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 {86} 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 a country so civilised as North America, there were no distinct races of sheep, for all had been mingled together.[180] The celebrated agriculturist Marshall[181] 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 the several British parks are uniform in character in each; but in the different parks, from not having mingled and crossed during many generations, they differ in a slight degree.
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 when 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[182] 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.
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When distinct breeds are allowed to cross freely, the result will be a heterogenous body; for instance, the dogs in Paraguay are far from uniform, and can no longer be affiliated to their parent-races.[183] The character which a crossed body of animals will ultimately assume must depend on several contingencies,--namely, on the relative numbers 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[184] 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 exceeds the other greatly in number, the latter will soon be wholly, or almost wholly, absorbed and lost.[185] Thus European pigs and dogs have been largely introduced into the islands of the Pacific Ocean, and the native races have been absorbed and lost in the course of about fifty or sixty years;[186] 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 and less of the character of the snake-rat, till at length all traces of it disappeared."[187] On the other hand, {88} 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.
How many generations are necessary for one species or race to absorb another by repeated crosses has often been discussed;[188] and the requisite number has probably been much exaggerated. Some writers have maintained that a dozen, or score, or even more generations, are necessary; but this in itself is improbable, for in the tenth generation there will be only 1-1024th part of foreign blood in the offspring. Gärtner found,[189] that with plants one species could be made to absorb another in from three to five generations, and he believes that this could always be effected in from six to seventh generations. In one instance, however, Kölreuter[190] speaks of the offspring of Mirabilis vulgaris, crossed during eight successive generations by M. longiflora, as resembling this latter species so closely, that the most scrupulous observer could detect "vix aliquam notabilem differentiam;"--he succeeded, as he says, "ad plenariam fere transmutationem." But this expression shows that the act of absorption was not even then absolutely complete, though these crossed plants contained only the 1-256th part of M. vulgaris. The conclusions of such accurate observers as Gärtner and Kölreuter are of far higher worth than those made without scientific aim by breeders. The most remarkable statement which I have met with of the persistent endurance of the effects of a single cross is given by Fleischmann,[191] who, in reference to German sheep, says "that the original coarse sheep have 5500 fibres of wool on a square inch; grades of the third or fourth Merino cross produced about 8000, the twentieth cross 27,000, the perfect pure Merino blood 40,000 to 48,000." So that in this case common German sheep crossed twenty times successively with Merinos have not by any means acquired wool as fine as that of the pure breed. In all cases, the rate of absorption will {89} depend largely on the conditions of life being favourable to any particular character; and we may suspect that there would be under the climate of Germany a constant tendency to degeneration in the wool of Merinos, unless prevented by careful selection; and thus perhaps the foregoing remarkable case may be explained. The rate of absorption must also depend on the amount of distinguishable difference between the two forms which are crossed, and especially, as Gärtner insists, on prepotency of transmission in the one form over the other. We have seen in the last chapter that one of two French breeds of sheep yielded up its character, when crossed with Merinos, very much slower than the other; and the common German sheep referred to by Fleischmann may present an analogous case. But in all cases there will be during many subsequent generations more or less liability to reversion, and it is this fact which has probably led authors to maintain that a score or more of generations are requisite for one race to absorb another. In considering the final result of the commingling of two or more breeds, we must not forget that the act of crossing in itself tends to bring back long-lost characters not proper to the immediate parent-forms.
With respect to the influence of the conditions of life on any two breeds which are allowed to cross freely, unless both are indigenous and have long been accustomed to the country where they live, they will, in all probability, be unequally affected by the conditions, and this will modify the result. Even with indigenous breeds, it will rarely or never occur that both are equally well adapted to the surrounding circumstances; more especially when permitted to roam freely, and not carefully tended, as will generally be the case with breeds allowed to cross. As a consequence of this, natural selection will to a certain extent come into action, and the best fitted will survive, and this will aid in determining the ultimate character of the commingled body.
