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1.【址:a g 9 559⒐ v i p】1  Next Chapter
2.  If there exist savages so barbarous as never to think of the inherited character of the offspring of their domestic animals, yet any one animal particularly useful to them, for any special purpose, would be carefully preserved during famines and other accidents, to which savages are so liable, and such choice animals would thus generally leave more offspring than the inferior ones; so that in this case there would be a kind of unconscious selection going on. We see the value set on animals even by the barbarians of Tierra del Fuego, by their killing and devouring their old women, in times of dearth, as of less value than their dogs.
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4.  Hence I look at individual differences, though of small interest to the systematist, as of high importance for us, as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. And I look at varieties which are in any degree more distinct and permanent, as steps leading to more strongly marked and more permanent varieties; and at these latter, as leading to sub-species, and to species. The passage from one stage of difference to another and higher stage may be, in some cases, due merely to the long-continued action of different physical conditions in two different regions; but I have not much faith in this view; and I attribute the passage of a variety, from a state in which it differs very slightly from its parent to one in which it differs more, to the action of natural selection in accumulating (as will hereafter be more fully explained) differences of structure in certain definite directions. Hence I believe a well-marked variety may be justly called an incipient species; but whether this belief be justifiable must be judged of by the general weight of the several facts and views given throughout this work.It need not be supposed that all varieties or incipient species necessarily attain the rank of species. They may whilst in this incipient state become extinct, or they may endure as varieties for very long periods, as has been shown to be the case by Mr Wollaston with the varieties of certain fossil land-shells in Madeira. If a variety were to flourish so as to exceed in numbers the parent species, it would then rank as the species, and the species as the variety; or it might come to supplant and exterminate the parent species; or both might co-exist, and both rank as independent species. But we shall hereafter have to return to this subject.
5.  Although I do not doubt that isolation is of considerable importance in the production of new species, on the whole I am inclined to believe that largeness of area is of more importance, more especially in the production of species, which will prove capable of enduring for a long period, and of spreading widely. Throughout a great and open area, not only will there be a better chance of favourable variations arising from the large number of individuals of the same species there supported, but the conditions of life are infinitely complex from the large number of already existing species; and if some of these many species become modified and improved, others will have to be improved in a corresponding degree or they will be exterminated. Each new form, also, as soon as it has been much improved, will be able to spread over the open and continuous area, and will thus come into competition with many others. Hence more new places will be formed, and the competition to fill them will be more severe, on a large than on a small and isolated area. Moreover, great areas, though now continuous, owing to oscillations of level, will often have recently existed in a broken condition, so that the good effects of isolation will generally, to a certain extent, have concurred. Finally, I conclude that, although small isolated areas probably have been in some respects highly favourable for the production of new species, yet that the course of modification will generally have been more rapid on large areas; and what is more important, that the new forms produced on large areas, which already have been victorious over many competitors, will be those that will spread most widely, will give rise to most new varieties and species, and will thus play an important part in the changing history of the organic world.We can, perhaps, on these views, understand some facts which will be again alluded to in our chapter on geographical distribution; for instance, that the productions of the smaller continent of Australia have formerly yielded, and apparently are now yielding, before those of the larger Europaeo-Asiatic area. Thus, also, it is that continental productions have everywhere become so largely naturalised on islands. On a small island, the race for life will have been less severe, and there will have been less modification and less extermination. Hence, perhaps, it comes that the flora of Madeira, according to Oswald Heer, resembles the extinct tertiary flora of Europe. All fresh-water basins, taken together, make a small area compared with that of the sea or of the land; and, consequently, the competition between fresh-water productions will have been less severe than elsewhere; new forms will have been more slowly formed, and old forms more slowly exterminated. And it is in fresh water that we find seven genera of Ganoid fishes, remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders now widely separated in the natural scale. These anomalous forms may almost be called living fossils; they have endured to the present day, from having inhabited a confined area, and from having thus been exposed to less severe competition.To sum up the circumstances favourable and unfavourable to natural selection, as far as the extreme intricacy of the subject permits. I conclude, looking to the future, that for terrestrial productions a large continental area, which will probably undergo many oscillations of level, and which consequently will exist for long periods in a broken condition, will be the most favourable for the production of many new forms of life, likely to endure long and to spread widely. For the area will first have existed as a continent, and the inhabitants, at this period numerous in individuals and kinds, will have been subjected to very severe competition. When converted by subsidence into large separate islands, there will still exist many individuals of the same species on each island: intercrossing on the confines of the range of each species will thus be checked: after physical changes of any kind, immigration will be prevented, so that new places in the polity of each island will have to be filled up by modifications of the old inhabitants; and time will be allowed for the varieties in each to become well modified and perfected. When, by renewed elevation, the islands shall be re-converted into a continental area, there will again be severe competition: the most favoured or improved varieties will be enabled to spread: there will be much extinction of the less improved forms, and the relative proportional numbers of the various inhabitants of the renewed continent will again be changed; and again there will be a fair field for natural selection to improve still further the inhabitants, and thus produce new species.That natural selection will always act with extreme slowness, I fully admit. Its action depends on there being places in the polity of nature, which can be better occupied by some of the inhabitants of the country undergoing modification of some kind. The existence of such places will often depend on physical changes, which are generally very slow, and on the immigration of better adapted forms having been checked. But the action of natural selection will probably still oftener depend on some of the inhabitants becoming slowly modified; the mutual relations of many of the other inhabitants being thus disturbed. Nothing can be effected, unless favourable variations occur, and variation itself is apparently always a very slow process. The process will often be greatly retarded by free intercrossing. Many will exclaim that these several causes are amply sufficient wholly to stop the action of natural selection. I do not believe so. On the other hand, I do believe that natural selection will always act very slowly, often only at long intervals of time, and generally on only a very few of the inhabitants of the same region at the same time. I further believe, that this very slow, intermittent action of natural selection accords perfectly well with what geology tells us of the rate and manner at which the inhabitants of this world have changed.Slow though the process of selection may be, if feeble man can do much by his powers of artificial selection, I can see no limit to the amount of change, to the beauty and infinite complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may be effected in the long course of time by nature's power of selection.
6.  I know of no case better adapted to show the importance of the laws of correlation in modifying important structures, independently of utility and, therefore, of natural selection, than that of the difference between the outer and inner flowers in some Compositous and Umbelliferous plants. Every one knows the difference in the ray and central florets of, for instance, the daisy, and this difference is often accompanied with the abortion of parts of the flower. But, in some Compositous plants, the seeds also differ in shape and sculpture; and even the ovary itself, with its accessory parts, differs, as has been described by Cassini. These differences have been attributed by some authors to pressure, and the shape of the seeds in the ray-florets in some Compositae countenances this idea; but, in the case of the corolla of the Umbelliferae, it is by no means, as Dr Hooker informs me, in species with the densest heads that the inner and outer flowers most frequently differ. It might have been thought that the development of the ray-petals by drawing nourishment from certain other parts of the flower had caused their abortion; but in some Compositae there is a difference in the seeds of the outer and inner florets without any difference in the corolla. Possibly, these several differences may be connected with some difference in the flow of nutriment towards the central and external flowers: we know, at least, that in irregular flowers, those nearest to the axis are oftenest subject to peloria, and become regular. I may add, as an instance of this, and of a striking case of correlation, that I have recently observed in some garden pelargoniums, that the central flower of the truss often loses the patches of darker colour in the two upper petals; and that when this occurs, the adherent nectary is quite aborted; when the colour is absent from only one of the two upper petals, the nectary is only much shortened.With respect to the difference in the corolla of the central and exterior flowers of a head or umbel, I do not feel at all sure that C. C. Sprengel's idea that the ray-florets serve to attract insects, whose agency is highly advantageous in the fertilisation of plants of these two orders, is so far-fetched, as it may at first appear: and if it be advantageous, natural selection may have come into play. But in regard to the differences both in the internal and external structure of the seeds, which are not always correlated with any differences in the flowers, it seems impossible that they can be in any way advantageous to the plant: yet in the Umbelliferae these differences are of such apparent importance the seeds being in some cases, according to Tausch, orthospermous in the exterior flowers and coelospermous in the central flowers, that the elder De Candolle founded his main divisions of the order on analogous differences. Hence we see that modifications of structure, viewed by systematists as of high value, may be wholly due to unknown laws of correlated growth, and without being, as far as we can see, of the slightest service to the species.We may often falsely attribute to correlation of growth, structures which are common to whole groups of species, and which in truth are simply due to inheritance; for an ancient progenitor may have acquired through natural selection some one modification in structure, and, after thousands of generations, some other and independent modification; and these two modifications, having been transmitted to a whole group of descendants with diverse habits, would naturally be thought to be correlated in some necessary manner. So, again, I do not doubt that some apparent correlations, occurring throughout whole orders, are entirely due to the manner alone in which natural selection can act. For instance, Alph. De Candolle has remarked that winged seeds are never found in fruits which do not open: I should explain the rule by the fact that seeds could not gradually become winged through natural selection, except in fruits which opened; so that the individual plants producing seeds which were a little better fitted to be wafted further, might get an advantage over those producing seed less fitted for dispersal; and this process could not possibly go on in fruit which did not open.The elder Geoffroy and Goethe propounded, at about the same period, their law of compensation or balancement of growth; or, as Goethe expressed it, 'in order to spend on one side, nature is forced to economise on the other side.' I think this holds true to a certain extent with our domestic productions: if nourishment flows to one part or organ in excess, it rarely flows, at least in excess, to another part; thus it is difficult to get a cow to give much milk and to fatten readily. The same varieties of the cabbage do not yield abundant and nutritious foliage and a copious supply of oil-bearing seeds. When the seeds in our fruits become atrophied, the fruit itself gains largely in size and quality. In our poultry, a large tuft of feathers on the head is generally accompanied by a diminished comb, and a large beard by diminished wattles. With species in a state of nature it can hardly be maintained that the law is of universal application; but many good observers, more especially botanists, believe in its truth. I will not, however, here give any instances, for I see hardly any way of distinguishing between the effects, on the one hand, of a part being largely developed through natural selection and another and adjoining part being reduced by this same process or by disuse, and, on the other hand, the actual withdrawal of nutriment from one part owing to the excess of growth in another and adjoining part.I suspect, also, that some of the cases of compensation which have been advanced, and likewise some other facts, may be merged under a more general principle, namely, that natural selection is continually trying to economise in every part of the organisation. If under changed conditions of life a structure before useful becomes less useful, any diminution, however slight, in its development, will be seized on by natural selection, for it will profit the individual not to have its nutriment wasted in building up an useless structure. I can thus only understand a fact with which I was much struck when examining cirripedes, and of which many other instances could be given: namely, that when a cirripede is parasitic within another and is thus protected, it loses more or less completely its own shell or carapace. This is the case with the male Ibla, and in a truly extraordinary manner with the Proteolepas: for the carapace in all other cirripedes consists of the three highly-important anterior segments of the head enormously developed, and furnished with great nerves and muscles; but in the parasitic and protected Proteolepas, the whole anterior part of the head is reduced to the merest rudiment attached to the bases of the prehensile antennae. Now the saving of a large and complex structure, when rendered superfluous by the parasitic habits of the Proteolepas, though effected by slow steps, would be a decided advantage to each successive individual of the species; for in the struggle for life to which every animal is exposed, each individual Proteolepas would have a better chance of supporting itself, by less nutriment being wasted in developing a structure now become useless.Thus, as I believe, natural selection will always succeed in the long run in reducing and saving every part of the organisation, as soon as it is rendered superfluous, without by any means causing some other part to be largely developed in a corresponding degree. And, conversely, that natural selection may perfectly well succeed in largely developing any organ, without requiring as a necessary compensation the reduction of some adjoining part.

