The Variation of Animals and Plants under Domestication — Volume 2
Inherited diseases afford evidence in some respects of less value than the foregoing cases, because diseases are not necessarily connected with any change in structure; but in other respects of more value, because the periods have been more carefully observed. Certain diseases are communicated to the child apparently by a process like inoculation, and the child is from the first affected; such cases may be here passed over. Large classes of diseases usually appear at certain ages, such as St. Vitus's dance in youth, consumption in early mid-life, gout later, and apoplexy still later; and these are naturally inherited at the same period. But even in diseases of this class, instances have been recorded, as with St. Vitus's dance, showing that an unusually early or late tendency to the disease is inheritable. (14/35. Dr. Prosper Lucas 'Hered. Nat.' tome 2 page 713.) In most cases the appearance of any inherited disease is largely determined by certain critical periods in each person's life, as well as by unfavourable conditions. There are many other diseases, which are not attached to any particular period, but which certainly tend to appear in the child at about the same age at which the parent was first attacked. An array of high authorities, ancient and modern, could be given in support of this proposition. The illustrious Hunter believed in it; and Piorry (14/36. 'L'Hered. dans les Maladies' 1840 page 135. For Hunter see Harlan 'Med. Researches' page 530.) cautions the physician to look closely to the child at the period when any grave inheritable disease attacked the parent. Dr. Prosper Lucas (14/37. 'L'Hered. Nat.' tome 2 page 850.), after collecting facts from every source, asserts that affections of all kinds, though not related to any particular period of life, tend to reappear in the offspring at whatever period of life they first appeared in the progenitor.
As the subject is important, it may be well to give a few instances, simply as illustrations, not as proof; for proof, recourse must be had to the authorities above quoted. Some of the following cases have been selected for the sake of showing that, when a slight departure from the rule occurs, the child is affected somewhat earlier in life than the parent. In the family of Le Compte blindness was inherited through three generations, and no less than twenty-seven children and grandchildren were all affected at about the same age; their blindness in general began to advance about the fifteenth or sixteenth year, and ended in total deprivation of sight at the age of about twenty-two. (14/38. Sedgwick 'Brit. and For. Med. — Chirurg. Review' April 1861 page 485. In some accounts the number of children and grandchildren is given as 37; but this seems to be an error judging from the paper first published in the 'Baltimore Med. and Phys. Reg.' 1809 of which Mr. Sedgwick has been so kind as to send me a copy.) In another case a father and his four children all became blind at twenty-one years old; in another, a grandmother grew blind at thirty-five, her daughter at nineteen, and three grandchildren at the ages of thirteen and eleven. (14/39. Prosper Lucas 'Hered. Nat.' tome 1 page 400.) So with deafness, two brothers, their father and paternal grandfather, all became deaf at the age of forty. (14/40. Sedgwick ibid July 1861 page 202.)
Esquirol gives several striking instances of insanity coming on at the same age, as that of a grandfather, father, and son, who all committed suicide near their fiftieth year. Many other cases could be given, as of a whole family who became insane at the age of forty. (14/41. Piorry page 109; Prosper Lucas tome 2 page 759.) Other cerebral affections sometimes follow the same rule, — for instance, epilepsy and apoplexy. A woman died of the latter disease when sixty-three years old; one of her daughters at forty-three, and the other at sixty-seven: the latter had twelve children, who all died from tubercular meningitis. (14/42. Prosper Lucas tome 2 page 748.) I mention this latter case because it illustrates a frequent occurrence, namely, a change in the precise nature of an inherited disease, though still affecting the same organ.
Asthma has attacked several members of the same family when forty years old, and other families during infancy. The most different diseases, such as angina pectoris, stone in the bladder, and various affections of the skin, have appeared in successive generations at nearly the same age. The little finger of a man began from some unknown cause to grow inwards, and the same finger in his two sons began at the same age to bend inwards in a similar manner. Strange and inexplicable neuralgic affections have caused parents and children to suffer agonies at about the same period of life. (14/43. Prosper Lucas tome 3 pages 678, 700, 702; Sedgwick ibid April 1863 page 449 and July 1863 page 162. Dr. J. Steinan 'Essay on Hereditary Disease' 1843 pages 27, 34.)
