GALLS. — Another class of facts, not relating to cultivated plants, deserves attention. I allude to the production of galls. Every one knows the curious, bright-red, hairy productions on the wild rose-tree, and the various different galls produced by the oak. Some of the latter resemble fruit, with one face as rosy as the rosiest apple. These bright colours can be of no service either to the gall-forming insect or to the tree, and probably are the direct result of the action of the light, in the same manner as the apples of Nova Scotia or Canada are brighter coloured than English apples. According to Osten Sacken’s latest revision, no less than fifty-eight kinds of galls are produced on the several species of oak, by Cynips with its sub-genera; and Mr. B.D. Walsh (23/48. See Mr. B.D. Walsh’s excellent papers in ‘Proc. Entomolog. Soc. Philadelphia’ December 1866 page 284. With respect to the willow see ibid 1864 page 546.) states that he can add many others to the list. One American species of willow, the Salix humilis, bears ten distinct kinds of galls. The leaves which spring from the galls of various English willows differ completely in shape from the natural leaves. The young shoots of junipers and firs, when punctured by certain insects, yield monstrous growths resembling flowers and fir-cones; and the flowers of some plants become from the same cause wholly changed in appearance. Galls are produced in every quarter of the world; of several sent to me by Mr. Thwaites from Ceylon, some were as symmetrical as a composite flower when in bud, others smooth and spherical like a berry; some protected by long spines, others clothed with yellow wool formed of long cellular hairs, others with regularly tufted hairs. In some galls the internal structure is simple, but in others it is highly complex; thus M. Lacaze-Duthiers (23/49. See his admirable ‘Histoire des Galles’ in ‘Annal. des Sc. Nat. Bot.’ 3rd series tome 19 1853 page 273.) has figured in the common ink-gall no less than seven concentric layers, composed of distinct tissue, namely, the epidermic, sub-epidermic, spongy, intermediate, and the hard protective layer formed of curiously thickened woody cells, and, lastly, the central mass, abounding with starch-granules on which the larvae feed.
Galls are produced by insects of various orders, but the greater number by species of Cynips. It is impossible to read M. Lacaze-Duthiers’ discussion and doubt that the poisonous secretion of the insect causes the growth of the gall; and every one knows how virulent is the poison secreted by wasps and bees, which belong to the same group with Cynips. Galls grow with extraordinary rapidity, and it is said that they attain their full size in a few days (23/50. Kirby and Spence ‘Entomology’ 1818 volume 1 page 450; Lacaze- Duthiers ibid page 284.); it is certain that they are almost completely developed before the larvae are hatched. Considering that many gall-insects are extremely small, the drop of secreted poison must be excessively minute; it probably acts on one or two cells alone, which, being abnormally stimulated, rapidly increase by a process of self-division. Galls, as Mr. Walsh (23/51. ‘Proc. Entomolog. Soc. Philadelphia’ 1864 page 558.) remarks, afford good, constant, and definite characters, each kind keeping as true to form as does any independent organic being. This fact becomes still more remarkable when we hear that, for instance, seven out of the ten different kinds of galls produced on Salix humilis are formed by gall-gnats (Cecidomyidae) which “though essentially distinct species, yet resemble one another so closely that in almost all cases it is difficult, and in most cases impossible, to distinguish the full-grown insects one from the other.” (23/52. Mr. B.D. Walsh ibid page 633 and December 1866 page 275.) For in accordance with a wide-spread analogy we may safely infer that the poison secreted by insects so closely allied would not differ much in nature; yet this slight difference is sufficient to induce widely different results. In some few cases the same species of gall-gnat produces on distinct species of willows galls which cannot be distinguished; the Cynips fecundatrix, also, has been known to produce on the Turkish oak, to which it is not properly attached, exactly the same kind of gall as on the European oak. (23/53. Mr. B.D. Walsh ibid 1864 pages 545, 411, 495; and December 1866 page 278. See also Lacaze-Duthiers.) These latter facts apparently prove that the nature of the poison is a more powerful agent in determining the form of the gall than the specific character of the tree which is acted on.
As the poisonous secretion of insects belonging to various orders has the special power of affecting the growth of various plants; as a slight difference in the nature of the poison suffices to produce widely different results; and lastly, as we know that the chemical compounds secreted by plants are eminently liable to be modified by changed conditions of life, we may believe it possible that various parts of a plant might be modified through the agency of its own altered secretions. Compare, for instance, the mossy and viscid calyx of a moss-rose, which suddenly appears through bud-variation on a Provence-rose, with the gall of red moss growing from the inoculated leaf of a wild rose, with each filament symmetrically branched like a microscopical spruce-fir, bearing a glandular tip and secreting odoriferous gummy matter. (23/54. Lacaze-Duthiers ibid pages 325, 328.) Or compare, on the one hand, the fruit of the peach, with its hairy skin, fleshy covering, hard shell and kernel, and on the other hand one of the more complex galls with its epidermic, spongy, and woody layers, surrounding tissue loaded with starch granules. These normal and abnormal structures manifestly present a certain degree of resemblance. Or, again, reflect on the cases above given of parrots which have had their plumage brightly decorated through some change in their blood, caused by having been fed on certain fishes, or locally inoculated with the poison of a toad. I am far from wishing to maintain that the moss-rose or the hard shell of the peach-stone or the bright colours of birds are actually due to any chemical change in the sap or blood; but these cases of galls and of parrots are excellently adapted to show us how powerfully and singularly external agencies may affect structure. With such facts before us, we need feel no surprise at the appearance of any modification in any organic being.
I may, also, here allude to the remarkable effects which parasitic fungi sometimes produce on plants. Reissek (23/55. ‘Linnaea’ volume 17 1843; quoted by Dr. M.T. Masters, Royal Institution, March 16, 1860.) has described a Thesium, affected by an Oecidium, which was greatly modified, and assumed some of the characteristic features of certain allied species, or even genera. Suppose, says Reissek, “the condition originally caused by the fungus to become constant in the course of time, the plant would, if found growing wild, be considered as a distinct species or even as belonging to a new genus.” I quote this remark to show how profoundly, yet in how natural a manner, this plant must have been modified by the parasitic fungus. Mr. Meehan (23/56. ‘Proc. Acad. Nat. Sc., Philadelphia’ June 16, 1874 and July 23, 1875.) also states that three species of Euphorbia and Portulaca olereacea, which naturally grow prostrate, become erect when they are attacked by the Oecidium. Euphorbia maculata in this case also becomes nodose, with the branchlets comparatively smooth and the leaves modified in shape, approaching in these respects to a distinct species, namely, the E. hypericifolia.
FACTS AND CONSIDERATIONS OPPOSED TO THE BELIEF THAT THE CONDITIONS OF LIFE ACT IN A POTENT MANNER IN CAUSING DEFINITE MODIFICATIONS OF STRUCTURE.
I have alluded to the slight differences in species naturally living in distinct countries under different conditions; and such differences we feel at first inclined to attribute, probably often with justice, to the definite action of the surrounding conditions. But it must be borne in mind that there exist many animals and plants which range widely and have been exposed to great diversities of climate, yet remain uniform in character. Some authors, as previously remarked, account for the varieties of our culinary and agricultural plants by the definite action of the conditions to which they have been exposed in the different parts of Great Britain; but there are about 200 plants (23/57. Hewett C. Watson ‘Cybele Britannica’ volume 1 1847 page 11.) which are found in every single English county; and these plants must have been exposed for an immense period to considerable differences of climate and soil, yet do not differ. So, again,, some animals and plants range over a large portion of the world, yet retain the same character.
Notwithstanding the facts previously given on the occurrence of highly peculiar local diseases and on the strange modifications of structure in plants caused by the inoculated poison of insects, and other analogous cases; still there are a multitude of variations–such as the modified skull of the niata ox and bulldog, the long horns of Caffre cattle, the conjoined toes of the solid-hoofed swine, the immense crest and protuberant skull of Polish fowls, the crop of the pouter-pigeon, and a host of other such cases–which we can hardly attribute to the definite action, in the sense before specified, of the external conditions of life. No doubt in every case there must have been some exciting cause; but as we see innumerable individuals exposed to nearly the same conditions, and one alone is affected, we may conclude that the constitution of the individual is of far higher importance than the conditions to which it has been exposed. It seems, indeed, to be a general rule that conspicuous variations occur rarely, and in one individual alone out of millions, though all may have been exposed, as far as we can judge, to nearly the same conditions. As the most strongly marked variations graduate insensibly into the most trifling, we are led by the same train of thought to attribute each slight variation much more to innate differences of constitution, however caused, than to the definite action of the surrounding conditions.
We are led to the same conclusion by considering the cases, formerly alluded to, of fowls and pigeons, which have varied and will no doubt go on varying in directly opposite ways, though kept during many generations under nearly the same conditions. Some, for instance, are born with their beaks, wings, tails, legs, etc., a little longer, and others with these same parts a little shorter. By the long-continued selection of such slight individual differences which occur in birds kept in the same aviary, widely different races could certainly be formed; and long-continued selection, important as is the result, does nothing but preserve the variations which arise, as it appears to us, spontaneously.
In these cases we see that domesticated animals vary in an indefinite number of particulars, though treated as uniformly as is possible. On the other hand, there are instances of animals and plants, which, though they have been exposed to very different conditions, both under nature and domestication, have varied in nearly the same manner. Mr. Layard informs me that he has observed amongst the Caffres of South Africa a dog singularly like an arctic Esquimaux dog. Pigeons in India present nearly the same wide diversities of colour as in Europe; and I have seen chequered and simply barred pigeons, and pigeons with blue and white loins, from Sierra Leone, Madeira, England, and India. New varieties of flowers are continually raised in different parts of Great Britain, but many of these are found by the judges at our exhibitions to be almost identical with old varieties. A vast number of new fruit-trees and culinary vegetables have been produced in North America: these differ from European varieties in the same general manner as the several varieties raised in Europe differ from one another; and no one has ever pretended that the climate of America has given to the many American varieties any general character by which they can be recognised. Nevertheless, from the facts previously advanced on the authority of Mr. Meehan with respect to American and European forest-trees it would be rash to affirm that varieties raised in the two countries would not in the course of ages assume a distinctive character. Dr. M. Masters has recorded a striking fact (23/58. ‘Gardener’s Chronicle’ 1857 page 629.) bearing on this subject: he raised numerous plants of Hybiscus syriacus from seed collected in South Carolina and the Holy Land, where the parent-plants must have been exposed to considerably different conditions; yet the seedlings from both localities broke into two similar strains, one with obtuse leaves and purple or crimson flowers, and the other with elongated leaves and more or less pink flowers.
We may, also, infer the prepotent influence of the constitution of the organism over the definite action of the conditions of life, from the several cases given in the earlier chapters of parallel series of varieties,–an important subject, hereafter to be more fully discussed. Sub-varieties of the several kinds of wheat, gourds, peaches, and other plants, and to a limited extent sub-varieties of the fowl, pigeon, and dog, have been shown either to resemble or to differ from one another in a closely corresponding or parallel manner. In other cases, a variety of one species resembles a distinct species; or the varieties of two distinct species resemble one another. Although these parallel resemblances no doubt often result from reversion to the former characters of a common progenitor; yet in other cases, when new characters first appear, the resemblance must be attributed to the inheritance of a similar constitution, and consequently to a tendency to vary in the same manner. We see something of a similar kind in the same monstrosity appearing and reappearing many times in the same species of animal, and, as Dr. Maxwell Masters has remarked to me, in the same species of plant.
We may at least conclude, that the amount of modification which animals and plants have undergone under domestication does not correspond with the degree to which they have been subjected to changed circumstances. As we know the parentage of domesticated birds far better than of most quadrupeds, we will glance through the list. The pigeon has varied in Europe more than almost any other bird; yet it is a native species, and has not been exposed to any extraordinary change of conditions. The fowl has varied equally, or almost equally, with the pigeon, and is a native of the hot jungles of India. Neither the peacock, a native of the same country, nor the guinea-fowl, an inhabitant of the dry deserts of Africa, has varied at all, or only in colour. The turkey, from Mexico, has varied but little. The duck, on the other hand, a native of Europe, has yielded some well-marked races; and as this is an aquatic bird, it must have been subjected to a far more serious change in its habits than the pigeon or even the fowl, which nevertheless have varied in a much higher degree. The goose, a native of Europe and aquatic like the duck, has varied less than any other domesticated bird, except the peacock.
Bud-variation is, also, important under our present point of view, in some few cases, as when all the eyes on the same tuber of the potato, or all the fruit on the same plum-tree, or all the flowers on the same plant, have suddenly varied in the same manner, it might be argued that the variation had been definitely caused by some change in the conditions to which the plants had been exposed; yet, in other cases, such an admission is extremely difficult. As new characters sometimes appear by bud-variation, which do not occur in the parent-species or in any allied species, we may reject, at least in these cases, the idea that they are due to reversion. Now it is well worth while to reflect maturely on some striking case of bud-variation, for instance that of the peach. This tree has been cultivated by the million in various parts of the world, has been treated differently, grown on its own roots and grafted on various stocks, planted as a standard, trained against a wall, or under glass; yet each bud of each sub-variety keeps true to its kind. But occasionally, at long intervals of time, a tree in England, or under the widely different climate of Virginia, produces a single bud, and this yields a branch which ever afterwards bears nectarines. Nectarines differ, as every one knows, from peaches in their smoothness, size, and flavour; and the difference is so great that some botanists have maintained that they are specifically distinct. So permanent are the characters thus suddenly acquired, that a nectarine produced by bud-variation has propagated itself by seed. To guard against the supposition that there is some fundamental distinction between bud and seminal variation, it is well to bear in mind that nectarines have likewise been produced from the stone of the peach; and, reversely, peaches from the stone of the nectarine. Now is it possible to conceive external conditions more closely alike than those to which the buds on the same tree are exposed? Yet one bud alone, out of the many thousands borne by the same tree, has suddenly, without any apparent cause, produced a nectarine. But the case is even stronger than this, for the same flower-bud has yielded a fruit, one-half or one-quarter a nectarine, and the other half or three-quarters a peach. Again, seven or eight varieties of the peach have yielded by bud-variation nectarines: the nectarines thus produced, no doubt, differ a little from one another; but still they are nectarines. Of course there must be some cause, internal or external, to excite the peach-bud to change its nature; but I cannot imagine a class of facts better adapted to force on our minds the conviction that what we call the external conditions of life are in many cases quite insignificant in relation to any particular variation, in comparison with the organisation or constitution of the being which varies.
