The Variation of Animals and Plants under Domestication Volume I Part 36

Supernumerary fingers and toes are eminently liable, as various authors have insisted, to be inherited. Polydactylism graduates (12/27. Vrolik has discussed this point at full length in a work published in Dutch, from which Sir J. Paget has kindly translated for me pa.s.sages. See, also, Isidore Geoffroy St. Hilaire 'Hist. des Anomalies' 1832 tome 1 page 684.) by multifarious steps from a mere cutaneous appendage, not including any bone, to a double hand. But an additional digit, supported on a metacarpal bone, and furnished with all the proper muscles, nerves, and vessels, is sometimes so perfect, that it escapes detection, unless the fingers are actually counted. Occasionally there are several supernumerary digits; but usually only one, making the total number six. This one may be attached to the inner or outer margin of the hand, representing either a thumb or little finger, the latter being the more frequent. Generally, through the law of correlation, both hands and both feet are similarly affected. Dr.

Burt Wilder has tabulated (12/28. 'Ma.s.sachusetts Medical Society' volume 2 No. 3; and 'Proc. Boston Soc. of Nat. Hist.' volume 14 1871 page 154.) a large number of cases, and finds that supernumerary digits are more common on the hands than on the feet, and that men are affected oftener than women. Both these facts can be explained on two principles which seem generally to hold good; firstly, that of two parts, the more specialised one is the more variable, and the arm is more highly specialised than the leg; and secondly that male animals are more variable than females.

The presence of a greater number of digits than five is a great anomaly, for this number is not normally exceeded by any existing mammal, bird, or reptile. Nevertheless, supernumerary digits are strongly inherited; they have been transmitted through five generations; and in some cases, after disappearing for one, two, or even three generations, have reappeared through reversion. These facts are rendered, as Professor Huxley has observed, more remarkable from its being known in most cases that the affected person has not married one similarly affected. In such cases the child of the fifth generation would have only 1-32nd part of the blood of his first sedigitated ancestor. Other cases are rendered remarkable by the affection gathering force, as Dr. Struthers has shown, in each generation, though in each the affected person married one not affected; moreover, such additional digits are often amputated soon after birth, and can seldom have been strengthened by use. Dr. Struthers gives the following instance: in the first generation an additional digit appeared on one hand; in the second, on both hands; in the third, three brothers had both hands, and one of the brothers a foot affected; and in the fourth generation all four limbs were affected. Yet we must not over-estimate the force of inheritance. Dr. Struthers a.s.serts that cases of non-inheritance and of the first appearance of additional digits in unaffected families are much more frequent than cases of inheritance. Many other deviations of structure, of a nature almost as anomalous as supernumerary digits, such as deficient phalanges (12/29. Dr. J.W. Ogle gives a case of the inheritance of deficient phalanges during four generations. He adds references to various recent papers on inheritance 'Brit. and For. Med.-Chirurg. Review' April 1872.), thickened joints, crooked fingers, etc., are, in like manner, strongly inherited, and are equally subject to intermission, together with reversion, though in such cases there is no reason to suppose that both parents had been similarly affected. (12/30. For these several statements see Dr. Struthers 'Edinburgh New Phil. Journal' July 1863 especially on intermissions in the line of descent. Prof. Huxley 'Lectures on our Knowledge of Organic Nature' 1863 page 97. With respect to inheritance, see Dr. Prosper Lucas 'L'Heredite Nat.' tome 1 page 325. Isid. Geoffroy 'Anom.'

tome 1 page 701. Sir A. Carlisle in 'Phil. Transact.' 1814 page 94. A.

Walker on 'Intermarriage' 1838 page 140 gives a case of five generations; as does Mr. Sedgwick in 'Brit. and Foreign Medico-Chirurg. Review' April 1863 page 462. On the inheritance of other anomalies in the extremities see Dr. H. Dobell in volume 46 of Medico-Chirurg. Transactions 1863; also Mr.

Sedgwick in op. cit. April 1863 page 460. With respect to additional digits in the negro see Prichard 'Physical History of Mankind.' Dr. Dieffenbach 'Jour. Royal Geograph. Soc.' 1841 page 208 says this anomaly is not uncommon with the Polynesians of the Chatham Islands; and I have heard of several cases with Hindus and Arabs.)