I will give only two other cases, which are interesting as illustrating the disappearance as well as the appearance of disease at the same age. Two brothers, their father, their paternal uncles, seven cousins, and their paternal grandfather, were all similarly affected by a skin-disease, called pityriasis versicolor; "the disease, strictly limited to the males of the family (though transmitted through the females), usually appeared at puberty, and disappeared at about the age of forty or forty-five years." The second case is that of four brothers, who when about twelve years old suffered almost every week from severe headaches, which were relieved only by a recumbent position in a dark room. Their father, paternal uncles, paternal grandfather, and granduncles all suffered in the same way from headaches, which ceased at the age of fifty-four or fifty-five in all those who lived so long. None of the females of the family were affected. (14/44. These cases are given by Mr. Sedgwick on the authority of Dr. H. Stewart in 'Med. — Chirurg. Review' April 1863 pages 449, 477.)]
It is impossible to read the foregoing accounts, and the many others which have been recorded, of diseases coming on during three or even more generations in several members of the same family at the same age, especially in the case of rare affections in which the coincidence cannot be attributed to chance, and to doubt that there is a strong tendency to inheritance in disease at corresponding periods of life. When the rule fails, the disease is apt to come on earlier in the child than in the parent; the exceptions in the other direction being very much rarer. Dr. Lucas (14/45. 'Hered. Nat.' tome 2 page 852.) alludes to several cases of inherited diseases coming on at an earlier period. I have already given one striking instance with blindness during three generations; and Mr. Bowman remarks that this frequently occurs with cataract. With cancer there seems to be a peculiar liability to earlier inheritance: Sir J. Paget, who has particularly attended to this subject, and tabulated a large number of cases, informs me that he believes that in nine cases out of ten the later generation suffers from the disease at an earlier period than the previous generation. He adds, "In the instances in which the opposite relation holds, and the members of later generations have cancer at a later age than their predecessors, I think it will be found that the non- cancerous parents have lived to extreme old ages." So that the longevity of a non-affected parent seems to have the power of influencing the fatal period in the offspring; and we thus apparently get another element of complexity in inheritance.
The facts, showing that with certain diseases the period of inheritance occasionally or even frequently advances, are important with respect to the general descent-theory, for they render it probable that the same thing would occur with ordinary modifications of structure. The final result of a long series of such advances would be the gradual obliteration of characters proper to the embryo and larva, which would thus come to resemble more and more closely the mature parent-form. But any structure which was of service to the embryo or larva would be preserved by the destruction at this stage of growth of each individual which manifested any tendency to lose its proper character at too early an age.
Finally, from the numerous races of cultivated plants and domestic animals, in which the seeds or eggs, the young or old, differ from one another and from those of the parent-species; — from the cases in which new characters have appeared at a particular period, and afterwards been inherited at the same period; — and from what we know with respect to disease, we must believe in the truth of the great principle of inheritance at corresponding periods of life.
SUMMARY OF THE THREE PRECEDING CHAPTERS.
Strong as is the force of inheritance, it allows the incessant appearance of new characters. These, whether beneficial or injurious, — of the most trifling importance, such as a shade of colour in a flower, a coloured lock of hair, or a mere gesture, — or of the highest importance, as when affecting the brain, or an organ so perfect and complex as the eye, — or of so grave a nature as to deserve to be called a monstrosity, — or so peculiar as not to occur normally in any member of the same natural class, — are often inherited by man, by the lower animals, and plants. In numberless cases it suffices for the inheritance of a peculiarity that one parent alone should be thus characterised. Inequalities in the two sides of the body, though opposed to the law of symmetry, may be transmitted. There is ample evidence that the effects of mutilations and of accidents, especially or perhaps exclusively when followed by disease, are occasionally inherited. There can be no doubt that the evil effects of the long-continued exposure of the parent to injurious conditions are sometimes transmitted to the offspring. So it is, as we shall see in a future chapter, with the effects of the use and disuse of parts, and of mental habits. Periodical habits are likewise transmitted, but generally, as it would appear, with little force.