It is known from the labours of Geoffroy Saint-Hilaire, and recently from those of Dareste and others, that eggs of the fowl, if shaken, placed upright, perforated, covered in part with varnish, etc., produce monstrous chickens. Now these monstrosities may be said to be directly caused by such unnatural conditions, but the modifications thus induced are not of a definite nature. An excellent observer, M. Camille Dareste (23/59. ‘Memoire sur la Production Artificielle des Monstruosites’ 1862 pages 8-12; ‘Recherches sur les Conditions, etc., chez les Monstres’ 1863 page 6. An abstract is given of Geoffroy’s Experiments by his son, in his ‘Vie, Travaux’ etc. 1847 page 290.), remarks “that the various species of monstrosities are not determined by specific causes; the external agencies which modify the development of the embryo act solely in causing a perturbation–a perversion in the normal course of development.” He compares the result to what we see in illness: a sudden chill, for instance, affects one individual alone out of many, causing either a cold, or sore-throat, rheumatism, or inflammation of the lungs or pleura. Contagious matter acts in an analogous manner. (23/60. Paget ‘Lectures on Surgical Pathology’ 1853 volume 1 page 483.) We may take a still more specific instance: seven pigeons were struck by rattle-snakes (23/61. ‘Researches upon the Venom of the Rattle-snake’ January 1861 by Dr. Mitchell page 67.): some suffered from convulsions; some had their blood coagulated, in others it was perfectly fluid; some showed ecchymosed spots on the heart, others on the intestines, etc.; others again showed no visible lesion in any organ. It is well known that excess in drinking causes different diseases in different men; but in the tropics the effects of intemperance differ from those caused in a cold climate (23/62. Mr. Sedgwick ‘British and Foreign Medico-Chirurg. Review’ July 1863 page 175.); and in this case we see the definite influence of opposite conditions. The foregoing facts apparently give us as good an idea as we are likely for a long time to obtain, how in many cases external conditions act directly, though not definitely, in causing modifications of structure.
SUMMARY. — There can be no doubt, from the facts given in this chapter, that extremely slight changes in the conditions of life sometimes, probably often, act in a definite manner on our domesticated productions; and, as the action of changed conditions in causing indefinite variability is accumulative, so it may be with their definite action. Hence considerable and definite modifications of structure probably follow from altered conditions acting during a long series of generations. In some few instances a marked effect has been produced quickly on all, or nearly all, the individuals which have been exposed to a marked change of climate, food, or other circumstance. This has occurred with European men in the United States, with European dogs in India, with horses in the Falkland Islands, apparently with various animals at Angora, with foreign oysters in the Mediterranean, and with maize transported from one climate to another. We have seen that the chemical compounds of some plants and the state of their tissues are readily affected by changed conditions. A relation apparently exists between certain characters and certain conditions, so that if the latter be changed the character is lost–as with the colours of flowers, the state of some culinary plants, the fruit of the melon, the tail of fat-tailed sheep, and the peculiar fleeces of other sheep.
The production of galls, and the change of plumage in parrots when fed on peculiar food or when inoculated by the poison of a toad, prove to us what great and mysterious changes in structure and colour, may be the definite result of chemical changes in the nutrient fluids or tissues.
We now almost certainly know that organic beings in a state of nature may be modified in various definite ways by the conditions to which they have been long exposed, as in the case of the birds and other animals in the northern and southern United States, and of American trees in comparison with their representatives in Europe. But in many cases it is most difficult to distinguish between the definite result of changed conditions, and the accumulation through natural selection of indefinite variations which have proved serviceable. If it profited a plant to inhabit a humid instead of an arid station, a fitting change in its constitution might possibly result from the direct action of the environment, though we have no grounds for believing that variations of the right kind would occur more frequently with plants inhabiting a station a little more humid than usual, than with other plants. Whether the station was unusually dry or humid, variations adapting the plant in a slight degree for directly opposite habits of life would occasionally arise, as we have good reason to believe from what we actually see in other cases.
The organisation or constitution of the being which is acted on, is generally a much more important element than the nature of the changed conditions, in determining the nature of the variation. We have evidence of this in the appearance of nearly similar modifications under different conditions, and of different modifications under apparently nearly the same conditions. We have still better evidence of this in closely parallel varieties being frequently produced from distinct races, or even distinct species; and in the frequent recurrence of the same monstrosity in the same species. We have also seen that the degree to which domesticated birds have varied, does not stand in any close relation with the amount of change to which they have been subjected.
To recur once again to bud-variations. When we reflect on the millions of buds which many trees have produced, before some one bud has varied, we are lost in wonder as to what the precise cause of each variation can be. Let us recall the case given by Andrew Knight of the forty-year-old tree of the yellow magnum bonum plum, an old variety which has been propagated by grafts on various stocks for a very long period throughout Europe and North America, and on which a single bud suddenly produced the red magnum bonum. We should also bear in mind that distinct varieties, and even distinct species,–as in the case of peaches, nectarines, and apricots,–of certain roses and camellias,– although separated by a vast number of generations from any progenitor in common, and although cultivated under diversified conditions, have yielded by bud-variation closely analogous varieties. When we reflect on these facts we become deeply impressed with the conviction that in such cases the nature of the variation depends but little on the conditions to which the plant has been exposed, and not in any especial manner on its individual character, but much more on the inherited nature or constitution of the whole group of allied beings to which the plant in question belongs. We are thus driven to conclude that in most cases the conditions of life play a subordinate part in causing any particular modification; like that which a spark plays, when a mass of combustibles bursts into flame–the nature of the flame depending on the combustible matter, and not on the spark. (23/63. Professor Weismann argues strongly in favour of this view in his ‘Saison-Dimorphismus der Schmetterlinge’ 1875 pages 40-43.)
No doubt each slight variation must have its efficient cause; but it is as hopeless an attempt to discover the cause of each, as to say why a chill or a poison affects one man differently from another. Even with modifications resulting from the definite action of the conditions of life, when all or nearly all the individuals, which have been similarly exposed, are similarly affected, we can rarely see the precise relation between cause and effect. In the next chapter it will be shown that the increased use or disuse of various organs produces an inherited effect. It will further be seen that certain variations are bound together by correlation as well as by other laws. Beyond this we cannot at present explain either the causes or nature of the variability of organic beings.
CHAPTER 2. XXIV. LAWS OF VARIATION — USE AND DISUSE, ETC.
In this and the two following chapters I shall discuss, as well as the difficulty of the subject permits, the several laws which govern Variability. These may be grouped under the effects of use and disuse, including changed habits and acclimatisation–arrest of development–correlated variation–the cohesion of homologous parts-the variability of multiple parts–compensation of growth–the position of buds with respect to the axis of the plant–and lastly, analogous variation. These several subjects so graduate into one another that their distinction is often arbitrary.
It may be convenient first briefly to discuss that coordinating and reparative power which is common, in a higher or lower degree, to all organic beings, and which was formerly designated by physiologists as nisus formativus.
Blumenbach and others (24/1. ‘An Essay on Generation’ English translation page 18; Paget ‘Lectures on Surgical Pathology’ 1853 volume 1 page 209.) have insisted that the principle which permits a Hydra, when cut into fragments, to develop itself into two or more perfect animals, is the same with that which causes a wound in the higher animals to heal by a cicatrice. Such cases as that of the Hydra are evidently analogous to the spontaneous division or fissiparous generation of the lowest animals, and likewise to the budding of plants. Between these extreme cases and that of a mere cicatrice we have every gradation. Spallanzani (24/2. ‘An Essay on Animal Reproduction’ English translation 1769 page 79.) by cutting off the legs and tail of a Salamander, got in the course of three months six crops of these members; so that 687 perfect bones were reproduced by one animal during one season. At whatever point the limb was cut off, the deficient part, and no more, was exactly reproduced. When a diseased bone has been removed, a new one sometimes “gradually assumes the regular form, and all the attachments of muscles, ligaments, etc., become as complete as before.” (24/3. Carpenter ‘Principles of Comp. Physiology’ 1854 page 479.)
This power of regrowth does not, however, always act perfectly; the reproduced tail of a lizard differs in the form of the scales from the normal tail: with certain Orthopterous insects the large hind legs are reproduced of smaller size (24/4. Charlesworth ‘Mag. of Nat. Hist.’ volume 1 1837 page 145.): the white cicatrice which in the higher animals unites the edges of a deep wound is not formed of perfect skin, for elastic tissue is not produced till long afterwards. (24/5. Paget ‘Lectures on Surgical Pathology’ volume 1 page 239.) “The activity of the nisus formativus,” says Blumenbach, “is in an inverse ratio to the age of the organised body.” Its power is also greater with animals, the lower they stand in the scale of organisation; and animals low in the scale correspond with the embryos of higher animals belonging to the same class. Newport’s observations (24/6. Quoted by Carpenter ‘Comp. Phys.’ page 479.) afford a good illustration of this fact, for he found that “myriapods, whose highest development scarcely carries them beyond the larva of perfect insects, can regenerate limbs and antennae up to the time of their last moult;” and so can the larvae of true insects, but, except in one order, not in the mature insect. Salamanders correspond in development with the tadpoles or larvae of the tailless Batrachians, and both possess to a large extent the power of regrowth; but not so the mature tailless Batrachians.
Absorption often plays an important part in the repair of injuries. When a bone is broken and does not unite, the ends are absorbed and rounded, so that a false joint is formed; or if the ends unite, but overlap, the projecting parts are removed. (24/7. Prof. Marey’s discussion on the power of co- adaptation in all parts of the organisation is excellent. ‘La Machine Animale’ 1873 chapter 9. See also Paget ‘Lectures’ etc. page 257.) A dislocated bone will form for itself a new socket. Displaced tendons and varicose veins excavate new channels in the bones against which they press. But absorption comes into action, as Virchow remarks, during the normal growth of bones; parts which are solid during youth become hollowed out for the medullary tissue as the bone increases in size. In trying to understand the many well- adapted cases of regrowth when aided by absorption, we should remember that almost all parts of the organisation, even whilst retaining the same form, undergo constant renewal; so that a part which is not renewed would be liable to absorption.
Some cases, usually classed under the so-called nisus formativus, at first appear to come under a distinct head; for not only are old structures reproduced, but new structures are formed. Thus, after inflammation “false membranes,” furnished with blood-vessels, lymphatics, and nerves, are developed; or a foetus escapes from the Fallopian tubes, and falls into the abdomen, “nature pours out a quantity of plastic lymph, which forms itself into organised membrane, richly supplied with blood-vessels,” and the foetus is nourished for a time. In certain cases of hydrocephalus the open and dangerous spaces in the skull are filled up with new bones, which interlock by perfect serrated sutures. (24/8. These cases are given by Blumenbach in his ‘Essay on Generation’ pages 52, 54.) But most physiologists, especially on the Continent, have now given up the belief in plastic lymph or blastema, and Virchow (24/9. ‘Cellular Pathology’ translation by Dr. Chance 1860 pages 27, 441.) maintains that every structure, new or old, is formed by the proliferation of pre-existing cells. On this view false membranes, like cancerous or other tumours, are merely abnormal developments of normal growths; and we can thus understand how it is that they resemble adjoining structures; for instance, that a “false membrane in the serous cavities acquires a covering of epithelium exactly like that which covers the original serous membrane; adhesions of the iris may become black apparently from the production of pigment-cells like those of the uvea.” (24/10. Paget ‘Lectures on Pathology’ volume 1 1853 page 357.)
No doubt the power of reparation, though not always perfect, is an admirable provision, ready for various emergencies, even for such as occur only at long intervals of time. (24/11. Paget ibid page 150.) Yet this power is not more wonderful than the growth and development of every single creature, more especially of those which are propagated by fissiparous generation. This subject has been here noticed, because we may infer that, when any part or organ is either greatly increased in size or wholly suppressed through variation and continued selection, the co-ordinating power of the organisation will continually tend to bring again all the parts into harmony with one another.
ON THE EFFECTS OF THE INCREASED USE AND DISUSE OF ORGANS.
It is notorious, and we shall immediately adduce proofs, that increased use or action strengthens muscles, glands, sense-organs, etc.; and that disuse, on the other hand, weakens them. It has been experimentally proved by Ranke (24/12. ‘Die Blutvertheilung, etc. der Organe’ 1871 as quoted by Jaeger ‘In Sachen Darwin’s’ 1874 page 48. See also H. Spencer ‘The Principles of Biology’ volume 2 1866 chapters 3-5.) that the flow of blood is greatly increased towards any part which is performing work, and sinks again when the part is at rest. Consequently, if the work is frequent, the vessels increase in size and the part is better nourished. Paget (24/13. ‘Lectures on Pathology’ 1853 volume 1 page 71.) also accounts for the long, thick, dark-coloured hairs which occasionally grow, even in young children, near old-standing inflamed surfaces or fractured bones by an increased flow of blood to the part. When Hunter inserted the spur of a cock into the comb, which is well supplied with blood-vessels, it grew in one case spirally to a length of six inches, and in another case forward, like a horn, so that the bird could not touch the ground with its beak. According to the interesting observations of M. Sedillot (24/14. ‘Comptes Rendus’ September 26, 1864 page 539.), when a portion of one of the bones of the leg of an animal is removed, the associated bone enlarges till it attains a bulk equal to that of the two bones, of which it has to perform the functions. This is best exhibited in dogs in which the tibia has been removed; the companion bone, which is naturally almost filiform and not one-fifth the size of the other, soon acquires a size equal to or greater than that of the tibia. Now, it is at first difficult to believe that increased weight acting on a straight bone could, by alternately increasing and diminishing the pressure, cause the blood to flow more freely in the vessels which permeate the periosteum and thus supply more nutriment to the bone. Nevertheless the observations adduced by Mr. Spencer (24/15. H. Spencer ‘The Principles of Biology’ volume 2 page 243.), on the strengthening of the bowed bones of rickety children, along their concave sides, leads to the belief that this is possible.