Additional digits have been observed in negroes as well as in other races of man, and in several of the lower animals, and have been inherited. Six toes have been described on the hind feet of the newt (Salamandra cristata), and are said to have occurred with the frog. It deserves notice, that the six-toed newt, though adult, preserved some of its larval characters; for part of the hyoidal apparatus, which is properly absorbed during the act of metamorphosis, was retained. It is also remarkable that in the case of man various structures in an embryonic or arrested state of development, such as a cleft-palate, bifid uterus, etc., are often accompanied by polydactylism. (12/31. Meckel and Isid G. St. Hilaire insist on this fact. See also M. A. Roujou 'Sur quelques a.n.a.logies du Type Humain'

page 61 published, I believe, in the 'Journal of the Anthropolog. Soc. of Paris' January 1872.) Six toes on the hinder feet are known to have been inherited for three generations of cats. In several breeds of the fowl the hinder toe is double, and is generally transmitted truly, as is well shown when Dorkings are crossed with common four-toed breeds (12/32. 'The Poultry Chronicle' 1854 page 559.) With animals which have properly less than five digits, the number is sometimes increased to five, especially on the front legs, though rarely carried beyond that number; but this is due to the development of a digit already existing in a more or less rudimentary state. Thus, the dog has properly four toes behind, but in the larger breeds a fifth toe is commonly, though not perfectly, developed. Horses, which properly have one toe alone fully developed with rudiments of the others, have been described with each foot bearing two or three small separate hoofs: a.n.a.logous facts have been noticed with cows, sheep, goats, and pigs. (12/33. The statements in this paragraph are taken from Isidore Geoffroy St. Hilaire 'Hist. des Anomalies' tome 1 pages 688-693. Mr.

Goodman gives, 'Phil. Soc. of Cambridge' November 25, 1872 the case of a cow with three well developed toes on each hind limb, besides the ordinary rudiments; and her calf by an ordinary bull had extra digits. This calf also bore two calves having extra digits.)

There is a famous case described by Mr. White of a child, three years old, with a thumb double from the first joint. He removed the lesser thumb, which was furnished with a nail; but to his astonishment it grew again and reproduced a nail. The child was then taken to an eminent London surgeon, and the newly-grown thumb was removed by its socket-joint, but again it grew and reproduced a nail. Dr. Struthers mentions a case of the partial regrowth of an additional thumb, amputated when a child was three months old; and the late Dr. Falconer communicated to me an a.n.a.logous instance. In the last edition of this work I also gave a case of the regrowth of a supernumerary little-finger after amputation; but having been informed by Dr. Bachmaier that several eminent surgeons expressed, at a meeting of the Anthropological Society of Munich, great doubt about my statements, I have made more particular inquiries. The full information thus gained, together with a tracing of the hand in its present state, has been laid before Sir J. Paget, and he has come to the conclusion that the degree of regrowth in this case is not greater than sometimes occurs with normal bones, especially with the humerus, when amputated at an early age. He further does not feel fully satisfied about the facts recorded by Mr. White. This being so, it is necessary for me to withdraw the view which I formerly advanced, with much hesitation, chiefly on the ground of the supposed regrowth of additional digits, namely, that their occasional development in man is a case of reversion to a lowly, organised progenitor provided with more than five digits.]

I may here allude to a cla.s.s of facts closely allied to, but somewhat different from, ordinary cases of inheritance. Sir H. Holland (12/34.

'Medical Notes and Reflections' 1839 pages 24, 34. See also Dr. P. Lucas 'L'Hered. Nat.' tome 2 page 33.) states that brothers and sisters of the same family are frequently affected, often at about the same age, by the same peculiar disease, not known to have previously occurred in the family.

He specifies the occurrence of diabetes in three brothers under ten years old; he also remarks that children of the same family often exhibit in common infantile diseases, the same peculiar symptoms. My father mentioned to me the case of four brothers who died between the ages of sixty and seventy, in the same highly peculiar comatose state. An instance has already been given of supernumerary digits appearing in four children out of six in a previously unaffected family. Dr. Devay states (12/35. 'Du Danger des Mariages Consanguins' 2nd edition 1862 page 103.) that two brothers married two sisters, their first-cousins, none of the four nor any relation being an albino; but the seven children produced from this double marriage were all perfect albinoes. Some of these cases, as Mr. Sedgwick (12/36. 'British and Foreign Medico-Chirurg. Review' July 1863 pages 183, 189.) has shown, are probably the result of reversion to a remote ancestor, of whom no record had been preserved; and all these cases are so far directly connected with inheritance that no doubt the children inherited a similar const.i.tution from their parents, and, from being exposed to nearly similar conditions of life, it is not surprising that they should be affected in the same manner and at the same period of life.