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1.  In man's methodical selection, a breeder selects for some definite object, and free intercrossing will wholly stop his work. But when many men, without intending to alter the breed, have a nearly common standard of perfection, and all try to get and breed from the best animals, much improvement and modification surely but slowly follow from this unconscious process of selection, notwithstanding a large amount of crossing with inferior animals. Thus it will be in nature; for within a confined area, with some place in its polity not so perfectly occupied as might be, natural selection will always tend to preserve all the individuals varying in the right direction, though in different degrees, so as better to fill up the unoccupied place. But if the area be large, its several districts will almost certainly present different conditions of life; and then if natural selection be modifying and improving a species in the several districts, there will be intercrossing with the other individuals of the same species on the confines of each. And in this case the effects of intercrossing can hardly be counterbalanced by natural selection always tending to modify all the individuals in each district in exactly the same manner to the conditions of each; for in a continuous area, the conditions will generally graduate away insensibly from one district to another. The intercrossing will most affect those animals which unite for each birth, which wander much, and which do not breed at a very quick rate. Hence in animals of this nature, for instance in birds, varieties will generally be confined to separated countries; and this I believe to be the case. In hermaphrodite organisms which cross only occasionally, and likewise in animals which unite for each birth, but which wander little and which can increase at a very rapid rate, a new and improved variety might be quickly formed on any one spot, and might there maintain itself in a body, so that whatever intercrossing took place would be chiefly between the individuals of the same new variety. A local variety when once thus formed might subsequently slowly spread to other districts. On the above principle, nurserymen always prefer getting seed from a large body of plants of the same variety, as the chance of intercrossing with other varieties is thus lessened.Even in the case of slow-breeding animals, which unite for each birth, we must not overrate the effects of intercrosses in retarding natural selection; for I can bring a considerable catalogue of facts, showing that within the same area, varieties of the same animal can long remain distinct, from haunting different stations, from breeding at slightly different seasons, or from varieties of the same kind preferring to pair together.
2.  In favour of this view, I may add, firstly, that C. livia, or the rock-pigeon, has been found capable of domestication in Europe and in India; and that it agrees in habits and in a great number of points of structure with all the domestic breeds. Secondly, although an English carrier or short-faced tumbler differs immensely in certain characters from the rock-pigeon, yet by comparing the several sub-breeds of these breeds, more especially those brought from distant countries, we can make an almost perfect series between the extremes of structure. Thirdly, those characters which are mainly distinctive of each breed, for instance the wattle and length of beak of the carrier, the shortness of that of the tumbler, and the number of tail-feathers in the fantail, are in each breed eminently variable; and the explanation of this fact will be obvious when we come to treat of selection. Fourthly, pigeons have been watched, and tended with the utmost care, and loved by many people. They have been domesticated for thousands of years in several quarters of the world; the earliest known record of pigeons is in the fifth Aegyptian dynasty, about 3000 B.C., as was pointed out to me by Professor Lepsius; but Mr Birch informs me that pigeons are given in a bill of fare in the previous dynasty. In the time of the Romans, as we hear from Pliny, immense prices were given for pigeons; 'nay, they are come to this pass, that they can reckon up their pedigree and race.' Pigeons were much valued by Akber Khan in India, about the year 1600; never less than 20,000 pigeons were taken with the court. 'The monarchs of Iran and Turan sent him some very rare birds;' and, continues the courtly historian, 'His Majesty by crossing the breeds, which method was never practised before, has improved them astonishingly.' About this same period the Dutch were as eager about pigeons as were the old Romans. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone, will be obvious when we treat of Selection. We shall then, also, see how it is that the breeds so often have a somewhat monstrous character. It is also a most favourable circumstance for the production of distinct breeds, that male and female pigeons can be easily mated for life; and thus different breeds can be kept together in the same aviary.I have discussed the probable origin of domestic pigeons at some, yet quite insufficient, length; because when I first kept pigeons and watched the several kinds, knowing well how true they bred, I felt fully as much difficulty in believing that they could ever have descended from a common parent, as any naturalist could in coming to a similar conclusion in regard to the many species of finches, or other large groups of birds, in nature. One circumstance has struck me much; namely, that all the breeders of the various domestic animals and the cultivators of plants, with whom I have ever conversed, or whose treatises I have read, are firmly convinced that the several breeds to which each has attended, are descended from so many aboriginally distinct species. Ask, as I have asked, a celebrated raiser of Hereford cattle, whether his cattle might not have descended from long horns, and he will laugh you to scorn. I have never met a pigeon, or poultry, or duck, or rabbit fancier, who was not fully convinced that each main breed was descended from a distinct species. Van Mons, in his treatise on pears and apples, shows how utterly he disbelieves that the several sorts, for instance a Ribston-pippin or Codlin-apple, could ever have proceeded from the seeds of the same tree. Innumerable other examples could be given. The explanation, I think, is simple: from long-continued study they are strongly impressed with the differences between the several races; and though they well know that each race varies slightly, for they win their prizes by selecting such slight differences, yet they ignore all general arguments, and refuse to sum up in their minds slight differences accumulated during many successive generations. May not those naturalists who, knowing far less of the laws of inheritance than does the breeder, and knowing no more than he does of the intermediate links in the long lines of descent, yet admit that many of our domestic races have descended from the same parents may they not learn a lesson of caution, when they deride the idea of species in a state of nature being lineal descendants of other species?Selection
3.  It is good thus to try in our imagination to give any form some advantage over another. Probably in no single instance should we know what to do, so as to succeed. It will convince us of our ignorance on the mutual relations of all organic beings; a conviction as necessary, as it seems to be difficult to acquire. 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.
4.  But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the diagram, though in itself made somewhat irregular. I am far from thinking that the most divergent varieties will invariably prevail and multiply: a medium form may often long endure, and may or may not produce more than one modified descendant; for natural selection will always act according to the nature of the places which are either unoccupied or not perfectly occupied by other beings; and this will depend on infinitely complex relations. But as a general rule, the more diversified in structure the descendants from any one species can be rendered, the more places they will be enabled to seize on, and the more their modified progeny will be increased. In our diagram the line of succession is broken at regular intervals by small numbered letters marking the successive forms which have become sufficiently distinct to be recorded as varieties. But these breaks are imaginary, and might have been inserted anywhere, after intervals long enough to have allowed the accumulation of a considerable amount of divergent variation.As all the modified descendants from a common and widely-diffused species, belonging to a large genus, will tend to partake of the same advantages which made their parent successful in life, they will generally go on multiplying in number as well as diverging in character: this is represented in the diagram by the several divergent branches proceeding from (A). The modified offspring from the later and more highly improved branches in the lines of descent, will, it is probable, often take the place of, and so destroy, the earlier and less improved branches: this is represented in the diagram by some of the lower branches not reaching to the upper horizontal lines. In some cases I do not doubt that the process of