The rocking of the stem of a tree increases in a marked manner the growth of the woody tissue in the parts which are strained. Prof. Sachs believes, from reasons which he assigns, that this is due to the pressure of the bark being relaxed in such parts, and not as Knight and H. Spencer maintain, to an increased flow of sap caused by the movement of the trunk. (24/16. Ibid volume 2 page 269. Sachs ‘Text-book of Botany’ 1875 page 734.) But hard woody tissue may be developed without the aid of any movement, as we see with ivy closely attached to an old wall. In all such cases, it is very difficult to distinguish between the effects of long-continued selection and those which follow from the increased action of the part, or directly from some other cause. Mr. H. Spencer (24/17. Ibid volume 2 page 273.) acknowledges this difficulty, and gives as an instance the thorns on trees and the shells of nuts. Here we have extremely hard woody tissue without the possibility of any movement, and without, as far as we can see, any other directly exciting cause; and as the hardness of these parts is of manifest service to the plant, we may look at the result as probably due to the selection of so-called spontaneous variations. Every one knows that hard work thickens the epidermis on the hands; and when we hear that with infants, long before birth, the epidermis is thicker on the palms and soles of the feet than on any other part of the body, as was observed with admiration by Albinus (24/18. Paget ‘Lectures on Pathology’ volume 2 page 209.), we are naturally inclined to attribute this to the inherited effects of long-continued use or pressure. We are tempted to extend the same view even to the hoofs of quadrupeds; but who will pretend to determine how far natural selection may have aided in the formation of structures of such obvious importance to the animal?
That use strengthens the muscles may be seen in the limbs of artisans who follow different trades; and when a muscle is strengthened, the tendons, and the crests of bone to which they are attached, become enlarged; and this must likewise be the case with the blood-vessels and nerves. On the other hand, when a limb is not used, as by Eastern fanatics, or when the nerve supplying it with nervous power is effectually destroyed, the muscles wither. So again, when the eye is destroyed the optic nerve becomes atrophied, sometimes even in the course of a few months. (24/19. Muller ‘Phys.’ English translation pages 54, 791. Prof. Reed has given (‘Physiological and Anat. Researches’ page 10) a curious account of the atrophy of the limbs of rabbits after the destruction of the nerve.) The Proteus is furnished with branchiae as well as with lungs: and Schreibers (24/20. Quoted by Lecoq in ‘Geograph. Bot.’ tome 1 1854 page 182.) found that when the animal was compelled to live in deep water, the branchiae were developed to thrice their ordinary size, and the lungs were partially atrophied. When, on the other hand, the animal was compelled to live in shallow water, the lungs became larger and more vascular, whilst the branchiae disappeared in a more or less complete degree. Such modifications as these are, however, of comparatively little value for us, as we do not actually know that they tend to be inherited.
In many cases there is reason to believe that the lessened use of various organs has affected the corresponding parts in the offspring. But there is no good evidence that this ever follows in the course of a single generation. It appears, as in the case of general or indefinite variability, that several generations must be subjected to changed habits for any appreciable result. Our domestic fowls, ducks, and geese have almost lost, not only in the individual but in the race, their power of flight; for we do not see a young fowl, when frightened, take flight like a young pheasant. Hence I was led carefully to compare the limb-bones of fowls, ducks, pigeons, and rabbits, with the same bones in the wild parent-species. As the measurements and weights were fully given in the earlier chapters I need here only recapitulate the results. With domestic pigeons, the length of the sternum, the prominence of its crest, the length of the scapulae and furculum, the length of the wings as measured from tip to tip of the radii, are all reduced relatively to the same parts in the wild pigeon. The wing and tail feathers, however, are increased in length, but this may have as little connection with the use of the wings or tail, as the lengthened hair on a dog with the amount of exercise which it has habitually taken. The feet of pigeons, except in the long-beaked races, are reduced in size. With fowls the crest of the sternum is less prominent, and is often distorted or monstrous; the wing-bones have become lighter relatively to the leg-bones, and are apparently a little shorter in comparison with those of the parent-form, the Gallus bankiva. With ducks, the crest of the sternum is affected in the same manner as in the foregoing cases: the furculum, coracoids, and scapulae are all reduced in weight relatively to the whole skeleton: the bones of the wings are shorter and lighter, and the bones of the legs longer and heavier, relatively to each other, and relatively to the whole skeleton, in comparison with the same bones in the wild-duck. The decreased weight and size of the bones, in the foregoing cases, is probably the indirect result of the reaction of the weakened muscles on the bones. I failed to compare the feathers of the wings of the tame and wild duck; but Gloger (24/21. ‘Das Abandern der Vogel’ 1833 s. 74.) asserts that in the wild duck the tips of the wing-feathers reach almost to the end of the tail, whilst in the domestic duck they often hardly reach to its base. He remarks also on the greater thickness of the legs, and says that the swimming membrane between the toes is reduced; but I was not able to detect this latter difference.
With the domesticated rabbit the body, together with the whole skeleton, is generally larger and heavier than in the wild animal, and the leg-bones are heavier in due proportion; but whatever standard of comparison be taken, neither the leg-bones nor the scapulae have increased in length proportionally with the increased dimensions of the rest of the skeleton. The skull has become in a marked manner narrower, and, from the measurements of its capacity formerly given, we may conclude, that this narrowness results from the decreased size of the brain, consequent on the mentally inactive life led by these closely-confined animals.
We have seen in the eighth chapter that silk-moths, which have been kept during many centuries closely confined, emerge from their cocoons with their wings distorted, incapable of flight, often greatly reduced in size, or even, according to Quatrefages, quite rudimentary. This condition of the wings may be largely owing to the same kind of monstrosity which often affects wild Lepidoptera when artificially reared from the cocoon; or it may be in part due to an inherent tendency, which is common to the females of many Bombycidae, to have their wings in a more or less rudimentary state; but part of the effect may be attributed to long-continued disuse.
From the foregoing facts there can be no doubt that with our anciently domesticated animals, certain bones have increased or decreased in size and weight owing to increased or decreased use; but they have not been modified, as shown in the earlier chapters, in shape or structure. With animals living a free life and occasionally exposed to severe competition the reduction would tend to be greater, as it would be an advantage to them to have the development of every superfluous part saved. With highly-fed domesticated animals, on the other hand, there seems to be no economy of growth, nor any tendency to the elimination of superfluous details. But to this subject I shall recur.
Turning now to more general observations, Nathusius has shown that with the improved races of the pig, the shortened legs and snout, the form of the articular condyles of the occiput, and the position of the jaws with the upper canine teeth projecting in a most anomalous manner in front of the lower canines, may be attributed to these parts not having been fully exercised. For the highly-cultivated races do not travel in search of food, nor root up the ground with their ringed muzzles. (24/22. Nathusius ‘Die Racen des Schweines’ 1860 s. 53, 57; ‘Vorstudien…Schweineschadel’ 1864 s. 103, 130, 133. Prof. Lucae supports and extends the conclusions of Von Nathusius: ‘Der Schadel des Maskenschweines’ 1870.) These modifications of structure, which are all strictly inherited, characterise several improved breeds, so that they cannot have been derived from any single domestic stock. With respect to cattle, Professor Tanner has remarked that the lungs and liver in the improved breeds “are found to be considerably reduced in size when compared with those possessed by animals having perfect liberty” (24/23. ‘Journal of Agriculture of Highland Soc.’ July 1860 page 321.); and the reduction of these organs affects the general shape of the body. The cause of the reduced lungs in highly-bred animals which take little exercise is obvious; and perhaps the liver may be affected by the nutritious and artificial food on which they largely subsist. Again, Dr. Wilckens asserts (24/24. ‘Landwirth. Wochenblatt’ No. 10.) that various parts of the body certainly differ in Alpine and lowland breeds of several domesticated animals, owing to their different habits of life; for instance, the neck and fore-legs in length, and the hoofs in shape.
It is well known that, when an artery is tied, the anastomosing branches, from being forced to transmit more blood, increase in diameter; and this increase cannot be accounted for by mere extension, as their coats gain in strength. With respect to glands, Sir J. Paget observes that “when one kidney is destroyed the other often becomes much larger, and does double work.” (24/25. ‘Lectures on Surgical Pathology’ 1853 volume 1 page 27.) If we compare the size of the udders and their power of secretion in cows which have been long domesticated, and in certain breeds of the goat in which the udders nearly touch the ground, with these organs in wild or half-domesticated animals, the difference is great. A good cow with us daily yields more than five gallons, or forty pints of milk, whilst a first-rate animal, kept, for instance, by the Damaras of South Africa (24/26. Andersson ‘Travels in South Africa’ page 318. For analogous cases in South America see Aug. St.-Hilaire ‘Voyage dans la Province de Goyaz’ tome 1 page 71.), “rarely gives more than two or three pints of milk daily, and, should her calf be taken from her, she absolutely refuses to give any.” We may attribute the excellence of our cows and of certain goats, partly to the continued selection of the best milking animals, and partly to the inherited effects of the increased action, through man’s art, of the secreting glands.
It is notorious that short-sight is inherited; and we have seen in the twelfth chapter from the statistical researches of M. Giraud-Teulon, that the habit of viewing near objects gives a tendency to short-sight. Veterinarians are unanimous that horses are affected with spavins, splints, ringbones, etc., from being shod and from travelling on hard roads, and they are almost equally unanimous that a tendency to these malformations is transmitted. Formerly horses were not shod in North Carolina, and it has been asserted that they did not then suffer from these diseases of the legs and feet. (24/27. Brickell ‘Nat. Hist. of North Carolina’ 1739 page 53.)
Our domesticated quadrupeds are all descended, as far as is known, from species having erect ears; yet few kinds can be named, of which at least one race has not drooping ears. Cats in China, horses in parts of Russia, sheep in Italy and elsewhere, the guinea-pig formerly in Germany, goats and cattle in India, rabbits, pigs, and dogs in all long-civilised countries have dependent ears. With wild animals, which constantly use their ears like funnels to catch every passing sound, and especially to ascertain the direction whence it comes, there is not, as Mr. Blyth has remarked, any species with drooping ears except the elephant. Hence the incapacity to erect the ears is certainly in some manner the result of domestication; and this incapacity has been attributed by various authors (24/28. Livingstone quoted by Youatt on ‘Sheep’ page 142. Hodgson in ‘Journal of Asiatic Soc. of Bengal’ volume 16 1847 page 1006 etc. etc. On the other hand Dr. Wilckens argues strongly against the belief that the drooping of the ears is the result of disuse: ‘Jahrbuch der deutschen Viehzucht’ 1866.) to disuse, for animals protected by man are not compelled habitually to use their ears. Col. Hamilton Smith (24/29. ‘Naturalist’s Library’ Dogs volume 2 1840 page 104.) states that in ancient effigies of the dog, “with the exception of one Egyptian instance, no sculpture of the earlier Grecian era produces representations of hounds with completely drooping ears; those with them half pendulous are missing in the most ancient; and this character increases, by degrees, in the works of the Roman period.” Godron also has remarked “that the pigs of the ancient Egyptians had not their ears enlarged and pendent.” (24/30. ‘De l’Espece’ tome 1 1859 page 367.) But it is remarkable that the drooping of the ear is not accompanied by any decrease in size; on the contrary, animals so different as fancy rabbits, certain Indian breeds of the goat, our petted spaniels, blood- hounds, and other dogs, have enormously elongated ears, so that it would appear as if their weight had caused them to droop, aided perhaps by disuse. With rabbits, the drooping of the much elongated ears has affected even the structure of the skull.
The tail of no wild animal, as remarked to me by Mr. Blyth, is curled; whereas pigs and some races of dogs have their tails much curled. This deformity, therefore, appears to be the result of domestication, but whether in any way connected with the lessened use of the tail is doubtful.
The epidermis on our hands is easily thickened, as every one knows, by hard work. In a district of Ceylon the sheep have “horny callosities that defend their knees, and which arise from their habit of kneeling down to crop the short herbage, and this distinguishes the Jaffna flocks from those of other portions of the island;” but it is not stated whether this peculiarity is inherited. (24/31. ‘Ceylon’ by Sir J.E. Tennent 1859 volume 2 page 531.)
The mucous membrane which lines the stomach is continuous with the external skin of the body; therefore it is not surprising that its texture should be affected by the nature of the food consumed, but other and more interesting changes likewise follow. Hunter long ago observed that the muscular coat of the stomach of a gull (Larus tridactylus) which had been fed for a year chiefly on grain was thickened; and, according to Dr. Edmondston, a similar change periodically occurs in the Shetland Islands in the stomach of the Larus argentatus, which in the spring frequents the cornfields and feeds on the seed. The same careful observer has noticed a great change in the stomach of a raven which had been long fed on vegetable food. In the case of an owl (Strix grallaria), similarly treated, Menetries states that the form of the stomach was changed, the inner coat became leathery, and the liver increased in size. Whether these modifications in the digestive organs would in the course of generations become inherited is not known. (24/32. For the foregoing statements see Hunter ‘Essays and Observations’ 1861 volume 2 page 329; Dr. Edmondston, as quoted in Macgillivray ‘British Birds’ volume 5 page 550: Menetries as quoted in Bronn ‘Geschichte der Natur’ b. 2 s. 110.)