Most of the facts. .h.i.therto given have served to ill.u.s.trate the force of inheritance, but we must now consider cases grouped as well as the subject allows into cla.s.ses, showing how feeble, capricious, or deficient the power of inheritance sometimes is. When a new peculiarity first appears, we can never predict whether it will be inherited. If both parents from their birth present the same peculiarity, the probability is strong that it will be transmitted to at least some of their offspring. We have seen that variegation is transmitted much more feebly by seed, taken from a branch which had become variegated through bud-variation, than from plants which were variegated as seedlings. With most plants the power of transmission notoriously depends on some innate capacity in the individual: thus Vilmorin (12/37. Verlot 'La Product. des Varietes' 1865 page 32.) raised from a peculiarly coloured balsam some seedlings, which all resembled their parent; but of these seedlings some failed to transmit the new character, whilst others transmitted it to all their descendants during several successive generations. So again with a variety of the rose, two plants alone out of six were found by Vilmorin to be capable of transmitting the desired character; numerous a.n.a.logous cases could be given.

[The weeping or pendulous growth of trees is strongly inherited in some cases, and, without any a.s.signable reason, feebly in other cases. I have selected this character as an instance of capricious inheritance, because it is certainly not proper to the parent-species, and because, both s.e.xes being borne on the same tree, both tend to transmit the same character.

Even supposing that there may have been in some instances crossing with adjoining trees of the same species, it is not probable that all the seedlings would have been thus affected. At Moccas Court there is a famous weeping oak; many of its branches "are 30 feet long, and no thicker in any part of this length than a common rope:" this tree transmits its weeping character, in a greater or less degree, to all its seedlings; some of the young oaks being so flexible that they have to be supported by props; others not showing the weeping tendency till about twenty years old.

(12/38. Loudon's 'Gardener's Mag.' volume 12 1836 page 368.) Mr. Rivers fertilised, as he informs me, the flowers of a new Belgian weeping thorn (Crataegus oxyacantha) with pollen from a crimson not-weeping variety, and three young trees, "now six or seven years old, show a decided tendency to be pendulous, but as yet are not so much so as the mother-plant." According to Mr. MacNab (12/39. Verlot 'La Product. des Varietes' 1865 page 94.), seedlings from a magnificent weeping birch (Betula alba), in the Botanic Garden at Edinburgh, grew for the first ten or fifteen years upright, but then all became weepers like their parent. A peach with pendulous branches, like those of the weeping willow, has been found capable of propagation by seed. (12/40. Bronn 'Geschichte der Natur' b. 2 s. 121. Mr. Meehan makes a similar statement in 'Proc. Nat. of Philadelphia' 1872 page 235.) Lastly, a weeping or rather a prostrate yew (Taxus baccata) was found in a hedge in Shrops.h.i.+re; it was a male, but one branch bore female flowers, and produced berries; these, being sown, produced seventeen trees all of which had exactly the same peculiar habit with the parent-tree. (12/41. Rev. W.A.

Leighton 'Flora of Shrops.h.i.+re' page 497; and Charlesworth 'Mag. of Nat.

Hist.' volume 1 1837 page 30. I possess prostrate trees produced from these seeds.)

These facts, it might have been thought, would have been sufficient to render it probable that a pendulous habit would in all cases be strictly inherited. But let us look to the other side. Mr. MacNab (12/42. Verlot op.

cit. page 93.) sowed seeds of the weeping beech (f.a.gus sylvatica), but succeeded in raising only common beeches. Mr. Rivers, at my request, raised a number of seedlings from three distinct varieties of weeping elm; and at least one of the parent-trees was so situated that it could not have been crossed by any other elm; but none of the young trees, now about a foot or two in height, show the least signs of weeping. Mr. Rivers formerly sowed above twenty thousand seeds of the weeping ash (Fraxinus excelsior), and not a single seedling was in the least degree pendulous: in Germany, M.

Borchmeyer raised a thousand seedlings, with the same result. Nevertheless, Mr. Anderson, of the Chelsea Botanic Garden, by sowing seed from a weeping ash, which was found before the year 1780, in Cambridges.h.i.+re, raised several pendulous trees. (12/43. For these several statements see Loudon's 'Gard. Magazine' volume 10 1834 pages 408, 180; and volume 9 1833 page 597.) Professor Henslow also informs me that some seedlings from a female weeping ash in the Botanic Garden at Cambridge were at first a little pendulous, but afterwards became quite upright: it is probable that this latter tree, which transmits to a certain extent its pendulous habit, was derived by a bud from the same original Cambridges.h.i.+re stock; whilst other weeping ashes may have had a distinct origin. But the crowning case, communicated to me by Mr. Rivers, which shows how capricious is the inheritance of a pendulous habit, is that a variety of another species of ash (F. lentiscifolia), now about twenty years old, which was formerly pendulous, "has long lost this habit, every shoot being remarkably erect; but seedlings formerly raised from it were perfectly prostrate, the stems not rising more than two inches above the ground." Thus the weeping variety of the common ash, which has been extensively propagated by buds during a long period, did not with Mr. Rivers, transmit its character to one seedling out of above twenty thousand; whereas the weeping variety of a second species of ash, which could not, whilst grown in the same garden, retain its own weeping character, transmitted to its character the pendulous habit in excess!

