it has even permitted of the formation of an aperture at the extremity of their snout, to allow the tongue to pass without having to separate the jaws.

Nothing is more remarkable than the effects of habit in herbivorous mammals.

A quadruped, whose environment and consequent needs have for long past inculcated the habit of browsing on grass, does nothing but walk about on the ground; and for the greater part of its life is obliged to stand on its four feet, generally making only few or moderate movements. The large portion of each day that this kind of animal has to pass in filling itself with the only kind of food that it cares for, has the result that it moves but little and only uses its feet for support in walking or running on the ground, and never for holding on, or climbing trees.

From this habit of continually consuming large quantities of foodmaterial, which distend the organs receiving it, and from the habit of making only moderate movements, it has come about that the body of these animals has greatly thickened, become heavy and massive and acquired a very great size: as is seen in elephants, rhinoceroses, oxen, buffaloes, horses, etc.

The habit of standing on their four feet during the greater part of the day, for the purpose of browsing, has brought into existence a thick horn which invests the extremity of their digits; and since these digits have no exercise and are never moved and serve no other purpose than that of support like the rest of the foot, most of them have become shortened, dwindled and, finally, even disappeared.

Thus in the pachyderms, some have five digits on their feet invested in horn, and their hoof is consequently divided into five parts; others have only four, and others again not more than three; but in the ruminants, which are apparently the oldest of the mammals that are permanently confined to the ground, there are not more than two digits on the feet and indeed, in the solipeds, there is only one (horse, donkey).

Nevertheless some of these herbivorous animals, especially the ruminants, are incessantly exposed to the attacks of carnivorous animals in the desert countries that they inhabit, and they can only find safety in headlong flight. Necessity has in these cases forced them to exert themselves in swift running, and from this habit their body has become more slender and their legs much finer; instances are furnished by the antelopes, gazelles, etc.

In our own climates, there are other dangers, such as those constituted by man, with his continual pursuit of red deer, roe deer and fallow deer; this has reduced them to the same necessity, has impelled them into similar habits, and had corresponding effects.

Since ruminants can only use their feet for support, and have little strength in their jaws, which only obtain exercise by cutting and browsing on the grass, they can only fight by blows with their heads, attacking one another with their crowns.

In the frequent fits of anger to which the males especially are subject, the efforts of their inner feeling cause the fluids to flow more strongly towards that part of their head; in some there is hence deposited a secretion of horny matter, and in others of bony matter mixed with horny matter, which gives rise to solid protuberances: thus we have the origin of horns and antlers, with which the head of most of these animals is armed.

It is interesting to observe the result of habit in the peculiar shape and size of the giraffe (Camelo-pardalis): this animal, the largest of the mammals, is known to live in the interior of Africa in places where the soil is nearly always arid and barren, so that it is obliged to browse on the leaves of trees and to make constant efforts to reach them. From this habit long maintained in all its race, it has resulted that the animal's fore-legs have become longer than its hind legs, and that its neck is lengthened to such a degree that the giraffe, without standing up on its hind legs, attains a height of six metres (nearly 20 feet).

Among birds, ostriches, which have no power of flight and are raised on very long legs, probably owe their singular shape to analogous circumstances.

The effect of habit is quite as remarkable in the carnivorous mammals as in the herbivores; but it exhibits results of a different kind.

Those carnivores, for instance, which have become accustomed to climbing, or to scratching the ground for digging holes, or to tearing their prey, have been under the necessity of using the digits of their feet: now this habit has promoted the separation of their digits, and given rise to the formation of the claws with which they are armed.

But some of the carnivores are obliged to have recourse to pursuit in order to catch their prey: now some of these animals were compelled by their needs to contract the habit of tearing with their claws, which they are constantly burying deep in the body of another animal in order to lay hold of it, and then make efforts to tear out the part seized. These repeated efforts must have resulted in its claws reaching a size and curvature which would have greatly impeded them in walking or running on stony ground: in such cases the animal has been compelled to make further efforts to draw back its claws, which are so projecting and hooked as to get in its way. From this there has gradually resulted the formation of those peculiar sheaths, into which cats, tigers, lions, etc. withdraw their claws when they are not using them.

Hence we see that efforts in a given direction, when they are long sustained or habitually made by certain parts of a living body, for the satisfaction of needs established by nature or environment, cause an enlargement of these parts and the acquisition of a size and shape that they would never have obtained, if these efforts had not become the normal activities of the animals exerting them. Instances are everywhere furnished by observations on all known animals.

Can there be any more striking instance than that which we find in the kangaroo? This animal, which carries its young in a pouch under the abdomen, has acquired the habit of standing upright, so as to rest only on its hind legs and tail; and of moving only by means of a succession of leaps, during which it maintains its erect attitude in order not to disturb its young. And the following is the result:

1. Its fore legs, which it uses very little and on which it only supports itself for a moment on abandoning its erect attitude, have never acquired a development proportional to that of the other parts, and have remained meagre, very short and with very little strength.

2. The hind legs, on the contrary, which are almost continually in action either for supporting the whole body or for making leaps, have acquired a great development and become very large and strong.