The increased or diminished length of the intestines, which apparently results from changed diet, is a more remarkable case, because it is characteristic of certain animals in their domesticated condition, and therefore must be inherited. The complex absorbent system, the blood-vessels, nerves, and muscles, are necessarily all modified together with the intestines. According to Daubenton, the intestines of the domestic cat are one-third longer than those of the wild cat of Europe; and although this species is not the parent- stock of the domestic animal, yet, as Isidore Geoffroy has remarked, the several species of cats are so closely allied that the comparison is probably a fair one. The increased length appears to be due to the domestic cat being less strictly carnivorous in its diet than any wild feline species; for instance, I have seen a French kitten eating vegetables as readily as meat. According to Cuvier, the intestines of the domesticated pig exceed greatly in proportionate length those of the wild boar. In the tame and wild rabbit the change is of an opposite nature, and probably results from the nutritious food given to the tame rabbit. (24/33. These statements on the intestines are taken from Isidore Geoffroy Saint-Hilaire ‘Hist. Nat. Gen.’ tome 3 pages 427, 441.)
CHANGED AND INHERITED HABITS OF LIFE.
This subject, as far as the mental powers of animals are concerned, so blends into instinct, that I will here only remind the reader of such cases as the tameness of our domesticated animals–the pointing or retrieving of dogs– their not attacking the smaller animals kept by man–and so forth. How much of these changes ought to be attributed to mere habit, and how much to the selection of individuals which have varied in the desired manner, irrespectively of the special circumstances under which they have been kept, can seldom be told.
We have already seen that animals may be habituated to a changed diet; but some additional instances may be given. In the Polynesian Islands and in China the dog is fed exclusively on vegetable matter, and the taste for this kind of food is to a certain extent inherited. (24/34. Gilbert White ‘Nat. Hist. Selborne’ 1825 volume 2 page 121.) Our sporting dogs will not touch the bones of game birds, whilst most other dogs devour them with greediness. In some parts of the world sheep have been largely fed on fish. The domestic hog is fond of barley, the wild boar is said to disdain it; and the disdain is partially inherited, for some young wild pigs bred in captivity showed an aversion for this grain, whilst others of the same brood relished it. (24/35. Burdach ‘Traite de Phys.’ tome 2 page 267 as quoted by Dr. P. Lucas ‘L’Hered. Nat.’ tome 1 page 388.) One of my relations bred some young pigs from a Chinese sow by a wild Alpine boar; they lived free in the park, and were so tame that they came to the house to be fed; but they would not touch swill, which was devoured by the other pigs. An animal when once accustomed to an unnatural diet, which can generally be effected only during youth, dislikes its proper food, as Spallanzani found to be the case with a pigeon which had been long fed on meat. Individuals of the same species take to new food with different degrees of readiness; one horse, it is stated, soon learned to eat meat, whilst another would have perished from hunger rather than have partaken of it. (24/36. This and several other cases are given by Colin ‘Physiologie Comp. des Animaux Dom.’ 1854 tome 1 page 426.) The caterpillars of the Bombyx hesperus feed in a state of nature on the leaves of the Cafe diable, but, after having been reared on the Ailanthus, they would not touch the Cafe diable, and actually died of hunger. (24/37. M. Michely de Cayenne in ‘Bull. Soc. d’Acclimat.’ tome 8 1861 page 563.)
It has been found possible to accustom marine fish to live in fresh water; but as such changes in fish and other marine animals have been chiefly observed in a state of nature, they do not properly belong to our present subject. The period of gestation and of maturity, as shown in the earlier chapters,–the season and the frequency of the act of breeding,–have all been greatly modified under domestication. With the Egyptian goose the rate of change with respect to the season has been recorded. (24/38. Quatrefages ‘Unite de l’Espece Humaine’ 1861 page 79.) The wild drake pairs with one female, the domestic drake is polygamous. Certain breeds of fowls have lost the habit of incubation. The paces of the horse, and the manner of flight of certain breeds of the pigeon, have been modified and are inherited. Cattle, horses, and pigs have learnt to browse under water in the St. John’s River, East Florida, where the Vallisneria has been largely naturalised. The cows were observed by Prof. Wyman to keep their heads immersed for “a period varying from fifteen to thirty-five seconds.” (24/39. ‘The American Naturalist’ April 1874 page 237.) The voice differs much in certain kinds of fowls and pigeons. Some varieties are clamorous and others silent, as the Call and common duck, or the Spitz and pointer dog. Every one knows how the breeds of the dog differ from one another in their manner of hunting, and in their ardour after different kinds of game or vermin.
With plants the period of vegetation is easily changed and is inherited, as in the case of summer and winter wheat, barley, and vetches; but to this subject we shall immediately return under acclimatisation. Annual plants sometimes become perennial under a new climate, as I hear from Dr. Hooker is the case with the stock and mignonette in Tasmania. On the other hand, perennials sometimes become annuals, as with the Ricinus in England, and as, according to Captain Mangles, with many varieties of the heartsease. Von Berg (24/40. ‘Flora’ 1835 b. 2 page 504.) raised from seed of Verbascum phoeniceum, which is usually a biennial, both annual and perennial varieties. Some deciduous bushes become evergreen in hot countries. (24/41. Alph. de Candolle ‘Geograph. Bot.’ tome 2 page 1078.) Rice requires much water, but there is one variety in India which can be grown without irrigation. (24/42. Royle ‘Illustrations of the Botany of the Himalaya’ page 19.) Certain varieties of the oat and of our other cereals are best fitted for certain soils. (24/43. ‘Gardener’s Chronicle’ 1850 pages 204, 219.) Endless similar facts could be given in the animal and vegetable kingdoms. They are noticed here because they illustrate analogous differences in closely allied natural species, and because such changed habits of life, whether due to habit, or to the direct action of external conditions, or to so-called spontaneous variability, would be apt to lead to modifications of structure.
ACCLIMATISATION. — From the previous remarks we are naturally led to the much disputed subject of acclimatisation. There are two distinct questions: Do varieties descended from the same species differ in their power of living under different climates? And secondly, if they so differ, how have they become thus adapted? We have seen that European dogs do not succeed well in India, and it is asserted (24/44. Rev. R. Everest ‘Journal As. Soc. of Bengal’ volume 3 page 19.), that no one has there succeeded in keeping the Newfoundland long alive; but then it may be argued, and probably with truth, that these northern breeds are specifically distinct from the native dogs which flourish in India. The same remark may be made with respect to different breeds of sheep, of which, according to Youatt (24/45. Youatt on ‘Sheep’ 1838 page 491.), not one brought “from a torrid climate lasts out the second year,” in the Zoological Gardens. But sheep are capable of some degree of acclimatisation, for Merino sheep bred at the Cape of Good Hope have been found far better adapted for India than those imported from England. (24/46. Royle ‘Prod. Resources of India’ page 153.) It is almost certain that all the breeds of the fowl are descended from one species; but the Spanish breed, which there is good reason to believe originated near the Mediterranean (24/47. Tegetmeier ‘Poultry Book’ 1866 page 102.), though so fine and vigorous in England, suffers more from frost than any other breed. The Arrindy silk moth introduced from Bengal, and the Ailanthus moth from the temperate province of Shan Tung, in China, belong to the same species, as we may infer from their identity in the caterpillar, cocoon, and mature states (24/48. Dr. R. Paterson in a paper communicated to Bot. Soc. of Canada quoted in the ‘Reader’ 1863 November 13.); yet they differ much in constitution: the Indian form “will flourish only in warm latitudes,” the other is quite hardy and withstands cold and rain.
Plants are more strictly adapted to climate than are animals. The latter when domesticated withstand such great diversities of climate, that we find nearly the same species in tropical and temperate countries; whilst the cultivated plants are widely dissimilar. Hence a larger field is open for inquiry in regard to the acclimatisation of plants than of animals. It is no exaggeration to say that with almost every plant which has long been cultivated, varieties exist which are endowed with constitutions fitted for very different climates; I will select only a few of the more striking cases, as it would be tedious to give all. In North America numerous fruit-trees have been raised, and in horticultural publications,–for instance, in that by Downing,–lists are given of the varieties which are best able to withstand the severe climate of the northern States and Canada. Many American varieties of the pear, plum, and peach are excellent in their own country, but until recently, hardly one was known that succeeded in England; and with apples (24/49. See remarks by Editor in ‘Gardener’s Chronicle’ 1848 page 5.), not one succeeds. Though the American varieties can withstand a severer winter than ours, the summer here is not hot enough. Fruit-trees have also originated in Europe with different constitutions, but they are not much noticed, because nurserymen here do not supply wide areas. The Forelle pear flowers early, and when the flowers have just set, and this is the critical period, they have been observed, both in France and England, to withstand with complete impunity a frost of 18 deg and even 14 deg Fahr., which killed the flowers, whether fully expanded or in bud, of all other kinds of pears. (24/50. ‘Gardener’s Chronicle’ 1860 page 938. Remarks by Editor and quotation from Decaisne.) This power in the flower of resisting cold and afterwards producing fruit does not invariably depend, as we know on good authority (24/51. J. de Jonghe of Brussels in ‘Gardener’s Chronicle’ 1857 page 612.), on general constitutional vigour. In proceeding northward, the number of varieties which are found capable of resisting the climate rapidly decreases, as may be seen in the list of the varieties of the cherry, apple, and pear, which can be cultivated in the neighbourhood of Stockholm. (24/52. Ch. Martius ‘Voyage Bot. Cotes Sept. de la Norvege’ page 26.) Near Moscow, Prince Troubetzkoy planted for experiment in the open ground several varieties of the pear, but one alone, the Poire sans Pepins, withstood the cold of winter. (24/53. ‘Journal de l’Acad. Hort. de Gand’ quoted in ‘Gardener’s Chronicle’ 1859 page 7.) We thus see that our fruit-trees, like distinct species of the same genus, certainly differ from each other in their constitutional adaptation to different climates.
With the varieties of many plants, the adaptation to climate is often very close. Thus it has been proved by repeated trials “that few if any of the English varieties of wheat are adapted for cultivation in Scotland” (24/54. ‘Gardener’s Chronicle’ 1851 page 396.); but the failure in this case is at first only in the quantity, though ultimately in the quality, of the grain produced. The Rev. M.J. Berkeley sowed wheat-seed from India, and got “the most meagre ears,” on land which would certainly have yielded a good crop from English wheat. (24/55. Ibid 1862 page 235.) In these cases varieties have been carried from a warmer to a cooler climate; in the reverse case, as “when wheat was imported directly from France into the West Indian Islands, it produced either wholly barren spikes or furnished with only two or three miserable seeds, while West Indian seed by its side yielded an enormous harvest.” (24/56. On the authority of Labat quoted in ‘Gardener’s Chronicle’ 1862 page 235.) Here is another case of close adaptation to a slightly cooler climate; a kind of wheat which in England may be used indifferently either as a winter or summer variety, when sown under the warmer climate of Grignan, in France, behaved exactly as if it had been a true winter wheat. (24/57. MM. Edwards and Colin ‘Annal. des Sc. Nat.’ 2nd series Bot. tome 5 page 22.)
Botanists believe that all the varieties of maize belong to the same species; and we have seen that in North America, in proceeding northward, the varieties cultivated in each zone produce their flowers and ripen their seed within shorter and shorter periods. So that the tall, slowly maturing southern varieties do not succeed in New England, and the New English varieties do not succeed in Canada. I have not met with any statement that the southern varieties are actually injured or killed by a degree of cold which the northern varieties can withstand with impunity, though this is probable; but the production of early flowering and early seeding varieties deserves to be considered as one form of acclimatisation. Hence it has been found possible, according to Kalm, to cultivate maize further and further northwards in America. In Europe, also, as we learn from the evidence given by Alph. De Candolle, the culture of maize has extended since the end of the last century thirty leagues north of its former boundary. (24/58. ‘Geograph. Bot.’ page 337.) On the authority of Linnaeus (24/59. ‘Swedish Acts’ English translation 1739-40 volume 1. Kalm in his ‘Travels’ volume 2 page 166 gives an analogous case with cotton-plants raised in New Jersey from Carolina seed.), I may quote an analogous case, namely, that in Sweden tobacco raised from home-grown seed ripens its seed a month sooner and is less liable to miscarry than plants raised from foreign seed.
With the Vine, differently from the maize, the line of practical culture has retreated a little southward since the middle ages (24/60. De Candolle ‘Geograph. Bot.’ page 339.); but this seems due to commerce being now easier, so that it is better to import wine from the south than to make it in northern districts. Nevertheless the fact of the vine not having spread northward shows that acclimatisation has made no progress during several centuries. There is, however, a marked difference in the constitution of the several varieties,– some being hardy, whilst others, like the muscat of Alexandria, require a very high temperature to come to perfection. According to Labat (24/61. ‘Gardener’s Chronicle’ 1862 page 235.), vines taken from France to the West Indies succeed with extreme difficulty, whilst those imported from Madeira or the Canary Islands thrive admirably.
Gallesio gives a curious account of the naturalisation of the Orange in Italy. During many centuries the sweet orange was propagated exclusively by grafts, and so often suffered from frosts, that it required protection. After the severe frost of 1709, and more especially after that of 1763, so many trees were destroyed, that seedlings from the sweet orange were raised, and, to the surprise of the inhabitants, their fruit was found to be sweet. The trees thus raised were larger, more productive, and hardier than the old kinds; and seedlings are now continually raised. Hence Gallesio concludes that much more was effected for the naturalisation of the orange in Italy by the accidental production of new kinds during a period of about sixty years, than had been effected by grafting old varieties during many ages. (24/62. Gallesio ‘Teoria della Riproduzione Veg.’ 1816 page 125; and ‘Traite du Citrus’ 1811 page 359.) I may add that Risso (24/63. ‘Essai sur l’Hist. des Orangers’ 1813 page 20 etc.) describes some Portuguese varieties of the orange as extremely sensitive to cold, and as much tenderer than certain other varieties.