Many a.n.a.logous facts could be given, showing how apparently capricious is the principle of inheritance. All the seedlings from a variety of the Barberry (B. vulgaris) with red leaves inherited the same character; only about one-third of the seedlings of the copper Beech (f.a.gus sylvatica) had purple leaves. Not one out of a hundred seedlings of a variety of the Cerasus padus, with yellow fruit, bore yellow fruit: one-twelfth of the seedlings of the variety of Cornus mascula, with yellow fruit, came true (12/44. These statements are taken from Alph. De Candolle 'Bot. Geograph.'

page 1083.): and lastly, all the trees raised by my father from a yellow- berried holly (Ilex aquifolium), found wild, produced yellow berries.

Vilmorin (12/45. Verlot op. cit. page 38.) observed in a bed of Saponaria calabrica an extremely dwarf variety, and raised from it a large number of seedlings; some of these partially resembled their parent, and he selected their seed; but the grandchildren were not in the least dwarfed: on the other hand, he observed a stunted and bushy variety of Tagetes signata growing in the midst of the common varieties by which it was probably crossed; for most of the seedlings raised from this plant were intermediate in character, only two perfectly resembling their parent; but seed saved from these two plants reproduced the new variety so truly, that hardly any selection has since been necessary.

Flowers transmit their colour truly, or most capriciously. Many annuals come true: thus I purchased German seeds of thirty-four named sub-varieties of one RACE of ten-week stocks (Matthiola annua), and raised a hundred and forty plants, all of which, with the exception of a single plant, came true. In saying this, however, it must be understood that I could distinguish only twenty kinds out of the thirty-four named sub-varieties; nor did the colour of the flower always correspond with the name affixed to the packet; but I say that they came true, because in each of the thirty- six short rows every plant was absolutely alike, with the one single exception. Again, I procured packets of German seed of twenty-five named varieties of common and quilled asters, and raised a hundred and twenty- four plants; of these, all except ten were true in the above limited sense; and I considered even a wrong shade of colour as false.

It is a singular circ.u.mstance that white varieties generally transmit their colour much more truly than any other variety. This fact probably stands in close relation with one observed by Verlot (12/46. Op. cit. page 59.), namely, that flowers which are normally white rarely vary into any other colour. I have found that the white varieties of Delphinium consolida and of the Stock are the truest. It is, indeed, sufficient to look through a nurseryman's seed-list, to see the large number of white varieties which can be propagated by seed. The several coloured varieties of the sweet-pea (Lathyrus odoratus) are very true; but I hear from Mr. Masters, of Canterbury, who has particularly attended to this plant, that the white variety is the truest. The hyacinth, when propagated by seed, is extremely inconstant in colour, but "white hyacinths almost always give by seed white-flowered plants" (12/47. Alph. De Candolle 'Geograph. Bot.' page 1082.); and Mr. Masters informs me that the yellow varieties also reproduce their colour, but of different shades. On the other hand, pink and blue varieties, the latter being the natural colour, are not nearly so true: hence, as Mr. Masters has remarked to me, "we see that a garden variety may acquire a more permanent habit than a natural species;" but it should have been added, that this occurs under cultivation, and therefore under changed conditions.

With many flowers, especially perennials, nothing can be more fluctuating than the colour of the seedlings, as is notoriously the case with verbenas, carnations, dahlias, cinerarias, and others. (12/48. See 'Cottage Gardener'

April 10, 1860 page 18 and September 10, 1861 page 456; 'Gardener's Chronicle' 1845 page 102.) I sowed seed of twelve named varieties of Snapdragon (Antirrhinum majus), and utter confusion was the result. In most cases the extremely fluctuating colour of seedling plants is probably in chief part due to crosses between differently-coloured varieties during previous generations. It is almost certain that this is the case with the polyanthus and coloured primrose (Primula veris and vulgaris), from their reciprocally dimorphic structure (12/49. Darwin in 'Journal of Proc. Linn.