modification will be confined to a single line of descent, and the number of the descendants will not be increased; although the amount of divergent modification may have been increased in the successive generations. This case would be represented in the diagram, if all the lines proceeding from (A) were removed, excepting that from a1 to a10 In the same way, for instance, the English race-horse and English pointer have apparently both gone on slowly diverging in character from their original stocks, without either having given off any fresh branches or races.After ten thousand generations, species (A) is supposed to have produced three forms, a10, f10, and m10, which, from having diverged in character during the successive generations, will have come to differ largely, but perhaps unequally, from each other and from their common parent. If we suppose the amount of change between each horizontal line in our diagram to be excessively small, these three forms may still be only well-marked varieties; or they may have arrived at the doubtful category of sub-species; but we have only to suppose the steps in the process of modification to be more numerous or greater in amount, to convert these three forms into well-defined species: thus the diagram illustrates the steps by which the small differences distinguishing varieties are increased into the larger differences distinguishing species. By continuing the same process for a greater number of generations (as shown in the diagram in a condensed and simplified manner), we get eight species, marked by the letters between a14 and m14, all descended from (A). Thus, as I believe, species are multiplied and genera are formed.In a large genus it is probable that more than one species would vary. In the diagram I have assumed that a second species (I) has produced, by analogous steps, after ten thousand generations, either two well-marked varieties (w10 and z10) or two species, according to the amount of change supposed to be represented between the horizontal lines. After fourteen thousand generations, six new species, marked by the letters n14 to z14, are supposed to have been produced. In each genus, the species, which are already extremely different in character, will generally tend to produce the greatest number of modified descendants; for these will have the best chance of filling new and widely different places in the polity of nature: hence in the diagram I have chosen the extreme species (A), and the nearly extreme species (I), as those which have largely varied, and have given rise to new varieties and species. The other nine species (marked by capital letters) of our original genus, may for a long period continue transmitting unaltered descendants; and this is shown in the diagram by the dotted lines not prolonged far upwards from want of space.But during the process of modification, represented in the diagram, another of our principles, namely that of extinction, will have played an important part. As in each fully stocked country natural selection necessarily acts by the selected form having some advantage in the struggle for life over other forms, there will be a constant tendency in the improved descendants of any one species to supplant and exterminate in each stage of descent their predecessors and their original parent. For it should be remembered that the competition will generally be most severe between those forms which are most nearly related to each other in habits, constitution, and structure. Hence all the intermediate forms between the earlier and later states, that is between the less and more improved state of a species, as well as the original parent-species itself, will generally tend to become extinct. So it probably will be with many whole collateral lines of descent, which will be conquered by later and improved lines of descent. If, however, the modified offspring of a species get into some distinct country, or become quickly adapted to some quite new station, in which child and parent do not come into competition, both may continue to exist.If then our diagram be assumed to represent a considerable amount of modification, species (A) and all the earlier varieties will have become extinct, having been replaced by eight new species (a14 to m14); and (I) will have been replaced by six (n14 to z14) new species.
5.  But we may go further than this. The original species of our genus were supposed to resemble each other in unequal degrees, as is so generally the case in nature; species (A) being more nearly related to B, C, and D, than to the other species; and species (I) more to G, H, K, L, than to the others. These two species (A) and (I), were also supposed to be very common and widely diffused species, so that they must originally have had some advantage over most of the other species of the genus. Their modified descendants, fourteen in number at the fourteen-thousandth generation, will probably have inherited some of the same advantages: they have also been modified and improved in a diversified manner at each stage of descent, so as to have become adapted to many related places in the natural economy of their country. It seems, therefore, to me extremely probable that they will have taken the places of, and thus exterminated, not only their parents (A) and (I), but likewise some of the original species which were most nearly related to their parents. Hence very few of the original species will have transmitted offspring to the fourteen-thousandth generation. We may suppose that only one (F), of the two species which were least closely related to the other nine original species, has transmitted descendants to this late stage of descent.The new species in our diagram descended from the original eleven species, will now be fifteen in number. Owing to the divergent tendency of natural selection, the extreme amount of difference in character between species a14 and z14 will be much greater than that between the most different of the original eleven species. The new species, moreover, will be allied to each other in a widely different manner. Of the eight descendants from (A) the three marked a14, q14, p14, will be nearly related from having recently branched off from a14; b14 and f14, from having diverged at an earlier period from a5, will be in some degree distinct from the three first-named species; and lastly, o14, e14, and m14, will be nearly related one to the other, but from having diverged at the first commencement of the process of modification, will be widely different from the other five species, and may constitute a sub-genus or even a distinct genus. The six descendants from (I) will form two sub-genera or even genera. But as the original species (I) differed largely from (A), standing nearly at the extreme points of the original genus, the six descendants from (I) will, owing to inheritance, differ considerably from the eight descendants from (A); the two groups, moreover, are supposed to have gone on diverging in different directions. The intermediate species, also (and this is a very important consideration), which connected the original species (A) and (I), have all become, excepting (F), extinct, and have left no descendants. Hence the six new species descended from (I), and the eight descended from (A), will have to be ranked as very distinct genera, or even as distinct sub-families.Thus it is, as I believe, that two or more genera are produced by descent, with modification, from two or more species of the same genus. And the two or more parent-species are supposed to have descended from some one species of an earlier genus. In our diagram, this is indicated by the broken lines, beneath the capital letters, converging in sub-branches downwards towards a single point; this point representing a single species, the supposed single parent of our several new sub-genera and genera.
6.  I mean by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood. The most obvious case is, that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded, affect the structure of the adult; in the same manner as any malconformation affecting the early embryo, seriously affects the whole organisation of the adult. The several parts of the body which are homologous, and which, at an early embryonic period, are alike, seem liable to vary in an allied manner: we see this in the right and left sides of the body varying in the same manner; in the front and hind legs, and even in the jaws and limbs, varying together, for the lower jaw is believed to be homologous with the limbs. These tendencies, I do not doubt, may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side; and if this had been of any great use to the breed it might probably have been rendered permanent by natural selection.Homologous parts, as has been remarked by some authors, tend to cohere; this is often seen in monstrous plants; and nothing is more common than the union of homologous parts in normal structures, as the union of the petals of the corolla into a tube. Hard parts seem to affect the form of adjoining soft parts; it is believed by some authors that the diversity in the shape of the pelvis in birds causes the remarkable diversity in the shape of their kidneys. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. In snakes, according to Schlegel, the shape of the body and the manner of swallowing determine the position of several of the most important viscera.