Hence we are led to look at inheritance as the rule, and non-inheritance as the anomaly. But this power often appears to us in our ignorance to act capriciously, transmitting a character with inexplicable strength or feebleness. The very same peculiarity, as the weeping habit of trees, silky feathers, etc., may be inherited either firmly or not at all by different members of the same group, and even by different individuals of the same species, though treated in the same manner. In this latter case we see that the power of transmission is a quality which is merely individual in its attachment. As with single characters, so it is with the several concurrent slight differences which distinguish sub-varieties or races; for of these, some can be propagated almost as truly as species, whilst others cannot be relied on. The same rule holds good with plants, when propagated by bulbs, offsets, etc., which in one sense still form parts of the same individual, for some varieties retain or inherit through successive bud-generations their character far more truly than others.
Some characters not proper to the parent-species have certainly been inherited from an extremely remote epoch, and may therefore be considered as firmly fixed. But it is doubtful whether length of inheritance in itself gives fixedness of character; though the chances are obviously in favour of any character which has long been transmitted true or unaltered still being transmitted true as long as the conditions of life remain the same. We know that many species, after having retained the same character for countless ages, whilst living under their natural conditions, when domesticated have varied in the most diversified manner, that is, have failed to transmit their original form; so that no character appears to be absolutely fixed. We can sometimes account for the failure of inheritance by the conditions of life being opposed to the development of certain characters; and still oftener, as with plants cultivated by grafts and buds, by the conditions causing new and slight modifications incessantly to appear. In this latter case it is not that inheritance wholly fails, but that new characters are continually superadded. In some few cases, in which both parents are similarly characterised, inheritance seems to gain so much force by the combined action of the two parents, that it counteracts its own power, and a new modification is the result.
In many cases the failure of the parents to transmit their likeness is due to the breed having been at some former period crossed; and the child takes after his grandparent or more remote ancestor of foreign blood. In other cases, in which the breed has not been crossed, but some ancient character has been lost through variation, it occasionally reappears through reversion, so that the parents apparently fail to transmit their own likeness. In all cases, however, we may safely conclude that the child inherits all its characters from its parents, in whom certain characters are latent, like the secondary sexual characters of one sex in the other. When, after a long succession of bud- generations, a flower or fruit becomes separated into distinct segments, having the colours or other attributes of both parent-forms, we cannot doubt that these characters were latent in the earlier buds, though they could not then be detected, or could be detected only in an intimately commingled state. So it is with animals of crossed parentage, which with advancing years occasionally exhibit characters derived from one of their two parents, of which not a trace could at first be perceived. Certain monstrosities, which resemble what naturalists call the typical form of the group in question, apparently come under the same law of reversion. It is assuredly an astonishing fact that the male and female sexual elements, that buds, and even full-grown animals, should retain characters, during several generations in the case of crossed breeds, and during thousands of generations in the case of pure breeds, written as it were in invisible ink, yet ready at any time to be evolved under certain conditions.
What these conditions precisely are, we do not know. But any cause which disturbs the organisation or constitution seems to be sufficient. A cross certainly gives a strong tendency to the reappearance of long-lost characters, both corporeal and mental. In the case of plants, this tendency is much stronger with those species which have been crossed after long cultivation and which therefore have had their constitutions disturbed by this cause as well as by crossing, than with species which have always lived under their natural conditions and have then been crossed. A return, also, of domesticated animals and cultivated plants to a wild state favours reversion; but the tendency under these circumstances has been much exaggerated.
When individuals of the same family which differ somewhat, and when races or species are crossed, the one is often prepotent over the other in transmitting its character. A race may possess a strong power of inheritance, and yet when crossed, as we have seen with trumpeter-pigeons, yield to the prepotency of every other race. Prepotency of transmission may be equal in the two sexes of the same species, but often runs more strongly in one sex. It plays an important part in determining the rate at which one race can be modified or wholly absorbed by repeated crosses with another. We can seldom tell what makes one race or species prepotent over another; but it sometimes depends on the same character being present and visible in one parent, and latent or potentially present in the other.