Soc. Bot.' 1862 page 94.); and these are plants which florists speak of as never coming true by seed: but if care be taken to prevent crossing, neither species is by any means very inconstant, in colour; thus I raised twenty-three plants from a purple primrose, fertilised by Mr. J. Scott with its pollen, and eighteen came up purple of different shades, and only five reverted to the ordinary yellow colour: again, I raised twenty plants from a bright-red cowslip, similarly treated by Mr. Scott, and every one perfectly resembled its parent in colour, as likewise did, with the exception of a single plant, 72 grandchildren. Even with the most variable flowers, it is probable that each delicate shade of colour might be permanently fixed so as to be transmitted by seed, by cultivation in the same soil, by long-continued selection, and especially by the prevention of crosses. I infer this from certain annual larkspurs (Delphinium consolida and ajacis), of which common seedlings present a greater diversity of colour than any other plant known to me; yet on procuring seed of five named German varieties of D. consolida, only nine plants out of ninety-four were false; and the seedlings of six varieties of D. ajacis were true in the same manner and degree as with the stocks above described. A distinguished botanist maintains that the annual species of Delphinium are always self-fertilised; therefore I may mention that thirty-two flowers on a branch of D. consolida, enclosed in a net, yielded twenty-seven capsules, with an average of 17.2 seed in each; whilst five flowers, under the same net, which were artificially fertilised, in the same manner as must be effected by bees during their incessant visits, yielded five capsules with an average of 35.2 fine seed; and this shows that the agency of insects is necessary for the full fertility of this plant. a.n.a.logous facts could be given with respect to the crossing of many other flowers, such as carnations, etc., of which the varieties fluctuate much in colour.

As with flowers, so with our domesticated animals, no character is more variable than colour, and probably in no animal more so than with the horse. Yet, with a little care in breeding, it appears that races of any colour might soon be formed. Hofacker gives the result of matching two hundred and sixteen mares of four different colours with like-coloured stallions, without regard to the colour of their ancestors; and of the two hundred and sixteen colts born, eleven alone failed to inherit the colour of their parents: Autenrieth and Ammon a.s.sert that, after two generations, colts of a uniform colour are produced with certainty. (12/50. Hofacker 'Ueber die Eigenschaften' etc. s. 10.)]

In a few rare cases peculiarities fail to be inherited, apparently from the force of inheritance being too strong. I have been a.s.sured by breeders of the canary-bird that to get a good jonquil-coloured bird it does not answer to pair two jonquils, as the colour then comes out too strong, or is even brown; but this statement is disputed by other breeders. So again, if two crested canaries are paired, the young birds rarely inherit this character (12/51. Bechstein 'Naturgesch. Deutschlands' b. 4 s. 462. Mr. Brent, a great breeder of canaries, informs me that he believes that these statements are correct.): for in crested birds a narrow s.p.a.ce of bare skin is left on the back of the head, where the feathers are up-turned to form the crest, and, when both parents are thus characterised, the bareness becomes excessive, and the crest itself fails to be developed. Mr. Hewitt, speaking of Laced Sebright Bantams, says (12/52. 'The Poultry Book' by W.B.

Tegetmeier 1866 page 245.) that, "why this should be so I know not, but I am confident that those that are best laced frequently produce offspring very far from perfect in their markings, whilst those exhibited by myself, which have so often proved successful, were bred from the union of heavily- laced birds with those that were scarcely sufficiently laced."

It is a singular fact that, although several deaf-mutes often occur in the same family, and though their cousins and other relations are often in the same condition, yet their parents are rarely deaf-mutes. To give a single instance: not one scholar out of 148, who were at the same time in the London Inst.i.tution, was the child of parents similarly affected. So again, when a male or female deaf-mute marries a sound person, their children are most rarely affected: in Ireland, out of 203 children thus produced one alone was mute. Even when both parents have been deaf-mutes, as in the case of forty-one marriages in the United States and of six in Ireland, only two deaf and dumb children were produced. Mr. Sedgwick (12/53. 'British and Foreign Med.-Chirurg. Review' July 1861 pages 200-204. Mr. Sedgwick has given such full details on this subject, with ample references, that I need refer to no other authorities), in commenting on this remarkable and fortunate failure in the power of transmission in the direct line, remarks that it may possibly be owing to "excess having reversed the action of some natural law in development." But it is safer in the present state of our knowledge to look at the whole case as simply unintelligible.