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2.  Although I do not doubt that isolation is of considerable importance in the production of new species, on the whole I am inclined to believe that largeness of area is of more importance, more especially in the production of species, which will prove capable of enduring for a long period, and of spreading widely. Throughout a great and open area, not only will there be a better chance of favourable variations arising from the large number of individuals of the same species there supported, but the conditions of life are infinitely complex from the large number of already existing species; and if some of these many species become modified and improved, others will have to be improved in a corresponding degree or they will be exterminated. Each new form, also, as soon as it has been much improved, will be able to spread over the open and continuous area, and will thus come into competition with many others. Hence more new places will be formed, and the competition to fill them will be more severe, on a large than on a small and isolated area. Moreover, great areas, though now continuous, owing to oscillations of level, will often have recently existed in a broken condition, so that the good effects of isolation will generally, to a certain extent, have concurred. Finally, I conclude that, although small isolated areas probably have been in some respects highly favourable for the production of new species, yet that the course of modification will generally have been more rapid on large areas; and what is more important, that the new forms produced on large areas, which already have been victorious over many competitors, will be those that will spread most widely, will give rise to most new varieties and species, and will thus play an important part in the changing history of the organic world.We can, perhaps, on these views, understand some facts which will be again alluded to in our chapter on geographical distribution; for instance, that the productions of the smaller continent of Australia have formerly yielded, and apparently are now yielding, before those of the larger Europaeo-Asiatic area. Thus, also, it is that continental productions have everywhere become so largely naturalised on islands. On a small island, the race for life will have been less severe, and there will have been less modification and less extermination. Hence, perhaps, it comes that the flora of Madeira, according to Oswald Heer, resembles the extinct tertiary flora of Europe. All fresh-water basins, taken together, make a small area compared with that of the sea or of the land; and, consequently, the competition between fresh-water productions will have been less severe than elsewhere; new forms will have been more slowly formed, and old forms more slowly exterminated. And it is in fresh water that we find seven genera of Ganoid fishes, remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders now widely separated in the natural scale. These anomalous forms may almost be called living fossils; they have endured to the present day, from having inhabited a confined area, and from having thus been exposed to less severe competition.To sum up the circumstances favourable and unfavourable to natural selection, as far as the extreme intricacy of the subject permits. I conclude, looking to the future, that for terrestrial productions a large continental area, which will probably undergo many oscillations of level, and which consequently will exist for long periods in a broken condition, will be the most favourable for the production of many new forms of life, likely to endure long and to spread widely. For the area will first have existed as a continent, and the inhabitants, at this period numerous in individuals and kinds, will have been subjected to very severe competition. When converted by subsidence into large separate islands, there will still exist many individuals of the same species on each island: intercrossing on the confines of the range of each species will thus be checked: after physical changes of any kind, immigration will be prevented, so that new places in the polity of each island will have to be filled up by modifications of the old inhabitants; and time will be allowed for the varieties in each to become well modified and perfected. When, by renewed elevation, the islands shall be re-converted into a continental area, there will again be severe competition: the most favoured or improved varieties will be enabled to spread: there will be much extinction of the less improved forms, and the relative proportional numbers of the various inhabitants of the renewed continent will again be changed; and again there will be a fair field for natural selection to improve still further the inhabitants, and thus produce new species.That natural selection will always act with extreme slowness, I fully admit. Its action depends on there being places in the polity of nature, which can be better occupied by some of the inhabitants of the country undergoing modification of some kind. The existence of such places will often depend on physical changes, which are generally very slow, and on the immigration of better adapted forms having been checked. But the action of natural selection will probably still oftener depend on some of the inhabitants becoming slowly modified; the mutual relations of many of the other inhabitants being thus disturbed. Nothing can be effected, unless favourable variations occur, and variation itself is apparently always a very slow process. The process will often be greatly retarded by free intercrossing. Many will exclaim that these several causes are amply sufficient wholly to stop the action of natural selection. I do not believe so. On the other hand, I do believe that natural selection will always act very slowly, often only at long intervals of time, and generally on only a very few of the inhabitants of the same region at the same time. I further believe, that this very slow, intermittent action of natural selection accords perfectly well with what geology tells us of the rate and manner at which the inhabitants of this world have changed.Slow though the process of selection may be, if feeble man can do much by his powers of artificial selection, I can see no limit to the amount of change, to the beauty and infinite complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may be effected in the long course of time by nature's power of selection.
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4.  BEF0RE entering on the subject of this chapter, I must make a few preliminary remarks, to show how the struggle for existence bears on Natural Selection. It has been seen in the last chapter that amongst organic beings in a state of nature there is some individual variability; indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life, and of one distinct organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and missletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera, and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow inevitably from the struggle for life. Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man's power of selection. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man's feeble efforts, as the works of Nature are to those of Art.We will now discuss in a little more detail the struggle for existence. In my future work this subject shall be treated, as it well deserves, at much greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult at least I have found it so than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, I am convinced that the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see, or we forget, that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings are destroyed by birds and beasts of prey; we do not always bear in mind, that though food may be now superabundant, it is not so at all seasons of each recurring year.I should premise that I use the term Struggle for Existence in a large and metaphorical sense, including dependence of one being on another, and including (which is more important) not only the life of the individual, but success in leaving progeny. Two canine animals in a time of dearth, may be truly said to struggle with each other which shall get food and live. But a plant on the edge of a desert is said to struggle for life against the drought, though more properly it should be said to be dependent on the moisture. A plant which annually produces a thousand seeds, of which on an average only one comes to maturity, may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. The missletoe is dependent on the apple and a few other trees, but can only in a far-fetched sense be said to struggle with these trees, for if too many of these parasites grow on the same tree, it will languish and die. But several seedling missletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the missletoe is disseminated by birds, its existence depends on birds; and it may metaphorically be said to struggle with other fruit-bearing plants, in order to tempt birds to devour and thus disseminate its seeds rather than those of other plants. In these several senses, which pass into each other, I use for convenience sake the general term of struggle for existence.A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season or occasional year, otherwise, on the principle of geometrical increase, its numbers would quickly become so inordinately great that no country could support the product. Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence, either one individual with another of the same species, or with the individuals of distinct species, or with the physical conditions of life. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms; for in this case there can be no artificial increase of food, and no prudential restraint from marriage. Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them.
5.   Seedlings from the same fruit, and the young of the same litter, sometimes differ considerably from each other, though both the young and the parents, as Muller has remarked, have apparently been exposed to exactly the same conditions of life; and this shows how unimportant the direct effects of the conditions of life are in comparison with the laws of reproduction, and of growth, and of inheritance; for had the action of the conditions been direct, if any of the young had varied, all would probably have varied in the same manner. To judge how much, in the case of any variation, we should attribute to the direct action of heat, moisture, light, food, &c., is most difficult: my impression is, that with animals such agencies have produced very little direct effect, though apparently more in the case of plants. Under this point of view, Mr Buckman's recent experiments on plants seem extremely valuable. When all or nearly all the individuals exposed to certain conditions are affected in the same way, the change at first appears to be directly due to such conditions; but in some cases it can be shown that quite opposite conditions produce similar changes of structure. Nevertheless some slight amount of change may, I think, be attributed to the direct action of the conditions of life as, in some cases, increased size from amount of food, colour from particular kinds of food and from light, and perhaps the thickness of fur from climate.Habit also has a deciding influence, as in the period of flowering with plants when transported from one climate to another. In animals it has a more marked effect; for instance, I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more, in proportion to the whole skeleton, than do the same bones in the wild-duck; and I presume that this change may be safely attributed to the domestic duck flying much less, and walking more, than its wild parent. The great and inherited development of the udders in cows and goats in countries where they are habitually milked, in comparison with the state of these organs in other countries, is another instance of the effect of use. Not a single domestic animal can be named which has not in some country drooping ears; and the view suggested by some authors, that the drooping is due to the disuse of the muscles of the ear, from the animals not being much alarmed by danger, seems probable.
6.  In some cases we might easily put down to disuse modifications of structure which are wholly, or mainly, due to natural selection. Mr. Wollaston has discovered the remarkable fact that 200 beetles, out of the 550 species inhabiting Madeira, are so far deficient in wings that they cannot fly; and that of the twenty-nine endemic genera, no less than twenty-three genera have all their species in this condition! Several facts, namely, that beetles in many parts of the world are very frequently blown to sea and perish; that the beetles in Madeira, as observed by Mr Wollaston, lie much concealed, until the wind lulls and the sun shines; that the proportion of wingless beetles is larger on the exposed Dezertas than in Madeira itself; and especially the extraordinary fact, so strongly insisted on by Mr. Wollaston, of the almost entire absence of certain large groups of beetles, elsewhere excessively numerous, and which groups have habits of life almost necessitating frequent flight; these several considerations have made me believe that the wingless condition of so many Madeira beetles is mainly due to the action of natural selection, but combined probably with disuse. For during thousands of successive generations each individual beetle which flew least, either from its wings having been ever so little less perfectly developed or from indolent habit, will have had the best chance of surviving from not being blown out to sea; and, on the other hand, those beetles which most readily took to flight will oftenest have been blown to sea and thus have been destroyed.The insects in Madeira which are not ground-feeders, and which, as the flower-feeding coleoptera and lepidoptera, must habitually use their wings to gain their subsistence, have, as Mr. Wollaston suspects, their wings not at all reduced, but even enlarged. This is quite compatible with the action of natural selection. For when a new insect first arrived on the island, the tendency of natural selection to enlarge or to reduce the wings, would depend on whether a greater number of individuals were saved by successfully battling with the winds, or by giving up the attempt and rarely or never flying. As with mariners ship-wrecked near a coast, it would have been better for the good swimmers if they had been able to swim still further, whereas it would have been better for the bad swimmers if they had not been able to swim at all and had stuck to the wreck.