The peach was known to Theophrastus, 322 B.C. (24/64. Alph. de Candolle ‘Geograph. Bot.’ page 882.) According to the authorities quoted by Dr. F. Rolle (24/65. ‘Ch. Darwin’s Lehre von der Entstehung’ etc. 1862 s. 87.), it was tender when first introduced into Greece, and even in the island of Rhodes only occasionally bore fruit. If this be correct, the peach, in spreading during the last two thousand years over the middle parts of Europe, must have become much hardier. At the present day different varieties differ much in hardiness: some French varieties will not succeed in England; and near Paris, the Pavie de Bonneuil does not ripen its fruit till very late in the season, even when grown on a wall; “it is, therefore, only fit for a very hot southern climate.” (24/66. Decaisne quoted in ‘Gardener’s Chronicle’ 1865 page 271.)
I will briefly give a few other cases. A variety of Magnolia grandiflora, raised by M. Roy, withstands a temperature several degrees lower than that which any other variety can resist. With camellias there is much difference in hardiness. One particular variety of the Noisette rose withstood the severe frost of 1860 “untouched and hale amidst a universal destruction of other Noisettes.” In New York the “Irish yew is quite hardy, but the common yew is liable to be cut down.” I may add that there are varieties of the sweet potato (Convolvulus batatas) which are suited for warmer, as well as for colder, climates. (24/67. For the magnolia see Loudon’s ‘Gardener’s Mag.’ volume 13 1837 page 21. For camellias and roses see ‘Gardener’s Chronicle’ 1860 page 384. For the yew ‘Journal of Hort.’ March 3, 1863 p 174. For sweet potatoes see Col. von Siebold in ‘Gardener’s Chronicle’ 1855 page 822.)
The plants as yet mentioned have been found capable of resisting an unusual degree of cold or heat, when fully grown. The following cases refer to plants whilst young. In a large bed of young Araucarias of the same age, growing close together and equally exposed, it was observed (24/68. The Editor ‘Gardener’s Chronicle’ 1861 page 239.), after the unusually severe winter of 1860-61, that, “in the midst of the dying, numerous individuals remained on which the frost had absolutely made no kind of impression.” Dr. Lindley, after alluding to this and other similar cases, remarks, “Among the lessons which the late formidable winter has taught us, is that, even in their power of resisting cold, individuals of the same species of plants are remarkably different.” Near Salisbury, there was a sharp frost on the night of May 24, 1836, and all the French beans (Phaseolus vulgaris) in a bed were killed except about one in thirty, which completely escaped. (24/69. Loudon’s ‘Gardener’s Mag.’ volume 12 1836 page 378.) On the same day of the month, but in the year 1864, there was a severe frost in Kent, and two rows of scarlet- runners (P. multiflorus) in my garden, containing 390 plants of the same age and equally exposed, were all blackened and killed except about a dozen plants. In an adjoining row of “Fulmer’s dwarf bean” (P. vulgaris), one single plant escaped. A still more severe frost occurred four days afterwards, and of the dozen plants which had previously escaped only three survived; these were not taller or more vigorous than the other young plants, but they escaped completely, with not even the tips of their leaves browned. It was impossible to behold these three plants, with their blackened, withered, and dead brethren all around, and not see at a glance that they differed widely in constitutional power of resisting frost.
This work is not the proper place to show that wild plants of the same species, naturally growing at different altitudes or under different latitudes, become to a certain extent acclimatised, as is proved by the different behaviour of their seedlings when raised in another country. In my ‘Origin of Species’ I have alluded to some cases, and I could add many others. One instance must suffice: Mr. Grigor, of Forres (24/70. ‘Gardener’s Chronicle’ 1865 page 699. Mr. G. Maw gives (‘Gardener’s Chronicle’ 1870 page 895) a number of striking cases; he brought home from southern Spain and northern Africa several plants, which he cultivated in England alongside specimens from northern districts; and he found a great difference not only in their hardiness during the winter, but in the behaviour of some of them during the summer.), states that seedlings of the Scotch fir (Pinus sylvestris), raised from seed from the Continent and from the forests of Scotland, differ much. “The difference is perceptible in one-year-old, and more so in two-year- old seedlings; but the effects of the winter on the second year’s growth almost uniformly make those from the Continent quite brown, and so damaged, that by the month of March they are quite unsaleable, while the plants from the native Scotch pine, under the same treatment, and standing alongside, although considerably shorter, are rather stouter and quite green, so that the beds of the one can be known from the other when seen from the distance of a mile.” Closely similar facts have been observed with seedling larches.
Hardy varieties would alone be valued or noticed in Europe; whilst tender varieties, requiring more warmth, would generally be neglected; but such occasionally arise. Thus Loudon (24/71. ‘Arboretum et Fruticetum’ volume 3 page 1376.) describes a Cornish variety of the elm which is almost an evergreen, and of which the shoots are often killed by the autumnal frosts, so that its timber is of little value. Horticulturists know that some varieties are much more tender than others: thus all the varieties of the broccoli are more tender than cabbages; but there is much difference in this respect in the sub-varieties of the broccoli; the pink and purple kinds are a little hardier than the white Cape broccoli, “but they are not to be depended on after the thermometer falls below 24 deg Fahr.;” the Walcheren broccoli is less tender than the Cape, and there are several varieties which will stand much severer cold than the Walcheren. (24/72. Mr. Robson in ‘Journal of Horticulture’ 1861 page 23.) Cauliflowers seed more freely in India than cabbages. (24/73. Dr. Bonavia ‘Report of the Agri.-Hort. Soc. of Oudh’ 1866.) To give one instance with flowers: eleven plants raised from a hollyhock, called the Queen of the Whites (24/74. ‘Cottage Gardener’ 1860 April 24 page 57.) were found to be much more tender than various other seedlings. It may be presumed that all tender varieties would succeed better under a climate warmer than ours. With fruit-trees, it is well known that certain varieties, for instance of the peach, stand forcing in a hot-house better than others; and this shows either pliability of organisation or some constitutional difference. The same individual cherry-tree, when forced, has been observed during successive years gradually to change its period of vegetation. (24/75. ‘Gardener’s Chronicle’ 1841 page 291.) Few pelargoniums can resist the heat of a stove, but Alba Multiflora will, as a most skilful gardener asserts, “stand pine-apple top and bottom heat the whole winter; without looking any more drawn than if it had stood in a common greenhouse; and Blanche Fleur seems as if it had been made on purpose for growing in winter, like many bulbs, and to rest all summer.” (24/76. Mr. Beaton in ‘Cottage Gardener’ March 20, 1860 page 377. Queen Mab will also stand stove heat. See ‘Gardener’s Chronicle’ 1845 page 226.) There can hardly be a doubt that the Alba Multiflora pelargonium must have a widely different constitution from that of most other varieties of this plant; it would probably withstand even an equatorial climate.
We have seen that according to Labat the vine and wheat require acclimatisation in order to succeed in the West Indies. Similar facts have been observed at Madras: “two parcels of mignonette-seed, one direct from Europe, the other saved at Bangalore (of which the mean temperature is much below that of Madras), were sown at the same time: they both vegetated equally favourably, but the former all died off a few days after they appeared above ground; the latter still survive, and are vigorous, healthy plants.” “So again, turnip and carrot seed saved at Hyderabad are found to answer better at Madras than seed from Europe or from the Cape of Good Hope.” (24/77. ‘Gardener’s Chronicle’ 1841 page 439.) Mr. J. Scott of the Calcutta Botanic Gardens, informs me that seeds of the sweet-pea (Lathyrus odoratus) imported from England produce plants, with thick, rigid stems and small leaves, which rarely blossom and never yield seed; plants raised from French seed blossom sparingly, but all the flowers are sterile; on the other hand, plants raised from sweet-peas grown near Darjeeling in Upper India, but originally derived from England, can be successfully cultivated on the plains of India; for they flower and seed profusely, and their stems are lax and scandent. In some of the foregoing cases, as Dr. Hooker has remarked to me, the greater success may perhaps be attributed to the seeds having been more fully ripened under a more favourable climate; but this view can hardly be extended to so many cases, including plants, which, from being cultivated under a climate hotter than their native one, become fitted for a still hotter climate. We may therefore safely conclude that plants can to a certain extent become accustomed to a climate either hotter or colder than their own; although the latter cases have been more frequently observed.
We will now consider the means by which acclimatisation may be effected, namely, through the appearance of varieties having a different constitution, and through the effects of habit. In regard to new varieties, there is no evidence that a change in the constitution of the offspring necessarily stands in any direct relation with the nature of the climate inhabited by the parents. On the contrary, it is certain that hardy and tender varieties of the same species appear in the same country. New varieties thus spontaneously arising become fitted to slightly different climates in two different ways; firstly, they may have the power, either as seedlings or when full-grown, of resisting intense cold, as with the Moscow pear, or of resisting intense heat, as with some kinds of Pelargonium, or the flowers may withstand severe frost, as with the Forelle pear. Secondly, plants may become adapted to climates widely different from their own, from flowering and fruiting either earlier or later in the season. In both these cases the power of acclimatisation by man consists simply in the selection and preservation of new varieties. But without any direct intention on his part of securing a hardier variety, acclimatisation may be unconsciously effected by merely raising tender plants from seed, and by occasionally attempting their cultivation further and further northwards, as in the case of maize, the orange and the peach.
How much influence ought to be attributed to inherited habit or custom in the acclimatisation of animals and plants is a much more difficult question. In many cases natural selection can hardly have failed to have come into play and complicated the result. It is notorious that mountain sheep resist severe weather and storms of snow which would destroy lowland breeds; but then mountain sheep have been thus exposed from time immemorial, and all delicate individuals will have been destroyed, and the hardiest preserved. So with the Arrindy silk-moths of China and India; who can tell how far natural selection may have taken a share in the formation of the two races, which are now fitted for such widely different climates? It seems at first probable that the many fruit-trees which are so well fitted for the hot summers and cold winters of North America, in contrast with their poor success under our climate, have become adapted through habit; but when we reflect on the multitude of seedlings annually raised in that country, and that none would succeed unless born with a fitting constitution, it is possible that mere habit may have done nothing towards their acclimatisation. On the other hand, when we hear that Merino sheep, bred during no great number of generations at the Cape of Good Hope–that some European plants raised during only a few generations in the cooler parts of India, withstand the hotter parts of that country much better than the sheep or seeds imported directly from England, we must attribute some influence to habit. We are led to the same conclusion when we hear from Naudin (24/78. Quoted by Asa Gray in ‘Am. Journ. of Sc.’ 2nd series January 1865 page 106.) that the races of melons, squashes, and gourds, which have long been cultivated in Northern Europe, are comparatively more precocious, and need much less heat for maturing their fruit, than the varieties of the same species recently brought from tropical regions. In the reciprocal conversion of summer and winter wheat, barley, and vetches into each other, habit produces a marked effect in the course of a very few generations. The same thing apparently occurs with the varieties of maize, which, when carried from the Southern States of America, or into Germany, soon became accustomed to their new homes. With vine-plants taken to the West Indies from Madeira, which are said to succeed better than plants brought directly from France, we have some degree of acclimatisation in the individual, independently of the production of new varieties by seed.
The common experience of agriculturists is of some value, and they often advise persons to be cautious in trying the productions of one country in another. The ancient agricultural writers of China recommend the preservation and cultivation of the varieties peculiar to each country. During the classical period, Columella wrote, “Vernaculum pecus peregrino longe praestantius est.” (24/79. For China see ‘Memoire sur les Chinois’ tome 11 1786 page 60. Columella is quoted by Carlier in ‘Journal de Physique’ tome 24 1784.)
I am aware that the attempt to acclimatise either animals or plants has been called a vain chimera. No doubt the attempt in most cases deserves to be thus called, if made independently of the production of new varieties endowed with a different constitution. With plants propagated by buds, habit rarely produces any effect; it apparently acts only through successive seminal generations. The laurel, bay, laurestinus, etc., and the Jerusalem artichoke, which are propagated by cuttings or tubers, are probably now as tender in England as when first introduced; and this appears to be the case with the potato, which until recently was seldom multiplied by seed. With plants propagated by seed, and with animals, there will be little or no acclimatisation unless the hardier individuals are either intentionally or unconsciously preserved. The kidney-bean has often been advanced as an instance of a plant which has not become hardier since its first introduction into Britain. We hear, however, on excellent authority (24/80. Messrs. Hardy and Son in ‘Gardener’s Chronicle’ 1856 page 589.) that some very fine seed, imported from abroad, produced plants “which blossomed most profusely, but were nearly all but abortive, whilst plants grown alongside from English seed podded abundantly;” and this apparently shows some degree of acclimatisation in our English plants. We have also seen that seedlings of the kidney-bean occasionally appear with a marked power of resisting frost; but no one, as far as I can hear, has ever separated such hardy seedlings, so as to prevent accidental crossing, and then gathered their seed, and repeated the process year after year. It may, however, be objected with truth that natural selection ought to have had a decided effect on the hardiness of our kidney- beans; for the tenderest individuals must have been killed during every severe spring, and the hardier preserved. But it should be borne in mind that the result of increased hardiness would simply be that gardeners, who are always anxious for as early a crop as possible, would sow their seed a few days earlier than formerly. Now, as the period of sowing depends much on the soil and elevation of each district, and varies with the season; and as new varieties have often been imported from abroad, can we feel sure that our kidney-beans are not somewhat hardier? I have not been able, by searching old horticultural works, to answer this question satisfactorily.