Characters may first appear in either sex, but oftener in the male than in the female, and afterwards be transmitted to the offspring of the same sex. In this case we may feel confident that the peculiarity in question is really present though latent in the opposite sex! hence the father may transmit through his daughter any character to his grandson; and the mother conversely to her granddaughter. We thus learn, and the fact is an important one, that transmission and development are distinct powers. Occasionally these two powers seem to be antagonistic, or incapable of combination in the same individual; for several cases have been recorded in which the son has not directly inherited a character from his father, or directly transmitted it to his son, but has received it by transmission through his non-affected mother, and transmitted it through his non-affected daughter. Owing to inheritance being limited by sex, we see how secondary sexual characters may have arisen under nature; their preservation and accumulation being dependent on their service to either sex.
At whatever period of life a new character first appears, it generally remains latent in the offspring until a corresponding age is attained, and then is developed. When this rule fails, the child generally exhibits the character at an earlier period than the parent. On this principle of inheritance at corresponding periods, we can understand how it is that most animals display from the germ to maturity such a marvellous succession of characters.
Finally, though much remains obscure with respect to Inheritance, we may look at the following laws as fairly well established.
FIRSTLY, a tendency in every character, new and old, to be transmitted by seminal and bud generation, though often counteracted by various known and unknown causes.
SECONDLY, reversion or atavism, which depends on transmission and development being distinct powers: it acts in various degrees and manners through both seminal and bud generation.
THIRDLY, prepotency of transmission, which may be confined to one sex, or be common to both sexes.
FOURTHLY, transmission, as limited by sex, generally to the same sex in which the inherited character first appeared; and this in many, probably most cases, depends on the new character having first appeared at a rather late period of life.
FIFTHLY, inheritance at corresponding periods of life, with some tendency to the earlier development of the inherited character.
In these laws of Inheritance, as displayed under domestication, we see an ample provision for the production, through variability and natural selection, of new specific forms.
CHAPTER 2.XV
ON CROSSING.
FREE INTERCROSSING OBLITERATES THE DIFFERENCES BETWEEN ALLIED BREEDS. WHEN THE NUMBERS OF TWO COMMINGLING BREEDS ARE UNEQUAL, ONE ABSORBS THE OTHER. THE RATE OF ABSORPTION DETERMINED BY PREPOTENCY OF TRANSMISSION, BY THE CONDITIONS OF LIFE, AND BY NATURAL SELECTION. ALL ORGANIC BEINGS OCCASIONALLY INTERCROSS; APPARENT EXCEPTIONS. ON CERTAIN CHARACTERS INCAPABLE OF FUSION; CHIEFLY OR EXCLUSIVELY THOSE WHICH HAVE SUDDENLY APPEARED IN THE INDIVIDUAL. ON THE MODIFICATION OF OLD RACES, AND THE FORMATION OF NEW RACES BY CROSSING. SOME CROSSED RACES HAVE BRED TRUE FROM THEIR FIRST PRODUCTION. ON THE CROSSING OF DISTINCT SPECIES IN RELATION TO THE FORMATION OF DOMESTIC RACES.
In the two previous chapters, when discussing reversion and prepotency, I was necessarily led to give many facts on crossing. In the present chapter I shall consider the part which crossing plays in two opposed directions, — firstly, in obliterating characters, and consequently in preventing the formation of new races; and secondly, in the modification of old races, or in the formation of new and intermediate races, by a combination of characters. I shall also show that certain characters are incapable of fusion.