How long a time it would require before such a crossed body of animals would assume within a limited area a uniform character no one can say; that they would ultimately become uniform from free intercrossing, and from the survival of the fittest, we may feel assured; but the character thus acquired would rarely or never, as we may infer from the several previous {90} considerations, be exactly intermediate between that of the two parent-breeds. With respect to the very slight differences by which the individuals of the same sub-variety, or even of allied varieties, are characterised, it is obvious that free crossing would soon obliterate such small distinctions. The formation of new varieties, independently of selection, would also thus be prevented; except when the same variation continually recurred from the action of some strongly predisposing cause. Hence we may conclude that free crossing has in all cases played an important part in giving to all the members of the same domestic race, and of the same natural species, uniformity of character, though largely modified by natural selection and by the direct action of the surrounding conditions.
On the possibility of all organic beings occasionally intercrossing.--But it may be asked, can free crossing occur with hermaphrodite animals and plants? All the higher animals, and the few insects which have been domesticated, have separated sexes, and must inevitably unite for each birth. With respect to the crossing of hermaphrodites, the subject is too large for the present volume, and will be more properly treated in a succeeding work. In my 'Origin of Species,' however, I have given a short abstract of the reasons which induce me to believe that all organic beings occasionally cross, though perhaps in some cases only at long intervals of time.[192] I will here just recall the fact that many plants, though hermaphrodite in structure, are unisexual in function;--such as those called by C. K. Sprengel dichogamous, in which the pollen and stigma of the same flower are matured at different periods; or those called by me reciprocally dimorphic, in which the flower's own pollen is not fitted to fertilise its own stigma; or again, the many kinds in which curious mechanical contrivances exist, effectually preventing self-fertilisation. There are, however, many hermaphrodite plants which are not in any way specially constructed to favour intercrossing, but which nevertheless commingle almost as freely as animals with separated sexes. This is the case with cabbages, radishes, and onions, as I know from {91} having experimented on them: even the peasants of Liguria say that cabbages must be prevented "from falling in love" with each other. In the orange tribe, Gallesio[193] remarks that the amelioration of the various kinds is checked by their continual and almost regular crossing. So it is with numerous other plants.
Nevertheless some cultivated plants can be named which rarely intercross, as the common pea, or which never intercross, as I have reason to believe is the case with the sweet-pea (Lathyrus odoratus); yet the structure of these flowers certainly favours an occasional cross. The varieties of the tomato and aubergine (Solanum) and pimenta (Pimenta vulgaris?) are said[194] never to cross, even when growing alongside each other. But it should be observed that these are all exotic plants, and we do not know how they would behave in their native country when visited by the proper insects.
It must also be admitted that some few natural species appear under our present state of knowledge to be perpetually self-fertilised, as in the case of the Bee Ophrys (O. apifera), though adapted in its structure to be occasionally crossed. The Leersia oryzoides produces minute enclosed flowers which cannot possibly be crossed, and these alone, to the exclusion of the ordinary flowers, have as yet been known to yield seed.[195] A few additional and analogous cases could be advanced. But these facts do not make me doubt that it is a general law of nature that the individuals of the same species occasionally intercross, and that some great advantage is derived from this act. It is well known (and I shall hereafter have to give instances) that some plants, both indigenous and naturalised, rarely or never produce flowers; or, if they flower, never produce seeds. But no one is thus led to doubt that it is a general law of nature that phanerogamic plants should produce flowers, and that these flowers should produce seed. When they fail, we believe that such plants would perform their proper functions under different conditions, or that they formerly did so and will do so again. On analogous grounds, I believe that the few flowers {92} which do not now intercross, either would do so under different conditions, or that they formerly fertilised each other at intervals--the means for effecting this being generally still retained--and they will do so again at some future period, unless indeed they become extinct. On this view alone, many points in the structure and action of the reproductive organs in hermaphrodite plants and animals are intelligible,--for instance, the male and female organs never being so completely enclosed as to render access from without impossible. Hence we may conclude that the most important of all the means for giving uniformity to the individuals of the same species, namely, the capacity of occasionally intercrossing, is present, or has been formerly present, with all organic beings.