3. Lastly, the tail, which is in this case much used for supporting the animal and carrying out its chief movements, has acquired an extremely remarkable thickness and strength at its base.

These well-known facts are surely quite sufficient to establish the results of habitual use on an organ or any other part of animals. If on observing in an animal any organ particularly well-developed, strong, and powerful, it is alleged that its habitual use has nothing to do with it, that its continued disuse involves it in no loss, and finally, that this organ has always been the same since the creation of the species to which the animal belongs, then I ask, Why can our domestic ducks no longer fly like wild ducks? I can, in short, cite a multitude of instances among ourselves, which bear witness to the differences that accrue to us from the use or disuse of any of our organs, although these differences are not preserved in the new individuals which arise by reproduction: for if they were their effects would be far greater.

I shall show in Part II., that when the will guides an animal to any action, the organs which have to carry out that action are immediately stimulated to it by the influx of subtle fluids (the nervous fluid), which become the determining factor of the movements required. This fact is verified by many observations, and cannot now be called in question.

Hence it follows that numerous repetitions of these organised activities strengthen, stretch, develop and even create the organs necessary to them. We have only to watch attentively what is happening all around us, to be convinced that this is the true cause of organic development and changes.

Now every change that is wrought in an organ through a habit of frequently using it, is subsequently preserved by reproduction, if it is common to the individuals who unite together in fertilisation for the propagation of their species. Such a change is thus handed on to all succeeding individuals in the same environment, without their having to acquire it in the same way that it was actually created.

Furthermore, in reproductive unions, the crossing of individuals who have different qualities or structures is necessarily opposed to the permanent propagation of these qualities and structures. Hence it is that in man, who is exposed to so great a diversity of environment, the accidental qualities or defects which he acquires are not preserved and propagated by reproduction. If, when certain peculiarities of shape or certain defects have been acquired, two individuals who are both affected were always to unite together, they would hand on the same peculiarities; and if successive generations were limited to such unions, a special and distinct race would then be formed. But perpetual crossings between individuals, who have not the same peculiarities of shape, cause the disappearance of all peculiarities acquired by special action of the environment. Hence, we may be sure that if men were not kept apart by the distances of their habitations, the crossing in reproduction would soon bring about the disappearance of the general characteristics distinguishing different nations.

If I intended here to pass in review all the classes, orders, genera and species of existing animals, I should be able to show that the conformation and structure of individuals, their organs, faculties, etc., etc., are everywhere a pure result of the environment to which each species is exposed by its nature, and by the habits that the individuals composing it have been compelled to acquire; I should be able to show that they are not the result of a shape which existed from the beginning, and has driven animals into the habits they are known to possess.

It is known that the animal called the ai or sloth (Bradypus tridactylus) is permanently in a state of such extreme weakness that it only executes very slow and limited movements, and walks on the ground with difficulty. So slow are its movements that it is alleged that it can only take fifty steps in a day. It is known, moreover, that the organisation of this animal is entirely in harmony with its state of feebleness and incapacity for walking; and that if it wished to make other movements than those which it actually does make it could not do so.

Hence on the supposition that this animal had received its organisation from nature, it has been asserted that this organisation forced it into the habits and miserable state in which it exists.

This is very far from being my opinion; for I am convinced that the habits which the ai was originally forced to contract must necessarily have brought its organisation to its present condition.

If continual dangers in former times have led the individuals of this species to take refuge in trees, to live there habitually and feed on their leaves, it is clear that they must have given up a great number of movements which animals living on the ground are in a position to perform. All the needs of the ai will then be reduced to clinging to branches and crawling and dragging themselves among them, in order to reach the leaves, and then to remaining on the tree in a state of inactivity in order to avoid falling off. This kind of inactivity, moreover, must have been continually induced by the heat of the climate; for among warm-blooded animals, heat is more conducive to rest than to movement.

Now the individuals of the race of the ai have long maintained this habit of remaining in the trees, and of performing only those slow and little varied movements which suffice for their needs. Hence their organisation will gradually have come into accordance with their new habits; and from this it must follow:

1. That the arms of these animals, which are making continual efforts to clasp the branches of trees, will be lengthened;

2. That the claws of their digits will have acquired a great length and a hooked shape, through the continued efforts of the animal to hold on;

3. That their digits, which are never used in making independent movements, will have entirely lost their mobility, become united and have preserved only the faculty of flexion or extension all together

4. That their thighs, which are continually clasping either the trunk or large branches of trees, will have contracted a habit of always being separated, so as to lead to an enlargement of the pelvis and a backward direction of the cotyloid cavities;

5. Lastly, that a great many of their bones will be welded together, and that parts of their skeleton will consequently have assumed an arrangement and form adapted to the habits of these animals, and different from those which they would require for other habits.

This is a fact that can never be disputed; since nature shows us in innumerable other instances the power of environment over habit and that of habit over the shape, arrangement and proportions of the parts of animals.

Since there is no necessity to cite any further examples, we may now turn to the main point elaborated in this discussion.