The effects of free or uncontrolled breeding between the members of the same variety or of closely allied varieties are important; but are so obvious that they need not be discussed at much length. It is free intercrossing which chiefly gives uniformity, both under nature and under domestication, to the individuals of the same species or variety, when they live mingled together and are not exposed to any cause inducing excessive variability. The prevention of free crossing, and the intentional matching of individual animals, are the corner-stones of the breeder's art. No man in his senses would expect to improve or modify a breed in any particular manner, or keep an old breed true and distinct, unless he separated his animals. The killing of inferior animals in each generation comes to the same thing as their separation. In savage and semi-civilised countries, where the inhabitants have not the means of separating their animals, more than a single breed of the same species rarely or never exists. In former times, even in the United States, there were no distinct races of sheep, for all had been mingled together. (15/1. 'Communications to the Board of Agriculture' volume 1 page 367.) The celebrated agriculturist Marshall (15/2. 'Review of Reports, North of England' 1808 page 200.) remarks that "sheep that are kept within fences, as well as shepherded flocks in open countries, have generally a similarity, if not a uniformity, of character in the individuals of each flock;" for they breed freely together, and are prevented from crossing with other kinds; whereas in the unenclosed parts of England the unshepherded sheep, even of the same flock, are far from true or uniform, owing to various breeds having mingled and crossed. We have seen that the half-wild cattle in each of the several British parks are nearly uniform in character; but in the different parks, from not having mingled and crossed during many generations, they differ to a certain small extent.
We cannot doubt that the extraordinary number of varieties and sub-varieties of the pigeon, amounting to at least one hundred and fifty, is partly due to their remaining, differently from other domesticated birds, paired for life once matched. On the other hand, breeds of cats imported into this country soon disappear, for their nocturnal and rambling habits render it hardly possible to prevent free crossing. Rengger (15/3. 'Saugethiere von Paraguay' 1830 s. 212.) gives an interesting case with respect to the cat in Paraguay: in all the distant parts of the kingdom it has assumed, apparently from the effects of the climate, a peculiar character, but near the capital this change has been prevented, owing, as he asserts, to the native animal frequently crossing with cats imported from Europe. In all cases like the foregoing, the effects of an occasional cross will be augmented by the increased vigour and fertility of the crossed offspring, of which fact evidence will hereafter be given; for this will lead to the mongrels increasing more rapidly than the pure parent-breeds.
When distinct breeds are allowed to cross freely, the result will be a heterogeneous body; for instance, the dogs in Paraguay are far from uniform, and can no longer be affiliated to their parent-races. (15/4. Rengger 'Saugethiere' etc. s. 154.) The character which a crossed body of animals will ultimately assume must depend on several contingencies, — namely, on the relative members of the individuals belonging to the two or more races which are allowed to mingle; on the prepotency of one race over the other in the transmission of character; and on the conditions of life to which they are exposed. When two commingled breeds exist at first in nearly equal numbers, the whole will sooner or later become intimately blended, but not so soon, both breeds being equally favoured in all respects, as might have been expected. The following calculation (15/5. White 'Regular Gradation in Man' page 146.) shows that this is the case: if a colony with an equal number of black and white men were founded, and we assume that they marry indiscriminately, are equally prolific, and that one in thirty annually dies and is born; then "in 65 years the number of blacks, whites, and mulattoes would be equal. In 91 years the whites would be 1-10th, the blacks 1-10th, and the mulattoes, or people of intermediate degrees of colour, 8-10ths of the whole number. In three centuries not 1-100th part of the whites would exist."
When one of two mingled races exceed the other greatly in number, the latter will soon be wholly, or almost wholly, absorbed and lost. (15/6. Dr. W.F. Edwards in his 'Caracteres Physiolog. des Races Humaines' page 24 first called attention to this subject, and ably discussed it.) Thus European pigs and dogs have been largely introduced in the islands of the Pacific Ocean, and the native races have been absorbed and lost in the course of about fifty or sixty years (15/7. Rev. D. Tyerman and Bennett 'Journal of Voyages' 1821-1829 volume 1 page 300.); but the imported races no doubt were favoured. Rats may be considered as semi-domesticated animals. Some snake-rats (Mus alexandrinus) escaped in the Zoological Gardens of London "and for a long time afterwards the keepers frequently caught cross-bred rats, at first half-breds, afterwards with less of the character of the snake-rat, till at length all traces of it disappeared." (15/8. Mr. S.J. Salter 'Journal Linn. Soc.' volume 6 1862 page 71.) On the other hand, in some parts of London, especially near the docks, where fresh rats are frequently imported, an endless variety of intermediate forms may be found between the brown, black, and snake rat, which are all three usually ranked as distinct species.