On certain Characters not blending.--When two breeds are crossed their characters usually become intimately fused together; but some characters refuse to blend, and are transmitted in an unmodified state either from both parents or from one. When grey and white mice are paired, the young are not piebald nor of an intermediate tint, but are pure white or of the ordinary grey colour: so it is when white and common collared turtle-doves are paired. In breeding Game fowls, a great authority, Mr. J. Douglas, remarks, "I may here state a strange fact: if you cross a black with a white game, you get birds of both breeds of the clearest colour." Sir R. Heron crossed during many years white, black, brown, and fawn-coloured Angora rabbits, and never once got these colours mingled in the same animal, but often all four colours in the same litter.[196] Additional cases could be given, but this form of inheritance is very far from universal even with respect to the most distinct colours. When turnspit dogs and ancon sheep, both of which have dwarfed limbs, are crossed with common breeds, the offspring are not intermediate in structure, but take after either parent. When tailless or hornless animals are crossed with perfect animals, it frequently, but by no means invariably, happens that the offspring are {93} either perfectly furnished with these organs or are quite destitute of them. According to Rengger, the hairless condition of the Paraguay dog is either perfectly or not at all transmitted to its mongrel offspring; but I have seen one partial exception in a dog of this parentage which had part of its skin hairy, and part naked; the parts being distinctly separated as in a piebald animal. When Dorking fowls with five toes are crossed with other breeds, the chickens often have five toes on one foot and four on the other. Some crossed pigs raised by Sir R. Heron between the solid-hoofed and common pig had not all four feet in an intermediate condition, but two feet were furnished with properly divided, and two with united hoofs.
Analogous facts have been observed with plants: Major Trevor Clarke crossed the little, glabrous-leaved, annual stock (Matthiola), with pollen of a large, red-flowered, rough-leaved, biennial stock, called cocardeau by the French, and the result was that half the seedlings had glabrous and the other half rough leaves, but none had leaves in an intermediate state. That the glabrous seedlings were the product of the rough-leaved variety, and not accidentally of the mother-plant's own pollen, was shown by their tall and strong habit of growth.[197] In the succeeding generations raised from the rough-leaved crossed seedlings, some glabrous plants appeared, showing that the glabrous character, though incapable of blending with and modifying the rough leaves, was all the time latent in this family of plants. The numerous plants formerly referred to, which I raised from reciprocal crosses between the peloric and common Antirrhinum, offer a nearly parallel case; for in the first generation all the plants resembled the common form, and in the next generation, out of one hundred and thirty-seven plants, two alone were in an intermediate condition, the others perfectly resembling either the peloric or common form. Major Trevor Clarke also fertilised the above-mentioned red-flowered stock with pollen from the purple Queen stock, and about half the seedlings scarcely differed in habit, and not at all in the red colour of the flower, from the mother-plant, the other half bearing blossoms of a rich purple, closely like those of the paternal plant. Gärtner crossed many white and yellow-flowered species and varieties of Verbascum; and these colours were never blended, but the offspring bore either pure white or pure yellow blossoms; the former in the larger proportion.[198] Dr. Herbert raised many seedlings, as he informed me, from Swedish turnips crossed by two other varieties, and these never produced flowers of an intermediate tint, but always like one of their parents. I fertilised the purple sweet-pea (Lathyrus odoratus), which has a dark reddish-purple standard-petal and violet-coloured wings and keel, with pollen of the painted-lady sweet-pea, which has a pale cherry-coloured standard, and almost white wings and keel; and from the same pod I twice raised plants perfectly resembling both sorts; the greater number resembling the father. So perfect was the resemblance, that I should have thought there had {94} been some mistake, if the plants which were at first identical with the paternal variety, namely, the painted-lady, had not later in the season produced, as mentioned in a former chapter, flowers blotched and streaked with dark purple. I raised grandchildren and great-grandchildren from these crossed plants, and they continued to resemble the painted-lady, but during the later generations became rather more blotched with purple, yet none reverted completely to the original mother-plant, the purple sweet-pea. The following case is slightly different, but still shows the same principle: Naudin[199] raised numerous hybrids between the yellow Linaria vulgaris and the purple L. purpurea, and during three successive generations the colours kept distinct in different parts of the same flower.
