'Planning' is simply the result of experience read backward and projected into the future. To me the 'purposive' action of a beehive is simply the summation and integration of its units, and Natural Selection has put higher and higher premiums on the most 'purposeful' integration. It is the same way (to me) in the evolution of the middle ear, the steps in the Cynodonts (clearly shown by me in 1910 and by you later in Oudenodon) make it easier to see how such a wonderful device as the middle ear could arise without any predetermination or human-like planning, and in fact in the good old Darwinian way, if only we admit that as the 'twig is bent the tree's inclined' and that each stage conserves the advantages of its predecessors... The simple idea that planning is only experience read backward and combined by selection in suitable or successful combinations takes the mystery out of Nature and out of men's minds.

All appearances to the contrary, the only watchmaker in nature is the blind forces of physics, albeit deployed in very special way. A true watchmaker has foresight: he designs his cogs springs, and plans their interconnections, with a future purpose in his mind's eye. Natural selection, the blind, unconscious, automatic process which Darwin discovered, and which we now know is the explanation for the existence and apparently purposeful form of all life, has no purpose in mind. It has no mind and no mind's eye. It does not plan for the future. It has no vision, no foresight, no sight at all. If it can be said to play the role of watchmaker in nature, it is the blind watchmaker.

All that we can do, is to keep steadily in mind that each organic being is striving to increase at a geometrical ratio; that each at some period of its life, during some season of the year, during each generation or at intervals, has to struggle for life, and to suffer great destruction. When we reflect on this struggle, we may console ourselves with the full belief, that the war of nature is not incessant, that no fear is felt, that death is generally prompt, and that the vigorous, the healthy, and the happy survive and multiply.

As buds give rise by growth to fresh buds, and these, if vigorous, branch out and overtop on all sides many a feebler branch, so by generation I believe it has been with the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever branching and beautiful ramifications.

As natural selection works solely by and for the good of each being, all corporeal and mental endowments will tend to progress towards perfection.

Evolution triumphs because it turns to natural selection, the plodding accumulation of error.

I have called this principle, by which each slight variation, if useful, is preserved, by the term Natural Selection

In its essence, the theory of natural selection is primarily an attempt to give an account of the probable mechanism of the origin of the adaptations of the organisms to their environment, and only secondarily an attempt to explain evolution at large. Some modem biologists seem to believe that the word 'adaptation' has teleological connotations, and should therefore be expunged from the scientific lexicon. With this we must emphatically disagree. That adaptations exist is so evident as to be almost a truism, although this need not mean that ours is the best of all possible worlds. A biologist has no right to close his eyes to the fact that the precarious balance between a living being and its environment must be preserved by some mechanism or mechanisms if life is to endure.

In the study of this membrane [the retina] I for the first time felt my faith in Darwinism (hypothesis of natural selection) weakened, being amazed and confounded by the supreme constructive ingenuity revealed not only in the retina and in the dioptric apparatus of the vertebrates but even in the meanest insect eye. ... I felt more profoundly than in any other subject of study the shuddering sensation of the unfathomable mystery of life.

In true natural selection, if a body has what it takes to survive, its genes automatically survive because they are inside it. So the genes that survive tend to be, automatically, those genes that confer on bodies the qualities that assist them to survive.

It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us. These laws, taken in the largest sense, being Growth with Reproduction; Inheritance which is almost implied by reproduction; Variability from the indirect and direct action of the external conditions of life, and from use and disuse; a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to Natural Selection, entailing Divergence of Character and the Extinction of less-improved forms. Thus, from the war of nature, from famine and death, the most exalted object which we are capable of conceiving, namely, the production of the higher animals, directly follows. There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.

It is natural selection that gives direction to changes, orients chance, and slowly, progressively produces more complex structures, new organs, and new species. Novelties come from previously unseen association of old material. To create is to recombine.

It may metaphorically be said that natural selection is daily and hourly scrutinising, throughout the world, the slightest variations; rejecting those that are bad, preserving and adding up all that are good; silently and insensibly working, whenever and wherever opportunity offers, at the improvement of each organic being in relation to its organic and inorganic conditions of life. We see nothing of these slow changes in progress, until the hand of time has marked the lapse of ages...