It is a fact that all animals have special habits corresponding to their genus and species, and always possess an organisation that is completely in harmony with those habits.

It seems from the study of this fact that we may adopt one or other of the two following conclusions, and that neither of them can be verified.

Conclusion adopted hitherto: Nature (or her Author) in creating animals, foresaw all the possible kinds of environment in which they would have to live, and endowed each species with a fixed organisation and with a definite and invariable shape, which compel each species to live in the places and climates where we actually find them, and there to maintain the habits which we know in them.

My individual conclusion: Nature has produced all the species of animals in succession, beginning with the most imperfect or simplest, and ending her work with the most perfect, so as to create a gradually increasing complexity in their organisation; these animals have spread at large throughout all the habitable regions of the globe, and every species has derived from its environment the habits that we find in it and the structural modifications which observation shows us.

The former of these two conclusions is that which has been drawn hitherto, at least by nearly everyone: it attributes to every animal a fixed organisation and structure which never have varied and never do vary; it assumes, moreover, that none of the localities inhabited by animals ever vary; for if they were to vary, the same animals could no longer survive, and the possibility of finding other localities and transporting themselves thither would not be open to them.

The second conclusion is my own: it assumes that by the influence of environment on habit, and thereafter by that of habit on the state of the parts and even on organisation, the structure and organisation of any animal may undergo modifications, possibly very great, and capable of accounting for the actual condition in which all animals are found.

In order to show that this second conclusion is baseless, it must first be proved that no point on the surface of the earth ever undergoes variation as to its nature, exposure, high or low situation, climate, etc., etc.; it must then be proved that no part of animals undergoes even after long periods of time any modification due to a change of environment or to the necessity which forces them into a different kind of life and activity from what has been customary to them.

Now if a single case is sufficient to prove that an animal which has long been in domestication differs from the wild species whence it sprang, and if in any such domesticated species, great differences of conformation are found between the individuals exposed to such a habit and those which are forced into different habits, it will then be certain that the first conclusion is not consistent with the laws of nature, while the second, on the contrary, is entirely in accordance with them.

Everything then combines to prove my statement, namely: that it is not the shape either of the body or its parts which gives rise to the habits of animals and their mode of life; but that it is, on the contrary, the habits, mode of life and all the other influences of the environment which have in course of time built up the shape of the body and of the parts of animals. With new shapes, new faculties have been acquired, and little by little nature has succeeded in fashioning animals such as we actually see them.

Can there be any more important conclusion in the range of natural history, or any to which more attention should be paid than that which I have just set forth?

Let us conclude this Part I. with the principles and exposition of the natural order of animals.

Chapter III

CybeRose note: It has long been claimed that "acquired characters" cannot be inherited. To make this claim it was necessary for Darwinists, Mendelists and neo-Darwinists to discuss only those acquired characters that are not inherited. Weismann, for example, cut off the tails of some mice, bred them and observed that the offspring did not have short tails. This has nothing to do with Lamarck's discussion, though it is evidence against Darwin's theory of "gemmules". If, as Darwin assumed, gemmules are produced in the various organs, then flow to the reproductive parts to contribute to the transmission of heredity, the absence of tails would prevent any "tail gemmules" from being passed along to the next generation.

Lamarck, on the other hand, had nothing to say about hereditary transmisson of injuries. Instead he gave the example of Ranunculus aquatilis/hederaceus — a species that transforms itself from an aquatic form to a terrestrial form according to its current environment. We now call this phenomenon "phenotypic plasticity", and it is of great importance in evolution and in plant (or animal) breeding.

Simply stated, we can learn a great deal about the hereditary potentials of plants (and animals) by raising them in different habitats. For example, if we raise Polyantha roses from cuttings and place each in different soil ranging from poor to rich we will find that in most cases the plants show a great deal of plasticity in the number of flowers per cluster, but much less in the size of the individual flowers. Tea roses, to the contrary, often show much plasticity in flower size, but less in the number of flowers.

If we should cross a Polyantha with a Tea, no doubt we will find some variation in plastic response among the F1 hybrids, and even more among the F2 generation progeny. All the F1 seedlings may appear similar while growing in the same conditions, but if we test them all in varying conditions we will see the variation. At this point we may decide whether we want to breed hardier Tea roses or Polyanthas with more shapely flowers. This decision will lead us to make different selections for continued breeding.

Suppose we want a hardier Tea. We should choose the F1 seedlings that bear the largest flowers and express the greatest plasticity for flower size, a trait inherited from the Tea parent. Crossing among these we can be confident that in the F2 generation most of the offspring will express similar plasticity, though to a larger degree. Another generation or two should bring us Tea-sized blooms.

On the other hand, if we are aiming for improved Polyanthas, we should select for small flowers and plasticity of cluster size. Continuing in this direction and favoring those with the most shapely flowers we soon will have the desired "sweet heart" roses arranged in clusters.

In either case, testing the plasticity of the seedlings will get us to our goal much more quickly and will require smaller populations of seedlings than reliance on mere "Mendelian" segregation.