From such cases as the foregoing, in which the offspring of the first generation perfectly resemble either parent, we come by a small step to those cases in which differently coloured flowers borne on the same root resemble both parents, and by another step to those in which the same flower or fruit is striped or blotched with the two parental colours, or bears a single stripe of the colour or other characteristic quality of one of the parent-forms. With hybrids and mongrels it frequently or even generally happens that one part of the body resembles more or less closely one parent and another part the other parent; and here again some resistance to fusion, or, what comes to the same thing, some mutual affinity between the organic atoms of the same nature, apparently comes into play, for otherwise all parts of the body would be equally intermediate in character. So again, when the offspring of hybrids or mongrels, which are themselves nearly intermediate in character, revert either wholly or by segments to their ancestors, the principle of the affinity of similar, or the repulsion of dissimilar atoms, must come into action. To this principle, which seems to be extremely general, we shall recur in the chapter on pangenesis.
It is remarkable, as has been strongly insisted upon by Isidore Geoffroy St. Hilaire in regard to animals, that the transmission of characters without fusion occurs most rarely when species are crossed; I know of one exception alone, namely, with the hybrids naturally produced between the common and hooded crow (Corvus corone and cornix), which, however, are closely allied species, differing in nothing except colour. Nor have I met with any well-ascertained cases of transmission of this kind, even when one form is strongly prepotent over another, when two races are crossed which have been slowly formed by man's selection, and therefore resemble to a certain extent natural species. Such cases as puppies in the same litter closely resembling two distinct breeds, are probably due to super-foetation,--that is, to the influence of two fathers. All the characters above enumerated, which are transmitted in a perfect state to some of the offspring and not to others,--such as distinct colours, nakedness of skin, smoothness of leaves, absence of horns or tail, additional toes, pelorism, dwarfed structure, &c.,--have all been known to appear suddenly in individual animals and plants. From this fact, and from the several slight, aggregated differences which distinguish domestic races and species from {95} each other, not being liable to this peculiar form of transmission, we may conclude that it is in some way connected with the sudden appearance of the characters in question.
On the Modification of old Races and the Formation of new Races by Crossing.--We have hitherto chiefly considered the effects of crossing in giving uniformity of character; we must now look to an opposite result. There can be no doubt that crossing, with the aid of rigorous selection during several generations, has been a potent means in modifying old races, and in forming new ones. Lord Orford crossed his famous stud of greyhounds once with the bulldog, which breed was chosen from being deficient in scenting powers, and from having what was wanted, courage and perseverance. In the course of six or seven generations all traces of the external form of the bulldog were eliminated, but courage and perseverance remained. Certain pointers have been crossed, as I hear from the Rev. W. D. Fox, with the foxhound, to give them dash and speed. Certain strains of Dorking fowls have had a slight infusion of Game blood; and I have known a great fancier who on a single occasion crossed his turbit-pigeons with barbs, for the sake of gaining greater breadth of beak.
In the foregoing cases breeds have been crossed once, for the sake of modifying some particular character; but with most of the improved races of the pig, which now breed true, there have been repeated crosses,--for instance, the improved Essex owes its excellence to repeated crosses with the Neapolitan, together probably with some infusion of Chinese blood.[200] So with our British sheep: almost all the races, except the Southdown, have been largely crossed; "this, in fact, has been the history of our principal breeds."[201] To give an example, the "Oxfordshire Downs" now rank as an established breed.[202] They were produced about the year 1830 by crossing "Hampshire and in some instances Southdown ewes with Cotswold rams:" now the Hampshire ram was itself produced by repeated crosses between the native {96} Hampshire sheep and Southdowns; and the long-woolled Cotswold were improved by crosses with the Leicester, which latter again is believed to have been a cross between several long-woolled sheep. Mr. Spooner, after considering the various cases which have been carefully recorded, concludes "that from a judicious pairing of cross-bred animals it is practicable to establish a new breed." On the Continent the history of several crossed races of cattle and of other animals has been well ascertained. To give one instance: the King of Wurtemberg, after twenty-five years' careful breeding, that is after six or seven generations, made a new breed of cattle from a cross between a Dutch and Swiss breed, combined with other breeds.[203] The Sebright bantam, which breeds as true as any other kind of fowl, was formed about sixty years ago by a complicated cross.[204] Dark Brahmas, which are believed by some fanciers to constitute a distinct species, were undoubtedly formed[205] in the United States, within a recent period, by a cross between Chittagongs and Cochins. With plants I believe there is little doubt that some kinds of turnips, now extensively cultivated, are crossed races; and the history of a variety of wheat which was raised from two very distinct varieties, and which after six years' culture presented an even sample, has been recorded on good authority.[206]