It may very properly be asked whether the attempt to define distinct species, of a more or less permanent nature, such as we are accustomed to deal with amongst the higher plants and animals, is not altogether illusory amongst such lowly organised forms of life as the bacteria. No biologist nowadays believes in the absolute fixity of species ... but there are two circumstances which here render the problem of specificity even more difficult of solution. The bacteriologist is deprived of the test of mutual fertility or sterility, so valuable in determining specific limits amongst organisms in which sexual reproduction prevails. Further, the extreme rapidity with which generation succeeds generation amongst bacteria offers to the forces of variation and natural selection a field for their operation wholly unparalleled amongst higher forms of life.

It will be noticed that the fundamental theorem proved above bears some remarkable resemblances to the second law of thermodynamics. Both are properties of populations, or aggregates, true irrespective of the nature of the units which compose them; both are statistical laws; each requires the constant increase of a measurable quantity, in the one case the entropy of a physical system and in the other the fitness, measured by m, of a biological population. As in the physical world we can conceive the theoretical systems in which dissipative forces are wholly absent, and in which the entropy consequently remains constant, so we can conceive, though we need not expect to find, biological populations in which the genetic variance is absolutely zero, and in which fitness does not increase. Professor Eddington has recently remarked that 'The law that entropy always increases—the second law of thermodynamics—holds, I think, the supreme position among the laws of nature'. It is not a little instructive that so similar a law should hold the supreme position among the biological sciences. While it is possible that both may ultimately be absorbed by some more general principle, for the present we should note that the laws as they stand present profound differences—-(1) The systems considered in thermodynamics are permanent; species on the contrary are liable to extinction, although biological improvement must be expected to occur up to the end of their existence. (2) Fitness, although measured by a uniform method, is qualitatively different for every different organism, whereas entropy, like temperature, is taken to have the same meaning for all physical systems. (3) Fitness may be increased or decreased by changes in the environment, without reacting quantitatively upon that environment. (4) Entropy changes are exceptional in the physical world in being irreversible, while irreversible evolutionary changes form no exception among biological phenomena. Finally, (5) entropy changes lead to a progressive disorganization of the physical world, at least from the human standpoint of the utilization of energy, while evolutionary changes are generally recognized as producing progressively higher organization in the organic world.

Judged superficially, a progressive saturation of the germ plasm of a species with mutant genes a majority of which are deleterious in their effects is a destructive process, a sort of deterioration of the genotype which threatens the very existence of the species and can finally lead only to its extinction. The eugenical Jeremiahs keep constantly before our eyes the nightmare of human populations accumulating recessive genes that produce pathological effects when homozygous. These prophets of doom seem to be unaware of the fact that wild species in the state of nature fare in this respect no better than man does with all the artificality of his surroundings, and yet life has not come to an end on this planet. The eschatological cries proclaiming the failure of natural selection to operate in human populations have more to do with political beliefs than with scientific findings.

Mutation is random; natural selection is the very opposite of random

Natural selection is a theory of local adaptation to changing environments. It proposes no perfecting principles, no guarantee of general improvement,

Natural selection is not the wind which propels the vessel, but the rudder which, by friction, now on this side and now on that, shapes the course.

On the theory of natural selection we can clearly understand the full meaning of that old canon in natural history, 'Natura non facit saltum.' This canon, if we look only to the present inhabitants of the world, is not strictly correct, but if we include all those of past times, it must by my theory be strictly true.

Quantitative work shows clearly that natural selection is a reality, and that, among other things, it selects Mendelian genes, which are known to be distributed at random through wild populations, and to follow the laws of chance in their distribution to offspring. In other words, they are an agency producing variation of the kind which Darwin postulated as the raw material on which selection acts.

Such biological ideas as the 'survival of the fittest,' whatever their doubtful value in natural science, are utterly useless in attempting to understand society ... The life of a man in society, while it is incidentally a biological fact, has characteristics that are not reducible to biology and must be explained in the distinctive terms of a cultural analysis ... the physical well-being of men is a result of their social organization and not vice versa ... Social improvement is a product of advances in technology and social organization, not of breeding or selective elimination ... Judgments as to the value of competition between men or enterprises or nations must be based upon social and not allegedly biological consequences; and ... there is nothing in nature or a naturalistic philosophy of life to make impossible the acceptance of moral sanctions that can be employed for the common good.

The frequent allegation that the selective processes in the human species are no longer 'natural' is due to persistence of the obsolete nineteenth- century concept of 'natural' selection. The error of this view is made clear when we ask its proponents such questions as, why should the 'surviving fittest' be able to withstand cold and inclement weather without the benefit of fire and clothing? Is it not ludicrous to expect selection to make us good at defending ourselves against wild beasts when wild beasts are getting to be so rare that it is a privilege to see one outside of a zoo? Is it necessary to eliminate everyone who has poor teeth when our dentists stand ready to provide us with artificial ones? Is it a great virtue to be able to endure pain when anaesthetics are available?