应用

1.  Now let us turn to the effects of crossing the several species of the horse-genus. Rollin asserts, that the common mule from the ass and horse is particularly apt to have bars on its legs. I once saw a mule with its legs so much striped that any one at first would have thought that it must have been the product of a zebra; and Mr. W. C. Martin, in his excellent treatise on the horse, has given a figure of a similar mule. In four coloured drawings, which I have seen, of hybrids between the ass and zebra, the legs were much more plainly barred than the rest of the body; and in one of them there was a double shoulder-stripe. In Lord Moreton's famous hybrid from a chestnut mare and male quagga, the hybrid, and even the pure offspring subsequently produced from the mare by a black Arabian sire, were much more plainly barred across the legs than is even the pure quagga. Lastly, and this is another most remarkable case, a hybrid has been figured by Dr Gray (and he informs me that he knows of a second case) from the ass and the hemionus; and this hybrid, though the ass seldom has stripes on its legs and the hemionus has none and has not even a shoulder-stripe, nevertheless had all four legs barred, and had three short shoulder-stripes, like those on the dun Welch pony, and even had some zebra-like stripes on the sides of its face. With respect to this last fact, I was so convinced that not even a stripe of colour appears from what would commonly be called an accident, that I was led solely from the occurrence of the face-stripes on this hybrid from the ass and hemionus, to ask Colonel Poole whether such face-stripes ever occur in the eminently striped Kattywar breed of horses, and was, as we have seen, answered in the affirmative.What now are we to say to these several facts? We see several very distinct species of the horse-genus becoming, by simple variation, striped on the legs like a zebra, or striped on the shoulders like an ass. In the horse we see this tendency strong whenever a dun tint appears a tint which approaches to that of the general colouring of the other species of the genus. The appearance of the stripes is not accompanied by any change of form or by any other new character. We see this tendency to become striped most strongly displayed in hybrids from between several of the most distinct species. Now observe the case of the several breeds of pigeons: they are descended from a pigeon (including two or three sub-species or geographical races) of a bluish colour, with certain bars and other marks; and when any breed assumes by simple variation a bluish tint, these bars and other marks invariably reappear; but without any other change of form or character. When the oldest and truest breeds of various colours are crossed, we see a strong tendency for the blue tint and bars and marks to reappear in the mongrels. I have stated that the most probable hypothesis to account for the reappearance of very ancient characters, is that there is a tendency in the young of each successive generation to produce the long-lost character, and that this tendency, from unknown causes, sometimes prevails. And we have just seen that in several species of the horse-genus the stripes are either plainer or appear more commonly in the young than in the old. Call the breeds of pigeons, some of which have bred true for centuries, species; and how exactly parallel is the case with that of the species of the horse-genus! For myself, I venture confidently to look back thousands on thousands of generations, and I see an animal striped like a zebra, but perhaps otherwise very differently constructed, the common parent of our domestic horse, whether or not it be descended from one or more wild stocks, of the ass, the hemionus, quagga, and zebra.He who believes that each equine species was independently created, will, I presume, assert that each species has been created with a tendency to vary, both under nature and under domestication, in this particular manner, so as often to become striped like other species of the genus; and that each has been created with a strong tendency, when crossed with species inhabiting distant quarters of the world, to produce hybrids resembling in their stripes, not their own parents, but other species of the genus. To admit this view is, as it seems to me, to reject a real for an unreal, or at least for an unknown, cause. It makes the works of God a mere mockery and deception; I would almost as soon believe with the old and ignorant cosmogonists, that fossil shells had never lived, but had been created in stone so as to mock the shells now living on the sea-shore.
2.  Any variation which is not inherited is unimportant for us. But the number and diversity of inheritable deviations of structure, both those of slight and those of considerable physiological importance, is endless. Dr Prosper Lucas's treatise, in two large volumes, is the fullest and the best on this subject. No breeder doubts how strong is the tendency to inheritance: like produces like is his fundamental belief: doubts have been thrown on this principle by theoretical writers alone. When a deviation appears not unfrequently, and we see it in the father and child, we cannot tell whether it may not be due to the same original cause acting on both; but when amongst individuals, apparently exposed to the same conditions, any very rare deviation, due to some extraordinary combination of circumstances, appears in the parent say, once amongst several million individuals and it reappears in the child, the mere doctrine of chances almost compels us to attribute its reappearance to inheritance. Every one must have heard of cases of albinism, prickly skin, hairy bodies, &c. appearing in several members of the same family. If strange and rare deviations of structure are truly inherited, less strange and commoner deviations may be freely admitted to be inheritable. Perhaps the correct way of viewing the whole subject, would be, to look at the inheritance of every character whatever as the rule, and non-inheritance as the anomaly.The laws governing inheritance are quite unknown; no one can say why the same peculiarity in different individuals of the same species, and in individuals of different species, is sometimes inherited and sometimes not so; why the child often reverts in certain characters to its grandfather or grandmother or other much more remote ancestor; why a peculiarity is often transmitted from one sex to both sexes or to one sex alone, more commonly but not exclusively to the like sex. It is a fact of some little importance to us, that peculiarities appearing in the males of our domestic breeds are often transmitted either exclusively, or in a much greater degree, to males alone. A much more important rule, which I think may be trusted, is that, at whatever period of life a peculiarity first appears, it tends to appear in the offspring at a corresponding age, though sometimes earlier. In many cases this could not be otherwise: thus the inherited peculiarities in the horns of cattle could appear only in the offspring when nearly mature; peculiarities in the silkworm are known to appear at the corresponding caterpillar or cocoon stage. But hereditary diseases and some other facts make me believe that the rule has a wider extension, and that when there is no apparent reason why a peculiarity should appear at any particular age, yet that it does tend to appear in the offspring at the same period at which it first appeared in the parent. I believe this rule to be of the highest importance in explaining the laws of embryology. These remarks are of course confined to the first appearance of the peculiarity, and not to its primary cause, which may have acted on the ovules or male element; in nearly the same manner as in the crossed offspring from a short-horned cow by a long-horned bull, the greater length of horn, though appearing late in life, is clearly due to the male element.Having alluded to the subject of reversion, I may here refer to a statement often made by naturalists namely, that our domestic varieties, when run wild, gradually but certainly revert in character to their aboriginal stocks. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature. I have in vain endeavoured to discover on what decisive facts the above statement has so often and so boldly been made. There would be great difficulty in proving its truth: we may safely conclude that very many of the most strongly-marked domestic varieties could not possibly live in a wild state. In many cases we do not know what the aboriginal stock was, and so could not tell whether or not nearly perfect reversion had ensued. It would be quite necessary, in order to prevent the effects of intercrossing, that only a single variety should be turned loose in its new home. Nevertheless, as our varieties certainly do occasionally revert in some of their characters to ancestral forms, it seems to me not improbable, that if we could succeed in naturalising, or were to cultivate, during many generations, the several races, for instance, of the cabbage, in very poor soil (in which case, however, some effect would have to be attributed to the direct action of the poor soil), that they would to a large extent, or even wholly, revert to the wild aboriginal stock. Whether or not the experiment would succeed, is not of great importance for our line of argument; for by the experiment itself the conditions of life are changed. If it could be shown that our domestic varieties manifested a strong tendency to reversion, that is, to lose their acquired characters, whilst kept under unchanged conditions, and whilst kept in a considerable body, so that free intercrossing might check, by blending together, any slight deviations of structure, in such case, I grant that we could deduce nothing from domestic varieties in regard to species. But there is not a shadow of evidence in favour of this view: to assert that we could not breed our cart and race-horses, long and short-horned cattle and poultry of various breeds, and esculent vegetables, for an almost infinite number of generations, would be opposed to all experience. I may add, that when under nature the conditions of life do change, variations and reversions of character probably do occur; but natural selection, as will hereafter be explained, will determine how far the new characters thus arising shall be preserved.When we look to the hereditary varieties or races of our domestic animals and plants, and compare them with species closely allied together, we generally perceive in each domestic race, as already remarked, less uniformity of character than in true species. Domestic races of the same species, also, often have a somewhat monstrous character; by which I mean, that, although differing from each other, and from the other species of the same genus, in several trifling respects, they often differ in an extreme degree in some one part, both when compared one with another, and more especially when compared with all the species in nature to which they are nearest allied. With these exceptions (and with that of the perfect fertility of varieties when crossed, a subject hereafter to be discussed), domestic races of the same species differ from each other in the same manner as, only in most cases in a lesser degree than, do closely-allied species of the same genus in a state of nature. I think this must be admitted, when we find that there are hardly any domestic races, either amongst animals or plants, which have not been ranked by some competent judges as mere varieties, and by other competent judges as the descendants of aboriginally distinct species. If any marked distinction existed between domestic races and species, this source of doubt could not so perpetually recur. It has often been stated that domestic races do not differ from each other in characters of generic value. I think it could be shown that this statement is hardly correct; but naturalists differ most widely in determining what characters are of generic value; all such valuations being at present empirical. Moreover, on the view of the origin of genera which I shall presently give, we have no right to expect often to meet with generic differences in our domesticated productions.When we attempt to estimate the amount of structural difference between the domestic races of the same species, we are soon involved in doubt, from not knowing whether they have descended from one or several parent-species. This point, if could be cleared up, would be interesting; if, for instance, it could be shown that the greyhound, bloodhound, terrier, spaniel, and bull-dog, which we all know propagate their kind so truly, were the offspring of any single species, then such facts would have great weight in making us doubt about the immutability of the many very closely allied and natural species for instance, of the many foxes inhabiting different quarters of the world. I do not believe, as we shall presently see, that all our dogs have descended from any one wild species; but, in the case of some other domestic races, there is presumptive, or even strong, evidence in favour of this view.