On the whole the facts now given show that, though habit does something towards acclimatisation, yet that the appearance of constitutionally different individuals is a far more effective agent. As no single instance has been recorded either with animals or plants of hardier individuals having been long and steadily selected, though such selection is admitted to be indispensable for the improvement of any other character, it is not surprising that man has done little in the acclimatisation of domesticated animals and cultivated plants. We need not, however, doubt that under nature new races and new species would become adapted to widely different climates, by variation, aided by habit, and regulated by natural selection.
ARRESTS OF DEVELOPMENT: RUDIMENTARY AND ABORTED ORGANS.
Modifications of structure from arrested development, so great or so serious as to deserve to be called monstrosities, are not infrequent with domesticated animals, but, as they differ much from any normal structure, they require only a passing notice. Thus the whole head may be represented by a soft nipple-like projection, and the limbs by mere papillae. These rudiments of limbs are sometimes inherited, as has been observed in a dog. (24/81. Isid. Geoffroy Saint-Hilaire ‘Hist. Nat. des Anomalies’ 1836 tome 2 pages 210, 223, 224, 395; ‘Philosoph. Transact.’ 1775 page 313.)
Many lesser anomalies appear to be due to arrested development. What the cause of the arrest may be, we seldom know, except in the case of direct injury to the embryo. That the cause does not generally act at an extremely early embryonic period we may infer from the affected organ seldom being wholly aborted,–a rudiment being generally preserved. The external ears are represented by mere vestiges in a Chinese breed of sheep; and in another breed, the tail is reduced “to a little button, suffocated in a manner, by fat.” (24/82. Pallas quoted by Youatt on ‘Sheep’ page 25.) In tailless dogs and cats a stump is left. In certain breeds of fowls the comb and wattles are reduced to rudiments; in the Cochin-China breed scarcely more than rudiments of spurs exist. With polled Suffolk cattle, “rudiments of horns can often be felt at an early age” (24/83. Youatt on ‘Cattle’ 1834 page 174.); and with species in a state of nature, the relatively great development of rudimentary organs at an early period of life is highly characteristic of such organs. With hornless breeds of cattle and sheep, another and singular kind of rudiment has been observed, namely, minute dangling horns attached to the skin alone, and which are often shed and grow again. With hornless goats, according to Desmarest (24/84. ‘Encyclop. Method.’ 1820 page 483: see page 500, on the Indian zebu casting its horns. Similar cases in European cattle were given in the third chapter.), the bony protuberance which properly supports the horn exists as a mere rudiment.
With cultivated plants it is far from rare to find the petals, stamens, and pistils represented by rudiments, like those observed in natural species. So it is with the whole seed in many fruits; thus, near Astrakhan there is a grape with mere traces of seeds, “so small and lying so near the stalk that they are not perceived in eating the grape.” (24/85. Pallas ‘Travels’ English Translat. volume 1 page 243.) In certain varieties of the gourd, the tendrils, according to Naudin, are represented by rudiments or by various monstrous growths. In the broccoli and cauliflower the greater number of the flowers are incapable of expansion, and include rudimentary organs. In the Feather hyacinth (Muscari comosum) in its natural state the upper and central flowers are brightly coloured but rudimentary; under cultivation the tendency to abortion travels downwards and outwards, and all the flowers become rudimentary; but the abortive stamens and pistils are not so small in the lower as in the upper flowers. In the Viburnum opulus, on the other hand, the outer flowers naturally have their organs of fructification in a rudimentary state, and the corolla is of large size; under cultivation, the change spreads to the centre, and all the flowers become affected. In the compositae, the so- called doubling of the flowers consists in the greater development of the corolla of the central florets, generally accompanied with some degree of sterility; and it has been observed (24/86. Mr. Beaton in ‘Journal of Horticulture’ May 21, 1861 page 133.) that the progressive doubling invariably spreads from the circumference to the centre,–that is, from the ray florets, which so often include rudimentary organs, to those of the disc. I may add, as bearing on this subject, that with Asters, seeds taken from the florets of the circumference have been found to yield the greatest number of double flowers. (24/87. Lecoq ‘De la Fecondation’ 1862 page 233.) In the above cases we have a natural tendency in certain parts to be rudimentary, and this under culture spreads either to, or from, the axis of the plant. It deserves notice, as showing how the same laws govern the changes which natural species and artificial varieties undergo, that in the species of Carthamus, one of the Compositae, a tendency to the abortion of the pappus may be traced extending from the circumference to the centre of the disc as in the so-called doubling of the flowers in the members of the same family. Thus, according to A. de Jussieu (24/88. ‘Annales du Museum’ tome 6 page 319.), the abortion is only partial in Carthamus creticus, but more extended in C. lanatus; for in this species only two or three of the central seeds are furnished with a pappus, the surrounding seeds being either quite naked or furnished with a few hairs; and lastly in C. tinctorius, even the central seeds are destitute of pappus, and the abortion is complete.
With animals and plants under domestication, when an organ disappears, leaving only a rudiment, the loss has generally been sudden, as with hornless and tailless breeds; and such cases may be ranked as inherited monstrosities. But in some few cases the loss has been gradual, and has been effected partly by selection, as with the rudimentary combs and wattles of certain fowls. We have also seen that the wings of some domesticated birds have been slightly reduced by disuse, and the great reduction of the wings in certain silk-moths, with mere rudiments left, has probably been aided by disuse.
With species in a state of nature, rudimentary organs are extremely common. Such organs are generally variable, as several naturalists have observed; for, being useless, they are not regulated by natural selection, and they are more or less liable to reversion. The same rule certainly holds good with parts which have become rudimentary under domestication. We do not know through what steps under nature rudimentary organs have passed in being reduced to their present condition; but we so incessantly see in species of the same group the finest gradations between an organ in a rudimentary and perfect state, that we are led to believe that the passage must have been extremely gradual. It may be doubted whether a change of structure so abrupt as the sudden loss of an organ would ever be of service to a species in a state of nature; for the conditions to which all organisms are closely adapted usually change very slowly. Even if an organ did suddenly disappear in some one individual by an arrest of development, intercrossing with the other individuals of the same species would tend to cause its partial reappearance; so that its final reduction could only be effected by some other means. The most probable view is, that a part which is now rudimentary, was formerly, owing to changed habits of life, used less and less, being at the same time reduced in size by disuse, until at last it became quite useless and superfluous. But as most parts or organs are not brought into action during an early period of life, disuse or decreased action will not lead to their reduction until the organism arrives at a somewhat advanced age; and from the principle of inheritance at corresponding ages the reduction will be transmitted to the offspring at the same advanced stage of growth. The part or organ will thus retain its full size in the embryo, as we know to be the case with most rudiments. As soon as a part becomes useless, another principle, that of economy of growth, will come into play, as it would be an advantage to an organism exposed to severe competition to save the development of any useless part; and individuals having the part less developed will have a slight advantage over others. But, as Mr. Mivart has justly remarked, as soon as a part is much reduced, the saving from its further reduction will be utterly insignificant; so that this cannot be effected by natural selection. This manifestly holds good if the part be formed of mere cellular tissue, entailing little expenditure of nutriment. How then can the further reduction of an already somewhat reduced part be effected? That this has occurred repeatedly under Nature is shown by the many gradations which exist between organs in a perfect state and the merest vestiges of them. Mr. Romanes (24/89. I suggested in ‘Nature’ (volume 8 pages 432, 505) that with organisms subjected to unfavourable conditions all the parts would tend towards reduction, and that under such circumstances any part which was not kept up to its standard size by natural selection would, owing to intercrossing, slowly but steadily decrease. In three subsequent communications to ‘Nature’ (March 12, April 9, and July 2, 1874), Mr. Romanes gives his improved view.) has, I think, thrown much light on this difficult problem. His view, as far as it can be given in a few words, is as follows: all parts are somewhat variable and fluctuate in size round an average point. Now, when a part has already begun from any cause to decrease, it is very improbable that the variations should be as great in the direction of increase as of diminution; for the previous reduction shows that circumstances have not been favourable for its development; whilst there is nothing to check variations in the opposite direction. If this be so, the long continued crossing of many individuals furnished with an organ which fluctuates in a greater degree towards decrease than towards increase, will slowly but steadily lead to its diminution. With respect to the complete and absolute abortion of a part, a distinct principle, which will be discussed in the chapter on pangenesis, probably comes into action.
With animals and plants reared by man there is no severe or recurrent struggle for existence, and the principle of economy will not come into action, so that the reduction of an organ will not thus be aided. So far, indeed, is this from being the case, that in some few instances organs, which are naturally rudimentary in the parent-species, become partially redeveloped in the domesticated descendants. Thus cows, like most other ruminants, properly have four active and two rudimentary mamma; but in our domesticated animals, the latter occasionally become considerably developed and yield milk. The atrophied mammae, which, in male domesticated animals, including man, have in some rare cases grown to full size and secreted milk, perhaps offer an analogous case. The hind feet of dogs naturally include rudiments of a fifth toe, and in certain large breeds these toes, though still rudimentary, become considerably developed and are furnished with claws. In the common Hen, the spurs and comb are rudimentary, but in certain breeds these become, independently of age or disease of the ovaria, well developed. The stallion has canine teeth, but the mare has only traces of the alveoli, which, as I am informed by the eminent veterinarian Mr. G.T. Brown, frequently contain minute irregular nodules of bone. These nodules, however, sometimes become developed into imperfect teeth, protruding through the gums and coated with enamel; and occasionally they grow to a fourth or even a third of the length of the canines in the stallion. With plants I do not know whether the redevelopment of rudimentary organs occurs more frequently under culture than under nature. Perhaps the pear-tree may be a case in point, for when wild it bears thorns, which consist of branches in a rudimentary condition and serve as a protection, but, when the tree is cultivated, they are reconverted into branches.
CHAPTER 2. XXV. LAWS OF VARIATION, continued.–CORRELATED VARIABILITY.
All parts of the organisation are to a certain extent connected together; but the connection may be so slight that it hardly exists, as with compound animals or the buds on the same tree. Even in the higher animals various parts are not at all closely related; for one part may be wholly suppressed or rendered monstrous without any other part of the body being affected. But in some cases, when one part varies, certain other parts always, or nearly always, simultaneously vary; they are then subject to the law of correlated variation. The whole body is admirably co-ordinated for the peculiar habits of life of each organic being, and may be said, as the Duke of Argyll insists in his ‘Reign of Law’ to be correlated for this purpose. Again, in large groups of animals certain structures always co-exist: for instance, a peculiar form of stomach with teeth of peculiar form, and such structures may in one sense be said to be correlated. But these cases have no necessary connection with the law to be discussed in the present chapter; for we do not know that the initial or primary variations of the several parts were in any way related: slight modifications or individual differences may have been preserved, first in one and then in another part, until the final and perfectly co-adapted structure was acquired; but to this subject I shall presently recur. Again, in many groups of animals the males alone are furnished with weapons, or are ornamented with gay colours; and these characters manifestly stand in some sort of correlation with the male reproductive organs, for when the latter are destroyed these characters disappear. But it was shown in the twelfth chapter that the very same peculiarity may become attached at any age to either sex, and afterwards be exclusively transmitted to the same sex at a corresponding age. In these cases we have inheritance limited by both sex and age; but we have no reason for supposing that the original cause of the variation was necessarily connected with the reproductive organs, or with the age of the affected being.
In cases of true correlated variation, we are sometimes able to see the nature of the connection; but in most cases it is hidden from us, and certainly differs in different cases. We can seldom say which of two correlated parts first varies, and induces a change in the other; or whether the two are the effects of some common cause. Correlated variation is an important subject for us; for when one part is modified through continued selection, either by man or under nature, other parts of the organisation will be unavoidably modified. From this correlation it apparently follows that with our domesticated animals and plants, varieties rarely or never differ from one another by a single character alone.
One of the simplest cases of correlation is that a modification which arises during an early stage of growth tends to influence the subsequent development of the same part, as well as of other and intimately connected parts. Isidore Geoffroy Saint-Hilaire states (25/1. ‘Hist. des Anomalies’ tome 3 page 392. Prof. Huxley applies the same principle in accounting for the remarkable, though normal, differences in the arrangement of the nervous system in the Mollusca, in his paper on the Morphology of the Cephalous Mollusca in ‘Phil. Transact.’ 1853 page 56.) that this may constantly be observed with monstrosities in the animal kingdom; and Moquin-Tandon (25/2. ‘Elements de Teratologie Veg.’ 1841 page 13.) remarks, that, as with plants the axis cannot become monstrous without in some way affecting the organs subsequently produced from it, so axial anomalies are almost always accompanied by deviations of structure in the appended parts. We shall presently see that with short-muzzled races of the dog certain histological changes in the basal elements of the bones arrest their development and shorten them, and this affects the position of the subsequently developed molar teeth. It is probable that certain modifications in the larvae of insects would affect the structure of the mature insects. But we must be careful not to extend this view too far, for during the normal course of development, certain species pass through an extraordinary course of change, whilst other and closely allied species arrive at maturity with little change of structure.
Another simple case of correlation is that with the increased or decreased dimensions of the whole body, or of any particular part, certain organs are increased or diminished in number, or are otherwise modified. Thus pigeon fanciers have gone on selecting pouters for length of body, and we have seen that their vertebrae are generally increased not only in size but in number, and their ribs in breadth. Tumblers have been selected for their small bodies, and their ribs and primary wing-feathers are generally lessened in number. Fantails have been selected for their large widely-expanded tails, with numerous tail-feathers, and the caudal vertebrae are increased in size and number. Carriers have been selected for length of beak, and their tongues have become longer, but not in strict accordance with the length of beak. In this latter breed and in others having large feet, the number of the scutellae on the toes is greater than in the breeds with small feet. Many similar cases could be given. In Germany it has been observed that the period of gestation is longer in large than in small breeds of cattle. With our highly-improved breeds of all kinds, the periods of maturity and of reproduction have advanced with respect to the age of the animal; and, in correspondence with this, the teeth are now developed earlier than formerly, so that, to the surprise of agriculturists, the ancient rules for judging of the age of an animal by the state of its teeth are no longer trustworthy. (25/3. Prof. J.B. Simonds on the Age of the Ox, Sheep, etc. quoted in ‘Gardener’s Chronicle’ 1854 page 588.)