Although many congenital monstrosities are inherited, of which examples have already been given, and to which may be added the lately recorded case of the transmission during a century of hare-lip with a cleft-palate in the writer's own family (12/54. Mr. Sproule in 'British Medical Journal' April 18, 1863.) yet other malformations are rarely or never inherited. Of these latter cases, many are probably due to injuries in the womb or egg, and would come under the head of non-inherited injuries or mutilations. With plants, a long catalogue of inherited monstrosities of the most serious and diversified nature could easily be given; and with plants, there is no reason to suppose that monstrosities are caused by direct injuries to the seed or embryo.

With respect to the inheritance of structures mutilated by injuries or altered by disease, it was until lately difficult to come to any definite conclusion. Some mutilations have been practised for a vast number of generations without any inherited result. G.o.dron remarks (12/55. 'De l'Espece' tome 2 1859 page 299.) that different races of man have from time immemorial knocked out their upper incisors, cut off joints of their fingers, made holes of immense size through the lobes of their ears or through their nostrils, tatooed themselves, made deep gashes in various parts of their bodies, and there is no reason to suppose that these mutilations have ever been inherited. (12/56. Nevertheless Mr. Wetherell states, 'Nature' December 1870 page 168, that when he visited fifteen years ago the Sioux Indians, he was informed "by a physician, who has pa.s.sed much of his time with these tribes, that some times a child was born with these marks. This was confirmed by the U.S. Government Indian Agent.") Adhesions due to inflammation and pits from the small-pox (and formerly many consecutive generations must have been thus pitted) are not inherited. With respect to Jews, I have been a.s.sured by three medical men of the Jewish faith that circ.u.mcision, which has been practised for so many ages, has produced no inherited effect. Blumenbach, however, a.s.serts (12/57.

'Philosoph. Mag.' volume 4 1799 page 5.) that Jews are often born in Germany in a condition rendering circ.u.mcision difficult, so that a name is given them signifying "born circ.u.mcised;" and Professor Preyer informs me that this is the case in Bonn, such children being considered the special favourites of Jehovah. I have also heard from Dr. A. Newman, of Guy's Hospital, of the grandson of a circ.u.mcised Jew, the father not having been circ.u.mcised, in a similar condition. But it is possible that all these cases may be accidental coincidence, for Sir J. Paget has seen five sons of a lady and one son of her sister with adherent prepuces; and one of these boys was affected in a manner "which might be considered like that commonly produced by circ.u.mcision;" yet there was no suspicion of Jewish blood in the family of these two sisters. Circ.u.mcision is practised by Mahomedans, but at a much later age than by Jews; and Dr. Riedel, a.s.sistant Resident in North Celebes, writes to me that the boys there go naked until from six to ten years old; and he has observed that many of them, though not all, have their prepuces much reduced in length, and this he attributes to the inherited effects of the operation. In the vegetable kingdom oaks and other trees have borne galls from primeval times, yet they do not produce inherited excrescences; and many other such facts could be adduced.

Notwithstanding the above several negative cases, we now possess conclusive evidence that the effects of operations are sometimes inherited. Dr. Brown- Sequard (12/58. 'Proc. Royal Soc.' volume 10 page 297. 'Communication to the Brit. a.s.soc.' 1870. 'The Lancet' January 1875 page 7. The extracts are from this last paper. It appears that Obersteiner 'Stricker's Med.

Jahrbucher' 1875 No. 2 has confirmed Brown-Sequard's observations.) gives the following summary of his observations on guinea-pigs; and this summary is so important that I will quote the whole:--

["1st. Appearance of epilepsy in animals born of parents having been rendered epileptic by an injury to the spinal cord.

"2nd. Appearance of epilepsy also in animals born of parents having been rendered epileptic by the section of the sciatic nerve.

"3rd. A change in the shape of the ear in animals born of parents in which such a change was the effect of a division of the cervical sympathetic nerve.

"4th. Partial closure of the eyelids in animals born of parents in which that state of the eyelids had been caused either by the section of the cervical sympathetic nerve or the removal of the superior cervical ganglion.

"5th. Exophthalmia in animals born of parents in which an injury to the restiform body had produced that protrusion of the eyeball. This interesting fact I have witnessed a good many times, and I have seen the transmission of the morbid state of the eye continue through four generations. In these animals, modified by heredity, the two eyes generally protruded, although in the parents usually only one showed exophthalmia, the lesion having been made in most cases only on one of the corpora restiformia.

"6th. Haematoma and dry gangrene of the ears in animals born of parents in which these ear-alterations had been caused by an injury to the restiform body near the nib of the calamus.