3.  In plants the same gradual process of improvement, through the occasional preservation of the best individuals, whether or not sufficiently distinct to be ranked at their first appearance as distinct varieties, and whether or not two or more species or races have become blended together by crossing, may plainly be recognised in the increased size and beauty which we now see in the varieties of the heartsease, rose, pelargonium, dahlia, and other plants, when compared with the older varieties or with their parent-stocks. No one would ever expect to get a first-rate heartsease or dahlia from the seed of a wild plant. No one would expect to raise a first-rate melting pear from the seed of a wild pear, though he might succeed from a poor seedling growing wild, if it had come from a garden-stock. The pear, though cultivated in classical times, appears, from Pliny's description, to have been a fruit of very inferior quality. I have seen great surprise expressed in horticultural works at the wonderful skill of gardeners, in having produced such splendid results from such poor materials; but the art, I cannot doubt, has been simple, and, as far as the final result is concerned, has been followed almost unconsciously. It has consisted in always cultivating the best known variety, sowing its seeds, and, when a slightly better variety has chanced to appear, selecting it, and so onwards. But the gardeners of the classical period, who cultivated the best pear they could procure, never thought what splendid fruit we should eat; though we owe our excellent fruit, in some small degree, to their having naturally chosen and preserved the best varieties they could anywhere find.A large amount of change in our cultivated plants, thus slowly and unconsciously accumulated, explains, as I believe, the well-known fact, that in a vast number of cases we cannot recognise, and therefore do not know, the wild parent-stocks of the plants which have been longest cultivated in our flower and kitchen gardens. If it has taken centuries or thousands of years to improve or modify most of our plants up to their present standard of usefulness to man, we can understand how it is that neither Australia, the Cape of Good Hope, nor any other region inhabited by quite uncivilised man, has afforded us a single plant worth culture. It is not that these countries, so rich in species, do not by a strange chance possess the aboriginal stocks of any useful plants, but that the native plants have not been improved by continued selection up to a standard of perfection comparable with that given to the plants in countries anciently civilised.
4、  Chapter 4 - Natural Selection
5、  I know of no case better adapted to show the importance of the laws of correlation in modifying important structures, independently of utility and, therefore, of natural selection, than that of the difference between the outer and inner flowers in some Compositous and Umbelliferous plants. Every one knows the difference in the ray and central florets of, for instance, the daisy, and this difference is often accompanied with the abortion of parts of the flower. But, in some Compositous plants, the seeds also differ in shape and sculpture; and even the ovary itself, with its accessory parts, differs, as has been described by Cassini. These differences have been attributed by some authors to pressure, and the shape of the seeds in the ray-florets in some Compositae countenances this idea; but, in the case of the corolla of the Umbelliferae, it is by no means, as Dr Hooker informs me, in species with the densest heads that the inner and outer flowers most frequently differ. It might have been thought that the development of the ray-petals by drawing nourishment from certain other parts of the flower had caused their abortion; but in some Compositae there is a difference in the seeds of the outer and inner florets without any difference in the corolla. Possibly, these several differences may be connected with some difference in the flow of nutriment towards the central and external flowers: we know, at least, that in irregular flowers, those nearest to the axis are oftenest subject to peloria, and become regular. I may add, as an instance of this, and of a striking case of correlation, that I have recently observed in some garden pelargoniums, that the central flower of the truss often loses the patches of darker colour in the two upper petals; and that when this occurs, the adherent nectary is quite aborted; when the colour is absent from only one of the two upper petals, the nectary is only much shortened.With respect to the difference in the corolla of the central and exterior flowers of a head or umbel, I do not feel at all sure that C. C. Sprengel's idea that the ray-florets serve to attract insects, whose agency is highly advantageous in the fertilisation of plants of these two orders, is so far-fetched, as it may at first appear: and if it be advantageous, natural selection may have come into play. But in regard to the differences both in the internal and external structure of the seeds, which are not always correlated with any differences in the flowers, it seems impossible that they can be in any way advantageous to the plant: yet in the Umbelliferae these differences are of such apparent importance the seeds being in some cases, according to Tausch, orthospermous in the exterior flowers and coelospermous in the central flowers, that the elder De Candolle founded his main divisions of the order on analogous differences. Hence we see that modifications of structure, viewed by systematists as of high value, may be wholly due to unknown laws of correlated growth, and without being, as far as we can see, of the slightest service to the species.We may often falsely attribute to correlation of growth, structures which are common to whole groups of species, and which in truth are simply due to inheritance; for an ancient progenitor may have acquired through natural selection some one modification in structure, and, after thousands of generations, some other and independent modification; and these two modifications, having been transmitted to a whole group of descendants with diverse habits, would naturally be thought to be correlated in some necessary manner. So, again, I do not doubt that some apparent correlations, occurring throughout whole orders, are entirely due to the manner alone in which natural selection can act. For instance, Alph. De Candolle has remarked that winged seeds are never found in fruits which do not open: I should explain the rule by the fact that seeds could not gradually become winged through natural selection, except in fruits which opened; so that the individual plants producing seeds which were a little better fitted to be wafted further, might get an advantage over those producing seed less fitted for dispersal; and this process could not possibly go on in fruit which did not open.The elder Geoffroy and Goethe propounded, at about the same period, their law of compensation or balancement of growth; or, as Goethe expressed it, 'in order to spend on one side, nature is forced to economise on the other side.' I think this holds true to a certain extent with our domestic productions: if nourishment flows to one part or organ in excess, it rarely flows, at least in excess, to another part; thus it is difficult to get a cow to give much milk and to fatten readily. The same varieties of the cabbage do not yield abundant and nutritious foliage and a copious supply of oil-bearing seeds. When the seeds in our fruits become atrophied, the fruit itself gains largely in size and quality. In our poultry, a large tuft of feathers on the head is generally accompanied by a diminished comb, and a large beard by diminished wattles. With species in a state of nature it can hardly be maintained that the law is of universal application; but many good observers, more especially botanists, believe in its truth. I will not, however, here give any instances, for I see hardly any way of distinguishing between the effects, on the one hand, of a part being largely developed through natural selection and another and adjoining part being reduced by this same process or by disuse, and, on the other hand, the actual withdrawal of nutriment from one part owing to the excess of growth in another and adjoining part.I suspect, also, that some of the cases of compensation which have been advanced, and likewise some other facts, may be merged under a more general principle, namely, that natural selection is continually trying to economise in every part of the organisation. If under changed conditions of life a structure before useful becomes less useful, any diminution, however slight, in its development, will be seized on by natural selection, for it will profit the individual not to have its nutriment wasted in building up an useless structure. I can thus only understand a fact with which I was much struck when examining cirripedes, and of which many other instances could be given: namely, that when a cirripede is parasitic within another and is thus protected, it loses more or less completely its own shell or carapace. This is the case with the male Ibla, and in a truly extraordinary manner with the Proteolepas: for the carapace in all other cirripedes consists of the three highly-important anterior segments of the head enormously developed, and furnished with great nerves and muscles; but in the parasitic and protected Proteolepas, the whole anterior part of the head is reduced to the merest rudiment attached to the bases of the prehensile antennae. Now the saving of a large and complex structure, when rendered superfluous by the parasitic habits of the Proteolepas, though effected by slow steps, would be a decided advantage to each successive individual of the species; for in the struggle for life to which every animal is exposed, each individual Proteolepas would have a better chance of supporting itself, by less nutriment being wasted in developing a structure now become useless.Thus, as I believe, natural selection will always succeed in the long run in reducing and saving every part of the organisation, as soon as it is rendered superfluous, without by any means causing some other part to be largely developed in a corresponding degree. And, conversely, that natural selection may perfectly well succeed in largely developing any organ, without requiring as a necessary compensation the reduction of some adjoining part.