CORRELATED VARIATION OF HOMOLOGOUS PARTS. — Parts which are homologous tend to vary in the same manner; and this is what might have been expected, for such parts are identical in form and structure during an early period of embryonic development, and are exposed in the egg or womb to similar conditions. The symmetry, in most kinds of animals, of the corresponding or homologous organs on the right and left sides of the body, is the simplest case in point; but this symmetry sometimes fails, as with rabbits having only one ear, or stags with one horn, or with many-horned sheep which sometimes carry an additional horn on one side of their heads. With flowers which have regular corollas, all the petals generally vary in the same manner, as we see in the complicated and symmetrical pattern, on the flowers, for instance, of the Chinese pink; but with irregular flowers, though the petals are of course homologous, this symmetry often fails, as with the varieties of the Antirrhinum or snapdragon, or that variety of the kidney-bean (Phaseolus) which has a white standard-petal.
In the Vertebrata the front and hind limbs are homologous, and they tend to vary in the same manner, as we see in long and short legged, or in thick and thin legged races of the horse and dog. Isidore Geoffroy (25/4. ‘Hist. des Anomalies’ tome 1 page 674.) has remarked on the tendency of supernumerary digits in man to appear, not only on the right and left sides, but on the upper and lower extremities. Meckel has insisted (25/5. Quoted by Isid. Geoffroy ibid tome 1 page 635.) that, when the muscles of the arm depart in number or arrangement from their proper type, they almost always imitate those of the leg; and so conversely the varying muscles of the leg imitate the normal muscles of the arm.
In several distinct breeds of the pigeon and fowl, the legs and the two outer toes are heavily feathered, so that in the trumpeter pigeon they appear like little wings. In the feather-legged bantam the “boots” or feathers, which grow from the outside of the leg and generally from the two outer toes, have, according to the excellent authority of Mr. Hewitt (25/6. ‘The Poultry Book’ by W.B. Tegetmeier 1866 page 250.), been seen to exceed the wing-feathers in length, and in one case were actually nine and a half inches long! As Mr. Blyth has remarked to me, these leg-feathers resemble the primary wing- feathers, and are totally unlike the fine down which naturally grows on the legs of some birds, such as grouse and owls. Hence it may be suspected that excess of food has first given redundancy to the plumage, and then that the law of homologous variation has led to the development of feathers on the legs, in a position corresponding with those on the wing, namely, on the outside of the tarsi and toes. I am strengthened in this belief by the following curious case of correlation, which for a long time seemed to me utterly inexplicable, namely, that in pigeons of any breed, if the legs are feathered, the two outer toes are partially connected by skin. These two outer toes correspond with our third and fourth toes. (25/7. Naturalists differ with respect to the homologies of the digits of birds; but several uphold the view above advanced. See on this subject Dr. E.S. Morse in ‘Annals of the Lyceum of Nat. Hist. of New York’ volume 10 1872 page 16.) Now, in the wing of the pigeon or of any other bird, the first and fifth digits are aborted; the second is rudimentary and carries the so-called “bastard-wing;” whilst the third and fourth digits are completely united and enclosed by skin, together forming the extremity of the wing. So that in feather-footed pigeons, not only does the exterior surface support a row of long feathers, like wing-feathers, but the very same digits which in the wing are completely united by skin become partially united by skin in the feet; and thus by the law of the correlated variation of homologous parts we can understand the curious connection of feathered legs and membrane between the two outer toes.
Andrew Knight (24/8. A. Walker on ‘Intermarriage’ 1838 page 160.) has remarked that the face or head and the limbs usually vary together in general proportions. Compare, for instance, the limbs of a dray and race horse, or of a greyhound and mastiff. What a monster a greyhound would appear with the head of a mastiff! The modern bulldog, however, has fine limbs, but this is a recently-selected character. From the measurements given in the sixth chapter, we see that in several breeds of the pigeon the length of the beak and the size of the feet are correlated. The view which, as before explained, seems the most probable is, that disuse in all cases tends to diminish the feet, the beak becoming at the same time shorter through correlation; but that in some few breeds in which length of beak has been a selected point, the feet, notwithstanding disuse, have increased in size through correlation. In the following case some kind of correlation is seen to exist between the feet and beak: several specimens have been sent to Mr. Bartlett at different times, as hybrids between ducks and fowls, and I have seen one; these were, as might be expected, ordinary ducks in a semi-monstrous condition, and in all of them the swimming-web between the toes was quite deficient or much reduced, and in all the beak was narrow and ill-shaped.
With the increased length of the beak in pigeons, not only the tongue increases in length, but likewise the orifice of the nostrils. But the increased length of the orifice of the nostrils perhaps stands in closer correlation with the development of the corrugated skin or wattle at the base of the beak, for when there is much wattle round the eyes, the eyelids are greatly increased or even doubled in length.
There is apparently some correlation even in colour between the head and the extremities. Thus with horses a large white star or blaze on the forehead is generally accompanied by white feet. (25/9. ‘The Farrier and Naturalist’ volume 1 1828 page 456. A gentleman who has attended to this point, tells me that about three-fourths of white-faced horses have white legs.) With white rabbits and cattle, dark marks often co-exist on the tips of the ears and on the feet. In black and tan dogs of different breeds, tan-coloured spots over the eyes and tan-coloured feet almost invariably go together. These latter cases of connected colouring may be due either to reversion or to analogous variation,–subjects to which I shall hereafter return,–but this does not necessarily determine the question of their original correlation. Mr. H.W. Jackson informs me that he has observed many hundred white-footed cats, and he finds that all are more or less conspicuously marked with white on the front of the neck or chest.
The lopping forwards and downwards of the immense ears of fancy rabbits seems partly due to the disuse of the muscles, and partly to the weight and length of the ears, which have been increased by selection during many generations. Now, with the increased size and changed direction of the ears not only has the bony auditory meatus become changed in outline, direction, and greatly in size, but the whole skull has been slightly modified. This could be clearly seen in “half-lops”–that is, in rabbits with only one ear lopping forward– for the opposite sides of their skulls were not strictly symmetrical. This seems to me a curious instance of correlation, between hard bones and organs so soft and flexible, as well as so unimportant under a physiological point of view, as the external ears. The result no doubt is largely due to mere mechanical action, that is, to the weight of the ears, on the same principle that the skull of a human infant is easily modified by pressure.
The skin and the appendages of hair, feathers, hoofs, horns, and teeth, are homologous over the whole body. Every one knows that the colour of the skin and that of the hair usually vary together; so that Virgil advises the shepherd to look whether the mouth and tongue of the ram are black, lest the lambs should not be purely white. The colour of the skin and hair, and the odour emitted by the glands of the skin, are said (25/10. Godron ‘Sur l’Espece’ tome 2 page 217.) to be connected, even in the same race of men. Generally the hair varies in the same way all over the body in length, fineness, and curliness. The same rule holds good with feathers, as we see with the laced and frizzled breeds both of fowls and pigeons. In the common cock the feathers on the neck and loins are always of a particular shape, called hackles: now in the Polish breed, both sexes are characterised by a tuft of feathers on the head, and through correlation these feathers in the male always assume the form of hackles. The wing and tail-feathers, though arising from parts not homologous, vary in length together; so that long or short winged pigeons generally have long or short tails. The case of the Jacobin-pigeon is more curious, for the wing and tail feathers are remarkably long; and this apparently has arisen in correlation with the elongated and reversed feathers on the back of the neck, which form the hood.
The hoofs and hair are homologous appendages; and a careful observer, namely Azara (25/11. ‘Quadrupedes du Paraguay’ tome 2 page 333.), states that in Paraguay horses of various colours are often born with their hair curled and twisted like that on the head of a negro. This peculiarity is strongly inherited. But what is remarkable is that the hoofs of these horses “are absolutely like those of a mule.” The hair also of their manes and tails is invariably much shorter than usual, being only from four to twelve inches in length; so that curliness and shortness of the hair are here, as with the negro, apparently correlated.
With respect to the horns of sheep, Youatt (25/12. ‘On Sheep’ page 142.) remarks that “multiplicity of horns is not found in any breed of much value; it is generally accompanied by great length and coarseness of the fleece.” Several tropical breeds of sheep which are clothed with hair instead of wool, have horns almost like those of a goat. Sturm (25/13. ‘Ueber Racen, Kreuzungen’ etc. 1825 s. 24.) expressly declares that in different races the more the wool is curled the more the horns are spirally twisted. We have seen in the third chapter, where other analogous facts have been given, that the parent of the Mauchamp breed, so famous for its fleece, had peculiarly shaped horns. The inhabitants of Angora assert (25/14. Quoted from Conolly in ‘The Indian Field’ February 1859 volume 2 page 266.) that “only the white goats which have horns wear the fleece in the long curly locks that are so much admired; those which are not horned having a comparatively close coat.” From these cases we may infer that the hair or wool and the horns tend to vary in a correlated manner. (25/15. In the third chapter I have said that “the hair and horns are so closely related to each other, that they are apt to vary together.” Dr. Wilckens (“Darwin’s Theorie” ‘Jahrbuch der Deutschen Viehzucht’ 1866 1. Heft) translates my words into “lang- und grobhaarige Thiere sollen geneigter sein, lange und viele Horner zu bekommen” and he then justly disputes this proposition; but what I have really said, in accordance with the authorities just quoted, may, I think, be trusted.) Those who have tried hydropathy are aware that the frequent application of cold water stimulates the skin; and whatever stimulates the skin tends to increase the growth of the hair, as is well shown in the abnormal growth of hair near old inflamed surfaces. Now, Professor Low (25/16. ‘Domesticated Animals of the British Islands’ pages 307, 368. Dr. Wilckens argues (‘Landwirth. Wochenblatt’ Nr. 10 1869) to the same effect with respect to domestic animals in Germany.) is convinced that with the different races of British cattle thick skin and long hair depend on the humidity of the climate which they inhabit. We can thus see how a humid climate might act on the horns–in the first place directly on the skin and hair, and secondly by correlation on the horns. The presence or absence of horns, moreover, both in the case of sheep and cattle, acts, as will presently be shown, by some sort of correlation on the skull.
With respect to hair and teeth, Mr. Yarrell (25/17. ‘Proceedings Zoolog. Soc.’ 1833 page 113.) found many of the teeth deficient in three hairless “Egyptian dogs,” and in a hairless terrier. The incisors, canines, and the premolars suffered most, but in one case all the teeth, except the large tubercular molar on each side, were deficient. With man several striking cases have been recorded (25/18. Sedgwick ‘Brit. and Foreign Medico-Chirurg. Review’ April 1863 page 453.) of inherited baldness with inherited deficiency, either complete or partial, of the teeth. I may give an analogous case, communicated to me by Mr. W. Wedderburn, of a Hindoo family in Scinde, in which ten men, in the course of four generations, were furnished, in both jaws taken together, with only four small and weak incisor teeth and with eight posterior molars. The men thus affected have very little hair on the body, and become bald early in life. They also suffer much during hot weather from excessive dryness of the skin. It is remarkable that no instance has occurred of a daughter being thus affected; and this fact reminds us how much more liable men are in England to become bald than women. Though the daughters in the above family are never affected, they transmit the tendency to their sons; and no case has occurred of a son transmitting it to his sons. The affection thus appears only in alternate generations, or after longer intervals. There is a similar connection between hair and teeth, according to Mr. Sedgwick, in those rare cases in which the hair has been renewed in old age, for this has “usually been accompanied by a renewal of the teeth.” I have remarked in a former part of this volume that the great reduction in the size of the tusks in domestic boars probably stands in close relation with their diminished bristles, due to a certain amount of protection; and that the reappearance of the tusks in boars, which have become feral and are fully exposed to the weather, probably depends on the reappearance of the bristles. I may add, though not strictly connected with our present point, that an agriculturist (25/19. ‘Gardener’s Chronicle’ 1849 page 205.) asserts that “pigs with little hair on their bodies are most liable to lose their tails, showing a weakness of the tegumental structure. It may be prevented by crossing with a more hairy breed.”
In the previous cases deficient hair, and teeth deficient in number or size, are apparently connected. In the following cases abnormally redundant hair, and teeth either deficient or redundant, are likewise connected. Mr. Crawfurd (25/20. ‘Embassy to the Court of Ava’ volume 1 page 320.) saw at the Burmese Court a man, thirty years old, with his whole body, except the hands and feet, covered with straight silky hair, which on the shoulders and spine was five inches in length. At birth the ears alone were covered. He did not arrive at puberty, or shed his milk teeth, until twenty years old; and at this period he acquired five teeth in the upper jaw, namely, four incisors and one canine, and four incisor teeth in the lower jaw; all the teeth were small. This man had a daughter who was born with hair within her ears; and the hair soon extended over her body. When Captain Yule (25/21. ‘Narrative of a Mission to the Court of Ava in 1855’ page 94.) visited the Court, he found this girl grown up; and she presented a strange appearance with even her nose densely covered with soft hair. Like her father, she was furnished with incisor teeth alone. The King had with difficulty bribed a man to marry her, and of her two children, one, a boy fourteen months old, had hair growing out of his ears, with a beard and moustache. This strange peculiarity has, therefore, been inherited for three generations, with the molar teeth deficient in the grandfather and mother; whether these teeth would likewise fail in the infant could not then be told.