"7th. Absence of two toes out of the three of the hind leg, and sometimes of the three, in animals whose parents had eaten up their hind-leg toes which had become anaesthetic from a section of the sciatic nerve alone, or of that nerve and also of the crural. Sometimes, instead of complete absence of the toes, only a part of one or two or three was missing in the young, although in the parent not only the toes but the whole foot was absent (partly eaten off, partly destroyed by inflammation, ulceration, or gangrene).

"8th. Appearance of various morbid states of the skin and hair of the neck and face in animals born of parents having had similar alterations in the same parts, as effects of an injury to the sciatic nerve."]

It should be especially observed that Brown-Sequard has bred during thirty years many thousand guinea-pigs from animals which had not been operated upon, and not one of these manifested the epileptic tendency. Nor has he ever seen a guinea-pig born without toes, which was not the offspring of parents which had gnawed off their own toes owing to the sciatic nerve having been divided. Of this latter fact thirteen instances were carefully recorded, and a greater number were seen; yet Brown-Sequard speaks of such cases as one of the rarer forms of inheritance. It is a still more interesting fact--

["That the sciatic nerve in the congenitally toeless animal has inherited the power of pa.s.sing through all the different morbid states which have occurred in one of its parents from the time of the division till after its reunion with the peripheric end. It is not therefore simply the power of performing an action which is inherited, but the power of performing a whole series of actions, in a certain order."]

In most of the cases of inheritance recorded by Brown-Sequard only one of the two parents had been operated upon and was affected. He concludes by expressing his belief that "what is transmitted is the morbid state of the nervous system," due to the operation performed on the parents.

With the lower animals Dr. Prosper Lucas has collected a long list of inherited injuries. A few instances will suffice. A cow lost a horn from an accident with consequent suppuration, and she produced three calves which were hornless on the same side of the head. With the horse, there seems hardly a doubt that exostoses on the legs, caused by too much travelling on hard roads, are inherited. Blumenbach records the case of a man who had his little finger on the right hand almost cut off, and which in consequence grew crooked, and his sons had the same finger on the same hand similarly crooked. A soldier, fifteen years before his marriage, lost his left eye from purulent ophthalmia, and his two sons were microphthalmic on the same side. (12/59. This last case is quoted by Mr. Sedgwick in 'British and Foreign Medico-Chirurg. Review' April, 1861 page 484. For Blumenbach see above-cited paper. See also Dr. P. Lucas 'Traite de l'Hered. Nat.' tome 2 page 492. Also 'Transact. Linn. Soc.' volume 9 page 323. Some curious cases are given by Mr. Baker in the 'Veterinary' volume 13 page 723. Another curious case is given in the 'Annales des Scienc. Nat.' 1st series, tome 11 page 324.) In all cases in which a parent has had an organ injured on one side, and two or more of the offspring are born with the same organ affected on the same side, the chances against mere coincidence are almost infinitely great. Even when only a single child is born having exactly the same part of the body affected as that of his injured parent, the chances against coincidence are great; and Professor Rolleston has given me two such cases which have fallen under his own observation,--namely of two men, one of whom had his knee and the other his cheek severely cut, and both had children born with exactly the same spot marked or scarred. Many instances have been recorded of cats, dogs, and horses, which have had their tails, legs, etc., amputated or injured, producing offspring with the same parts ill-formed; but as it is not very rare for similar malformations to appear spontaneously, all such cases may be due to coincidence. It is, however, an argument on the other side that "under the old excise laws the shepherd-dog was only exempt from tax when without a tail, and for this reason it was always removed" (12/60. 'The Dog' by Stonehenge 1867 page 118.); and there still exist breeds of the shepherd-dog which are always born dest.i.tute of a tail. Finally, it must be admitted, more especially since the publication of Brown-Sequard's observations, that the effects of injuries, especially when followed by disease, or perhaps exclusively when thus followed, are occasionally inherited. (12/61. The Mot-mot habitually bites the barbs off the middle part of the two central tail-feathers, and as the barbs are congenitally somewhat reduced on the same part of these feathers, it seems extremely probable, as Mr. Salvin remarks, 'Proc. Zoolog. Soc.' 1873 page 429, that this is due to the inherited effects of long-continued mutilation.)

CAUSES OF NON-INHERITANCE.