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  • 舒赫 08-06

      There are many laws regulating variation, some few of which can be dimly seen, and will be hereafter briefly mentioned. I will here only allude to what may be called correlation of growth. Any change in the embryo or larva will almost certainly entail changes in the mature animal. In monstrosities, the correlations between quite distinct parts are very curious; and many instances are given in Isidore Geoffroy St Hilaire's great work on this subject. Breeders believe that long limbs are almost always accompanied by an elongated head. Some instances of correlation are quite whimsical; thus cats with blue eyes are invariably deaf; colour and constitutional peculiarities go together, of which many remarkable cases could be given amongst animals and plants. From the facts collected by Heusinger, it appears that white sheep and pigs are differently affected from coloured individuals by certain vegetable poisons. Hairless dogs have imperfect teeth; long-haired and coarse-haired animals are apt to have, as is asserted, long or many horns; pigeons with feathered feet have skin between their outer toes; pigeons with short beaks have small feet, and those with long beaks large feet. Hence, if man goes on selecting, and thus augmenting, any peculiarity, he will almost certainly unconsciously modify other parts of the structure, owing to the mysterious laws of the correlation of growth.The result of the various, quite unknown, or dimly seen laws of variation is infinitely complex and diversified. It is well worth while carefully to study the several treatises published on some of our old cultivated plants, as on the hyacinth, potato, even the dahlia, &c.; and it is really surprising to note the endless points in structure and constitution in which the varieties and sub varieties differ slightly from each other. The whole organization seems to have become plastic, and tends to depart in some small degree from that of the parental type.

  • 叶可可 08-06

      In plants the same gradual process of improvement, through the occasional preservation of the best individuals, whether or not sufficiently distinct to be ranked at their first appearance as distinct varieties, and whether or not two or more species or races have become blended together by crossing, may plainly be recognised in the increased size and beauty which we now see in the varieties of the heartsease, rose, pelargonium, dahlia, and other plants, when compared with the older varieties or with their parent-stocks. No one would ever expect to get a first-rate heartsease or dahlia from the seed of a wild plant. No one would expect to raise a first-rate melting pear from the seed of a wild pear, though he might succeed from a poor seedling growing wild, if it had come from a garden-stock. The pear, though cultivated in classical times, appears, from Pliny's description, to have been a fruit of very inferior quality. I have seen great surprise expressed in horticultural works at the wonderful skill of gardeners, in having produced such splendid results from such poor materials; but the art, I cannot doubt, has been simple, and, as far as the final result is concerned, has been followed almost unconsciously. It has consisted in always cultivating the best known variety, sowing its seeds, and, when a slightly better variety has chanced to appear, selecting it, and so onwards. But the gardeners of the classical period, who cultivated the best pear they could procure, never thought what splendid fruit we should eat; though we owe our excellent fruit, in some small degree, to their having naturally chosen and preserved the best varieties they could anywhere find.A large amount of change in our cultivated plants, thus slowly and unconsciously accumulated, explains, as I believe, the well-known fact, that in a vast number of cases we cannot recognise, and therefore do not know, the wild parent-stocks of the plants which have been longest cultivated in our flower and kitchen gardens. If it has taken centuries or thousands of years to improve or modify most of our plants up to their present standard of usefulness to man, we can understand how it is that neither Australia, the Cape of Good Hope, nor any other region inhabited by quite uncivilised man, has afforded us a single plant worth culture. It is not that these countries, so rich in species, do not by a strange chance possess the aboriginal stocks of any useful plants, but that the native plants have not been improved by continued selection up to a standard of perfection comparable with that given to the plants in countries anciently civilised.

  • 比尔·默里 08-06

       When a young naturalist commences the study of a group of organisms quite unknown to him, he is at first much perplexed to determine what differences to consider as specific, and what as varieties; for he knows nothing of the amount and kind of variation to which the group is subject; and this shows, at least, how very generally there is some variation. But if he confine his attention to one class within one country, he will soon make up his mind how to rank most of the doubtful forms. His general tendency will be to make many species, for he will become impressed, just like the pigeon or poultry-fancier before alluded to, with the amount of difference in the forms which he is continually studying; and he has little general knowledge of analogical variation in other groups and in other countries, by which to correct his first impressions. As he extends the range of his observations, he will meet with more cases of difficulty; for he will encounter a greater number of closely-allied forms. But if his observations be widely extended, he will in the end generally be enabled to make up his own mind which to call varieties and which species; but he will succeed in this at the expense of admitting much variation, and the truth of this admission will often be disputed by other naturalists. When, moreover, he comes to study allied forms brought from countries not now continuous, in which case he can hardly hope to find the intermediate links between his doubtful forms, he will have to trust almost entirely to analogy, and his difficulties will rise to a climax.Certainly no clear line of demarcation has as yet been drawn between species and sub-species that is, the forms which in the opinion of some naturalists come very near to, but do not quite arrive at the rank of species; or, again, between sub-species and well-marked varieties, or between lesser varieties and individual differences. These differences blend into each other in an insensible series; and a series impresses the mind with the idea of an actual passage.

  • 阿曼达·艾宾顿 08-06

      When a species, owing to highly favourable circumstances, increases inordinately in numbers in a small tract, epidemics at least, this seems generally to occur with our game animals often ensue: and here we have a limiting check independent of the struggle for life. But even some of these so-called epidemics appear to be due to parasitic worms, which have from some cause, possibly in part through facility of diffusion amongst the crowded animals, been disproportionably favoured: and here comes in a sort of struggle between the parasite and its prey.