The outlook seems grim. Natural selection under civilized conditions may lead mankind to evolve towards a state of genetic overspecialization for living in gadget-ridden environments. It is certainly up to man to decide whether this direction of his evolution is or is not desirable. If it is not, man has, or soon will have, the knowledge requisite to redirect the evolution of his species pretty much as he sees fit. Perhaps we should not be too dogmatic about this choice of direction. We may be awfully soft compared to paleolithic men when it comes to struggling, unaided by gadgets, with climatic difficulties and wild beasts. Most of us feel most of the time that this is not a very great loss. If our remote descendants grow to be even more effete than we are, they may conceivably be compensated by acquiring genotypes conducive to kindlier dispositions and greater intellectual capacities than those prevalent in mankind today.

The preservation of favourable variations and the rejection of injurious variations, I call Natural Selection, or Survival of the Fittest. Variations neither useful nor injurious would not be affected by natural selection and would be left a fluctuating element.

The process of mutation is the only known source of the raw materials of genetic variability, and hence of evolution. It is subject to experimental study, and considerable progress has been accomplished in this study in recent years. An apparent paradox has been disclosed. Although the living matter becomes adapted to its environment through formation of superior genetic patterns from mutational components, the process of mutation itself is not adaptive. On the contrary, the mutants which arise are, with rare exceptions, deleterious to their carriers, at least in the environments which the species normally encounters. Some of them are deleterious apparently in all environments. Therefore, the mutation process alone, not corrected and guided by natural selection, would result in degeneration and extinction rather than in improved adaptedness.

The process of natural selection has been summed up in the phrase 'survival of the fittest'. This, however, tells only part of the story. 'Survival of the existing' in many cases covers more of the truth. For in hosts of cases the survival of characters rests not on any special usefulness or fitness, but on the fact that individuals possessing these characters have inhabited or invaded a certain area. The principle of utility explains survivals among competing structures. It rarely accounts for qualities associated with geographic distribution.
The nature of animals which first colonize a district must determine what the future fauna will be. From their specific characters, which are neither useful nor harmful, will be derived for the most part the specific characters of their successors.
It is not essential to the meadow lark that he should have a black blotch on the breast or the outer tail-feather white. Yet all meadow larks have these characters just as all shore larks have the tiny plume behind the ear. Those characters of the parent stock, which may be harmful in the new relations, will be eliminated by natural selection. Those especially helpful will be intensified and modified, but the great body of characters, the marks by which we know the species, will be neither helpful nor hurtful. These will be meaningless streaks and spots, variations in size of parts, peculiar relations of scales or hair or feathers, little matters which can neither help nor hurt, but which have all the persistence heredity can give.

The Struggle for Existence amongst all organic beings throughout the world, which inevitably follows from their high geometrical powers of increase, will be treated of. This is the doctrine of Malthus, applied to the whole animal and vegetable kingdoms. As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.

The theory of evolution by natural selection is an ecological theory—founded on ecological observation by perhaps the greatest of all ecologists. It has been adopted by and brought up by the science of genetics, and ecologists, being modest people, are apt to forget their distinguished parenthood.

To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree. Yet reason tells me, that if numerous gradations from a perfect and complex eye to one very imperfect and simple, each grade being useful to its possessor, can be shown to exist; if further, the eye does vary ever so slightly, and the variations be inherited, which is certainly the case; and if any variation or modification in the organ be ever useful to an animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, can hardly be considered real.

We are the accidental result of an unplanned process … the fragile result of an enormous concatenation of improbabilities, not the predictable product of any definite process.

We live in a cultural milieu ... The idea that culture is our ecological niche is still applicable. The impact and force of natural selection on the human physique are conditioned by the dimensions of culture.

We may consequently state the fundamental theorem of Natural Selection in the form: The rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time.

When one ponders on the tremendous journey of evolution over the past three billion years or so, the prodigious wealth of structures it has engendered, and the extraordinarily effective teleonomic performances of living beings from bacteria to man, one may well find oneself beginning to doubt again whether all this could conceiveably be the product of an enormous lottery presided over by natural selection, blindly picking the rare winners from among numbers drawn at random. [Nevertheless,] a detailed review of the accumulated modern evidence [shows] that this conception alone is compatible with the facts.