A large number of cases of non-inheritance are intelligible on the principle, that a strong tendency to inheritance does exist, but that it is overborne by hostile or unfavourable conditions of life. No one would expect that our improved pigs, if forced during several generations to travel about and root in the ground for their own subsistence, would transmit, as truly as they now do their short muzzles and legs, and their tendency to fatten. Dray-horses a.s.suredly would not long transmit their great size and ma.s.sive limbs, if compelled to live on a cold, damp mountainous region; we have indeed evidence of such deterioration in the horses which have run wild on the Falkland Islands. European dogs in India often fail to transmit their true character. Our sheep in tropical countries lose their wool in a few generations. There seems also to be a close relation between certain peculiar pastures and the inheritance of an enlarged tail in fat-tailed sheep, which form one of the most ancient breeds in the world. With plants, we have seen that tropical varieties of maize lose their proper character in the course of two or three generations, when cultivated in Europe; and conversely so it is with European varieties cultivated in Brazil. Our cabbages, which here come so true by seed, cannot form heads in hot countries. According to Carriere (12/62. 'Production et Fixation des Varietes' 1865 page 72.) the purple- leafed beech and barberry transmit their character by seed far less truly in certain districts than in others. Under changed circ.u.mstances, periodical habits of life soon fail to be transmitted, as the period of maturity in summer and winter wheat, barley, and vetches. So it is with animals: for instance, a person, whose statement I can trust, procured eggs of Aylesbury ducks from that town, where they are kept in houses and are reared as early as possible for the London market; the ducks bred from these eggs in a distant part of England, hatched their first brood on January 24th, whilst common ducks, kept in the same yard and treated in the same manner, did not hatch till the end of March; and this shows that the period of hatching was inherited. But the grandchildren of these Aylesbury ducks completely lost their habit of early incubation, and hatched their eggs at the same time with the common ducks of the same place.

Many cases of non-inheritance apparently result from the conditions of life continually inducing fresh variability. We have seen that when the seeds of pears, plums, apples, etc., are sown, the seedlings generally inherit some degree of family likeness. Mingled with these seedlings, a few, and sometimes many, worthless, wild-looking plants commonly appear, and their appearance may be attributed to the principle of reversion. But scarcely a single seedling will be found perfectly to resemble the parent-form; and thus may be accounted for by constantly recurring variability induced by the conditions of life. I believe in this, because it has been observed that certain fruit-trees truly propagate their kind whilst growing on their own roots; but when grafted on other stocks, and by this process their natural state is manifestly affected, they produce seedlings which vary greatly, departing from the parental type in many characters. (12/63.

Downing 'Fruits of America' page 5: Sageret 'Pom. Phys.' pages 43, 72.) Metzger, as stated in the ninth chapter, found that certain kinds of wheat brought from Spain and cultivated in Germany, failed during many years to reproduce themselves truly; but at last, when accustomed to their new conditions, they ceased to be variable,--that is, they became amenable to the power of inheritance. Nearly all the plants which cannot be propagated with any approach to certainty by seed, are kinds which have been long propagated by buds, cuttings, offsets, tubers, etc., and have in consequence been frequently exposed during what may be called their individual lives to widely diversified conditions of life. Plants thus propagated become so variable, that they are subject, as we have seen in the last chapter, even to bud-variation. Our domesticated animals, on the other hand, are not commonly exposed during the life of the individual to such extremely diversified conditions, and are not liable to such extreme variability; therefore they do not lose the power of transmitting most of their characteristic features. In the foregoing remarks on non-inheritance, crossed breeds are of course excluded, as their diversity mainly depends on the unequal development of character derived from either parent or their ancestors.

CONCLUSION.

It has been shown in the early part of this chapter how commonly new characters of the most diversified nature, whether normal or abnormal, injurious or beneficial, whether affecting organs of the highest or most trifling importance, are inherited. It is often sufficient for the inheritance of some peculiar character, that one parent alone should possess it, as in most cases in which the rarer anomalies have been transmitted. But the power of transmission is extremely variable. In a number of individuals descended from the same parents, and treated in the same manner, some display this power in a perfect manner, and in some it is quite deficient; and for this difference no reason can be a.s.signed. The effects of injuries or mutilations are occasionally inherited; and we shall see in a future chapter that the long-continued use and disuse of parts produces an inherited effect. Even those characters which are considered the most fluctuating, such as colour, are with rare exceptions transmitted much more forcibly than is generally supposed. The wonder, indeed, in all cases is not that any character should be transmitted, but that the power of inheritance should ever fail. The checks to inheritance, as far as we know them, are, firstly, circ.u.mstances hostile to the particular character in question; secondly, conditions of life incessantly inducing fresh variability; and lastly, the crossing of distinct varieties during some previous generation, together with reversion or atavism-that is, the tendency in the child to resemble its grand-parents or more remote ancestors instead of its immediate parents. This latter subject will be discussed in the following chapter.

END OF VOLUME I.