A parallel case of a man fifty-five years old, and of his son, with their faces covered with hair, has recently occurred in Russia. Dr. Alex. Brandt has sent me an account of this case, together with specimens of the extremely fine hair from the cheeks. The man is deficient in teeth, possessing only four incisors in the lower and two in the upper jaw. His son, about three years old, has no teeth except four lower incisors. The case, as Dr. Brandt remarks in his letter, no doubt is due to an arrest of development in the hair and teeth. We here see how independent of the ordinary conditions of existence such arrests must be, for the lives of a Russian peasant and of a native of Burmah are as different as possible. (25/22. I owe to the kindness of M. Chauman, of St. Petersburg, excellent photographs of this man and his son, both of whom have since been exhibited in Paris and London.)
Here is another and somewhat different case communicated to me by Mr. Wallace on the authority of Dr. Purland, a dentist: Julia Pastrana, a Spanish dancer, was a remarkably fine woman, but she had a thick masculine beard and a hairy forehead; she was photographed, and her stuffed skin was exhibited as a show; but what concerns us is, that she had in both the upper and lower jaw an irregular double set of teeth, one row being placed within the other, of which Dr. Purland took a cast. From the redundancy of teeth her mouth projected, and her face had a gorilla-like appearance. These cases and those of the hairless dogs forcibly call to mind the fact, that the two orders of mammals–namely, the Edentata and Cetacea–which are the most abnormal in their dermal covering, are likewise the most abnormal either by deficiency or redundancy of teeth.
The organs of sight and hearing are generally admitted to be homologous with one another and with various dermal appendages; hence these parts are liable to be abnormally affected in conjunction. Mr. White Cowper says “that in all cases of double microphthalmia brought under his notice he has at the same time met with defective development of the dental system.” Certain forms of blindness seem to be associated with the colour of the hair; a man with black hair and a woman with light-coloured hair, both of sound constitution, married and had nine children, all of whom were born blind; of these children, five “with dark hair and brown iris were afflicted with amaurosis; the four others, with light-coloured hair and blue iris, had amaurosis and cataract conjoined.” Several cases could be given, showing that some relation exists between various affections of the eyes and ears; thus Liebreich states that out of 241 deaf-mutes in Berlin, no less than fourteen suffered from the rare disease called pigmentary retinitis. Mr. White Cowper and Dr. Earle have remarked that inability to distinguish different colours, or colour-blindness, “is often associated with a corresponding inability to distinguish musical sounds.” (25/23. These statements are taken from Mr. Sedgwick in the ‘Medico-Chirurg. Review’ July 1861 page 198; April 1863 pages 455 and 458. Liebreich is quoted by Professor Devay in his ‘Mariages Consanguins’ 1862 page 116.)
Here is a more curious case: white cats, if they have blue eyes, are almost always deaf. I formerly thought that the rule was invariable, but I have heard of a few authentic exceptions. The first two notices were published in 1829 and relate to English and Persian cats: of the latter, the Rev. W.T. Bree possessed a female, and he states, “that of the offspring produced at one and the same birth, such as, like the mother, were entirely white (with blue eyes) were, like her, invariably deaf; while those that had the least speck of colour on their fur, as invariably possessed the usual faculty of hearing.” (25/24. Loudon’s ‘Mag. of Nat. Hist.’ volume 1 1829 pages 66, 178. See also Dr. P. Lucas ‘L’Hered. Nat.’ tome 1 page 428 on the inheritance of deafness in cats. Mr. Lawson Tait states (‘Nature’ 1873 page 323) that only male cats are thus affected; but this must be a hasty generalisation. The first case recorded in England by Mr. Bree related to a female, and Mr. Fox informs me that he has bred kittens from a white female with blue eyes, which was completely deaf; he has also observed other females in the same condition.) The Rev. W. Darwin Fox informs me that he has seen more than a dozen instances of this correlation in English, Persian, and Danish cats; but he adds “that, if one eye, as I have several times observed, be not blue, the cat hears. On the other hand, I have never seen a white cat with eyes of the common colour that was deaf.” In France Dr. Sichel (25/25. ‘Annales des Sc. Nat.’ Zoolog. 3rd series 1847 tome 8 page 239.) has observed during twenty years similar facts; he adds the remarkable case of the iris beginning, at the end of four months, to grow dark-coloured, and then the cat first began to hear.
This case of correlation in cats has struck many persons as marvellous. There is nothing unusual in the relation between blue eyes and white fur; and we have already seen that the organs of sight and hearing are often simultaneously affected. In the present instance the cause probably lies in a slight arrest of development in the nervous system in connection with the sense-organs. Kittens during the first nine days, whilst their eyes are closed, appear to be completely deaf; I have made a great clanging noise with a poker and shovel close to their heads, both when they were asleep and awake, without producing any effect. The trial must not be made by shouting close to their ears, for they are, even when asleep, extremely sensitive to a breath of air. Now, as long as the eyes continue closed, the iris is no doubt blue, for in all the kittens which I have seen this colour remains for some time after the eyelids open. Hence, if we suppose the development of the organs of sight and hearing to be arrested at the stage of the closed eyelids, the eyes would remain permanently blue and the ears would be incapable of perceiving sound; and we should thus understand this curious case. As, however, the colour of the fur is determined long before birth, and as the blueness of the eyes and the whiteness of the fur are obviously connected, we must believe that some primary cause acts at a much earlier period.
The instances of correlated variability hitherto given have been chiefly drawn from the animal kingdom, and we will now turn to plants. Leaves, sepals, petals, stamens, and pistils are all homologous. In double flowers we see that the stamens and pistils vary in the same manner, and assume the form and colour of the petals. In the double columbine (Aquilegia vulgaris), the successive whorls of stamens are converted into cornucopias, which are enclosed within one another and resemble the true petals. In hose-in-hose flowers the sepals mock the petals. In some cases the flowers and leaves vary together in tint: in all the varieties of the common pea, which have purple flowers, a purple mark may be seen on the stipules.
M. Faivre states that with the varieties of Primula sinensis the colour of the flower is evidently correlated with the colour of the under side of the leaves; and he adds that the varieties with fimbriated flowers almost always have voluminous, balloon-like calyces. (25/26. ‘Revue des Cours Scientifiques’ June 5, 1869 page 430.) With other plants the leaves and fruit or seeds vary together in colour, as in a curious pale-leaved variety of the sycamore, which has recently been described in France (25/27. ‘Gardener’s Chronicle’ 1864 page 1202.), and as in the purple-leaved hazel, in which the leaves, the husk of the nut, and the pellicle round the kernel are all coloured purple. (25/28. Verlot gives several other instances ‘Des Varietes’ 1865 page 72.) Pomologists can predict to a certain extent, from the size and appearance of the leaves of their seedlings, the probable nature of the fruit; for, as Van Mons remarks (25/29. ‘Arbres Fruitiers’ 1836 tome 2 pages 204, 226.) variations in the leaves are generally accompanied by some modification in the flower, and consequently in the fruit. In the Serpent melon, which has a narrow tortuous fruit above a yard in length, the stem of the plant, the peduncle of the female flower, and the middle lobe of the leaf, are all elongated in a remarkable manner. On the other hand, several varieties of Cucurbita, which have dwarfed stems, all produce, as Naudin remarks, leaves of the same peculiar shape. Mr. G. Maw informs me that all the varieties of the scarlet Pelargoniums which have contracted or imperfect leaves have contracted flowers: the difference between “Brilliant” and its parent “Tom Thumb” is a good instance of this. It may be suspected that the curious case described by Risso (25/30. ‘Annales du Museum’ tome 20 page 188.), of a variety of the Orange which produces on the young shoots rounded leaves with winged petioles, and afterwards elongated leaves on long but wingless petioles, is connected with the remarkable change in form and nature which the fruit undergoes during its development.
In the following instance we have the colour and the form of the petals apparently correlated, and both dependent on the nature of the season. An observer, skilled in the subject, writes (25/31. ‘Gardener’s Chronicle’ 1843 page 877.), “I noticed, during the year 1842, that every Dahlia of which the colour had any tendency to scarlet, was deeply notched–indeed, to so great an extent as to give the petals the appearance of a saw; the indentures were, in some instances, more than a quarter of an inch deep.” Again, Dahlias which have their petals tipped with a different colour from the rest of the flower are very inconstant, and during certain years some, or even all the flowers, become uniformly coloured; and it has been observed with several varieties (25/32. Ibid 1845 page 102.) that when this happens the petals grow much elongated and lose their proper shape. This, however, may be due to reversion, both in colour and form, to the aboriginal species.
In this discussion on correlation, we have hitherto treated of cases in which we can partly understand the bond of connection; but I will now give cases in which we cannot even conjecture, or can only very obscurely see, the nature of the bond. Isidore Geoffroy Saint-Hilaire, in his work on Monstrosities, insists (25/33. ‘Hist. des Anomalies’ tome 3 page 402. See also M. Camille Dareste ‘Recherches sur les Conditions’ etc. 1863 pages 16, 48.), “que certaines anomalies coexistent rarement entr’elles, d’autres frequemment, d’autres enfin presque constamment, malgre la difference tres-grande de leur nature, et quoiqu’elles puissent paraitre COMPLETEMENT INDEPENDANTES les unes des autres.” We see something analogous in certain diseases: thus in a rare affection of the renal capsules (of which the functions are unknown), the skin becomes bronzed; and in hereditary syphilis, as I hear from Sir J. Paget, both the milk and the second teeth assume a peculiar and characteristic form. Professor Rolleston, also, informs me that the incisor teeth are sometimes furnished with a vascular rim in correlation with intra-pulmonary deposition of tubercles. In other cases of phthisis and of cyanosis the nails and finger- ends become clubbed like acorns. I believe that no explanation has been offered of these and of many other cases of correlated disease.
What can be more curious and less intelligible than the fact previously given, on the authority of Mr. Tegetmeier, that young pigeons of all breeds, which when mature have white, yellow, silver-blue, or dun-coloured plumage, come out of the egg almost naked; whereas pigeons of other colours when first born are clothed with plenty of down? White Pea-fowls, as has been observed both in England and France (25/34. Rev. E.S. Dixon ‘Ornamental Poultry’ 1848 page 111; Isidore Geoffroy ‘Hist. Anomalies’ tome 1 page 211.), and as I have myself seen, are inferior in size to the common coloured kind; and this cannot be accounted for by the belief that albinism is always accompanied by constitutional weakness; for white or albino moles are generally larger than the common kind.
To turn to more important characters: the niata cattle of the Pampas are remarkable from their short foreheads, upturned muzzles, and curved lower jaws. In the skull the nasal and premaxillary bones are much shortened, the maxillaries are excluded from any junction with the nasals, and all the bones are slightly modified, even to the plane of the occiput. From the analogous case of the dog, hereafter to be given, it is probable that the shortening of the nasal and adjoining bones is the proximate cause of the other modifications in the skull, including the upward curvature of the lower jaw, though we cannot follow out the steps by which these changes have been effected.
Polish fowls have a large tuft of feathers on their heads; and their skulls are perforated by numerous holes, so that a pin can be driven into the brain without touching any bone. That this deficiency of bone is in some way connected with the tuft of feathers is clear from tufted ducks and geese likewise having perforated skulls. The case would probably be considered by some authors as one of balancement or compensation. In the chapter on Fowls, I have shown that with Polish fowls the tuft of feathers was probably at first small; by continued selection it became larger, and then rested on a fibrous mass; and finally, as it became still larger, the skull itself became more and more protuberant until it acquired its present extraordinary structure. Through correlation with the protuberance of the skull, the shape and even the relative connection of the premaxillary and nasal bones, the shape of the orifice of the nostrils, the breadth of the frontal bone, the shape of the post-lateral processes of the frontal and squamosal bones, and the direction of the bony cavity of the ear, have all been modified. The internal configuration of the skull and the whole shape of the brain have likewise been altered in a truly marvellous manner.
After this case of the Polish fowl it would be superfluous to do more than refer to the details previously given on the manner in which the changed form of the comb has affected the skull, in various breeds of the fowl, causing by correlation crests, protuberances, and depressions on its surface.
With our cattle and sheep the horns stand in close connection with the size of the skull, and with the shape of the frontal bones; thus Cline (25/35. ‘On the Breeding of Domestic Animals’ 1829 page 6.) found that the skull of a horned ram weighed five times as much as that of a hornless ram of the same age. When cattle become hornless, the frontal bones are “materially diminished in breadth towards the poll;” and the cavities between the bony plates “are not so deep, nor do they extend beyond the frontals.” (25/36. Youatt on ‘Cattle’ 1834 page 283.)
It may be well here to pause and observe how the effects of correlated variability, of the increased use of parts, and of the accumulation of so- called spontaneous variations through natural selection, are in many cases inextricably commingled. We may borrow an illustration from Mr. Herbert Spencer, who remarks that, when the Irish elk acquired its gigantic horns, weighing above one hundred pounds, numerous co-ordinated changes of structure would have been indispensable,–namely, a thickened skull to carry the horns; strengthened cervical vertebrae, with strengthened ligaments; enlarged dorsal vertebrae to support the neck, with powerful fore-legs and feet; all these parts being supplied with proper muscles, blood-vessels, and nerves. How then could these admirably co-ordinated modifications of structure have been acquired? According to the doctrine which I maintain, the horns of the male elk were slowly gained through sexual selection,–that is, by the best-armed males conquering the worse-armed, and leaving a greater number of descendants. But it is not at all necessary that the several parts of the body should have simultaneously varied. Each stag presents individual characteristics, and in the same district those which had slightly heavier horns, or stronger necks, or stronger bodies, or were the most courageous, would secure the greater number of does, and consequently have a greater number of offspring. The offspring would inherit, in a greater or less degree, these same qualities, would occasionally intercross with one another, or with other individuals varying in some favourable manner; and of their offspring, those which were