  • 香港·华丽秀 08-05

    {  Seedlings from the same fruit, and the young of the same litter, sometimes differ considerably from each other, though both the young and the parents, as Muller has remarked, have apparently been exposed to exactly the same conditions of life; and this shows how unimportant the direct effects of the conditions of life are in comparison with the laws of reproduction, and of growth, and of inheritance; for had the action of the conditions been direct, if any of the young had varied, all would probably have varied in the same manner. To judge how much, in the case of any variation, we should attribute to the direct action of heat, moisture, light, food, &c., is most difficult: my impression is, that with animals such agencies have produced very little direct effect, though apparently more in the case of plants. Under this point of view, Mr Buckman's recent experiments on plants seem extremely valuable. When all or nearly all the individuals exposed to certain conditions are affected in the same way, the change at first appears to be directly due to such conditions; but in some cases it can be shown that quite opposite conditions produce similar changes of structure. Nevertheless some slight amount of change may, I think, be attributed to the direct action of the conditions of life as, in some cases, increased size from amount of food, colour from particular kinds of food and from light, and perhaps the thickness of fur from climate.Habit also has a deciding influence, as in the period of flowering with plants when transported from one climate to another. In animals it has a more marked effect; for instance, I find in the domestic duck that the bones of the wing weigh less and the bones of the leg more, in proportion to the whole skeleton, than do the same bones in the wild-duck; and I presume that this change may be safely attributed to the domestic duck flying much less, and walking more, than its wild parent. The great and inherited development of the udders in cows and goats in countries where they are habitually milked, in comparison with the state of these organs in other countries, is another instance of the effect of use. Not a single domestic animal can be named which has not in some country drooping ears; and the view suggested by some authors, that the drooping is due to the disuse of the muscles of the ear, from the animals not being much alarmed by danger, seems probable.

  • 林晓雯 08-04

      The Origin of Species}

  • 王汝云 08-04

      When a variation is of the slightest use to a being, we cannot tell how much of it to attribute to the accumulative action of natural selection, and how much to the conditions of life. Thus, it is well known to furriers that animals of the same species have thicker and better fur the more severe the climate is under which they have lived; but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations, and how much to the direct action of the severe climate? for it would appear that climate has some direct action on the hair of our domestic quadrupeds.

  • 周康成 08-04

      As we see that those variations which under domestication appear at any particular period of life, tend to reappear in the offspring at the same period; for instance, in the seeds of the many varieties of our culinary and agricultural plants; in the caterpillar and cocoon stages of the varieties of the silkworm; in the eggs of poultry, and in the colour of the down of their chickens; in the horns of our sheep and cattle when nearly adult; so in a state of nature, natural selection will be enabled to act on and modify organic beings at any age, by the accumulation of profitable variations at that age, and by their inheritance at a corresponding age. If it profit a plant to have its seeds more and more widely disseminated by the wind, I can see no greater difficulty in this being effected through natural selection, than in the cotton-planter increasing and improving by selection the down in the pods on his cotton-trees. Natural selection may modify and adapt the larva of an insect to a score of contingencies, wholly different from those which concern the mature insect. These modifications will no doubt affect, through the laws of correlation, the structure of the adult; and probably in the case of those insects which live only for a few hours, and which never feed, a large part of their structure is merely the correlated result of successive changes in the structure of their larvae. So, conversely, modifications in the adult will probably often affect the structure of the larva; but in all cases natural selection will ensure that modifications consequent on other modifications at a different period of life, shall not be in the least degree injurious: for if they became so, they would cause the extinction of the species.Natural selection will modify the structure of the young in relation to the parent, and of the parent in relation to the young. In social animals it will adapt the structure of each individual for the benefit of the community; if each in consequence profits by the selected change. What natural selection cannot do, is to modify the structure of one species, without giving it any advantage, for the good of another species; and though statements to this effect may be found in works of natural history, I cannot find one case which will bear investigation. A structure used only once in an animal's whole life, if of high importance to it, might be modified to any extent by natural selection; for instance, the great jaws possessed by certain insects, and used exclusively for opening the cocoon or the hard tip to the beak of nestling birds, used for breaking the egg. It has been asserted, that of the best short-beaked tumbler-pigeons more perish in the egg than are able to get out of it; so that fanciers assist in the act of hatching. Now, if nature had to make the beak of a full-grown pigeon very short for the bird's own advantage, the process of modification would be very slow, and there would be simultaneously the most rigorous selection of the young birds within the egg, which had the most powerful and hardest beaks, for all with weak beaks would inevitably perish: or, more delicate and more easily broken shells might be selected, the thickness of the shell being known to vary like every other structure.Sexual Selection

  • 哈吉克巴尔·古珀 08-03

       As we see that those variations which under domestication appear at any particular period of life, tend to reappear in the offspring at the same period; for instance, in the seeds of the many varieties of our culinary and agricultural plants; in the caterpillar and cocoon stages of the varieties of the silkworm; in the eggs of poultry, and in the colour of the down of their chickens; in the horns of our sheep and cattle when nearly adult; so in a state of nature, natural selection will be enabled to act on and modify organic beings at any age, by the accumulation of profitable variations at that age, and by their inheritance at a corresponding age. If it profit a plant to have its seeds more and more widely disseminated by the wind, I can see no greater difficulty in this being effected through natural selection, than in the cotton-planter increasing and improving by selection the down in the pods on his cotton-trees. Natural selection may modify and adapt the larva of an insect to a score of contingencies, wholly different from those which concern the mature insect. These modifications will no doubt affect, through the laws of correlation, the structure of the adult; and probably in the case of those insects which live only for a few hours, and which never feed, a large part of their structure is merely the correlated result of successive changes in the structure of their larvae. So, conversely, modifications in the adult will probably often affect the structure of the larva; but in all cases natural selection will ensure that modifications consequent on other modifications at a different period of life, shall not be in the least degree injurious: for if they became so, they would cause the extinction of the species.Natural selection will modify the structure of the young in relation to the parent, and of the parent in relation to the young. In social animals it will adapt the structure of each individual for the benefit of the community; if each in consequence profits by the selected change. What natural selection cannot do, is to modify the structure of one species, without giving it any advantage, for the good of another species; and though statements to this effect may be found in works of natural history, I cannot find one case which will bear investigation. A structure used only once in an animal's whole life, if of high importance to it, might be modified to any extent by natural selection; for instance, the great jaws possessed by certain insects, and used exclusively for opening the cocoon or the hard tip to the beak of nestling birds, used for breaking the egg. It has been asserted, that of the best short-beaked tumbler-pigeons more perish in the egg than are able to get out of it; so that fanciers assist in the act of hatching. Now, if nature had to make the beak of a full-grown pigeon very short for the bird's own advantage, the process of modification would be very slow, and there would be simultaneously the most rigorous selection of the young birds within the egg, which had the most powerful and hardest beaks, for all with weak beaks would inevitably perish: or, more delicate and more easily broken shells might be selected, the thickness of the shell being known to vary like every other structure.Sexual Selection

  • 王莉莉 08-01

    {  I must here introduce a short digression. In the case of animals and plants with separated sexes, it is of course obvious that two individuals must always unite for each birth; but in the case of hermaphrodites this is far from obvious. Nevertheless I am strongly inclined to believe that with all hermaphrodites two individuals, either occasionally or habitually, concur for the reproduction of their kind. This view, I may add, was first suggested by Andrew Knight. We shall presently see its importance; but I must here treat the subject with extreme brevity, though I have the materials prepared for an ample discussion. All vertebrate animals, all insects, and some other large groups of animals, pair for each birth. Modern research has much diminished the number of supposed hermaphrodites, and of real hermaphrodites a large number pair; that is, two individuals regularly unite for reproduction, which is all that concerns us. But still there are many hermaphrodite animals which certainly do not habitually pair, and a vast majority of plants are hermaphrodites. What reason, it may be asked, is there for supposing in these cases that two individuals ever concur in reproduction? As it is impossible here to enter on details, I must trust to some general considerations alone.In the first place, I have collected so large a body of facts, showing, in accordance with the almost universal belief of breeders, that with animals and plants a cross between different varieties, or between individuals of the same variety but of another strain, gives vigour and fertility to the offspring; and on the other hand, that close interbreeding diminishes vigour and fertility; that these facts alone incline me to believe that it is a general law of nature (utterly ignorant though we be of the meaning of the law) that no organic being self-fertilises itself for an eternity of generations; but that a cross with another individual is occasionally perhaps at very long intervals -- indispensable.

  • 林芝第 08-01

      When a species, owing to highly favourable circumstances, increases inordinately in numbers in a small tract, epidemics at least, this seems generally to occur with our game animals often ensue: and here we have a limiting check independent of the struggle for life. But even some of these so-called epidemics appear to be due to parasitic worms, which have from some cause, possibly in part through facility of diffusion amongst the crowded animals, been disproportionably favoured: and here comes in a sort of struggle between the parasite and its prey.

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