Until quite lately, cautious and experienced breeders, though not averse to a single infusion of foreign blood, were almost universally convinced that the attempt to establish a new race, intermediate between two widely distinct races, was hopeless: "they clung with superstitious tenacity to the doctrine of purity of blood, believing it to be the ark in which alone true safety could be found."[207] Nor was this conviction unreasonable: when two distinct races are crossed, the offspring of the first generation are generally nearly uniform in character; but even this sometimes fails to be the case, especially with crossed dogs and fowls, the young of which from the first are sometimes much {97} diversified. As cross-bred animals are generally of large size and vigorous, they have been raised in great numbers for immediate consumption. But for breeding they are found to be utterly useless; for though they may be themselves uniform in character, when paired together they yield during many generations offspring astonishingly diversified. The breeder is driven to despair, and concludes that he will never form an intermediate race. But from the cases already given, and from others which have been recorded, it appears that patience alone is necessary; as Mr. Spooner remarks, "nature opposes no barrier to successful admixture; in the course of time, by the aid of selection and careful weeding, it is practicable to establish a new breed." After six or seven generations the hoped-for result will in most cases be obtained; but even then an occasional reversion, or failure to keep true, may be expected. The attempt, however, will assuredly fail if the conditions of life be decidedly unfavourable to the characters of either parent-breed.[208]
Although the grandchildren and succeeding generations of cross-bred animals are generally variable in an extreme degree, some curious exceptions to the rule have been observed, both with crossed races and species. Thus Boitard and Corbié[209] assert that from a Pouter and a Runt "a Cavalier will appear, which we have classed amongst pigeons of pure race, because it transmits all its qualities to its posterity." The editor of the 'Poultry Chronicle'[210] bred some bluish fowls from a black Spanish cock and a Malay hen; and these remained true to colour "generation after generation." The Himalayan breed of rabbits was certainly formed by crossing two sub-varieties of the silver-grey rabbit; although it suddenly assumed its present character, which differs much from that of either parent-breed, yet it has ever since been easily and truly propagated. I crossed some Labrador and Penguin ducks, and recrossed the mongrels with Penguins; afterwards, most of the ducks reared during three generations were nearly uniform in character, being brown with a white crescentic mark on the lower part of the breast, {98} and with some white spots at the base of the beak; so that by the aid of a little selection a new breed might easily have been formed. In regard to crossed varieties of plants, Mr. Beaton remarks[211] that "Melville's extraordinary cross between the Scotch kale and an early cabbage is as true and genuine as any on record;" but in this case no doubt selection was practised. Gärtner[212] has given five cases of hybrids, in which the progeny kept constant; and hybrids between Dianthus armeria and deltoides remained true and uniform to the tenth generation. Dr. Herbert likewise showed me a hybrid from two species of Loasa which from its first production had kept constant during several generations.
We have seen in the earlier chapters, that some of our domesticated animals, such as dogs, cattle, pigs, &c., are almost certainly descended from more than one species, or wild race, if any one prefers to apply this latter term to forms which were enabled to keep distinct in a state of nature. Hence the crossing of aboriginally distinct species probably came into play at an early period in the formation of our present races. From Rütimeyer's observations there can be little doubt that this occurred with cattle; but in most cases some one of the forms which were allowed to cross freely, will, it is probable, have absorbed and obliterated the others. For it is not likely that semi-civilized men would have taken the necessary pains to modify by selection their commingled, crossed, and fluctuating stock. Nevertheless, those animals which were best adapted to their conditions of life would have survived through natural selection; and by this means crossing will often have indirectly aided in the formation of primeval domesticated breeds.
Within recent times, as far as animals are concerned, the crossing of distinct species has done little or nothing in the formation or modification of our races. It is not yet known whether the species of silk-moth which have been recently crossed in France will yield permanent races. In the fourth chapter I alluded with some hesitation to the statement that a new breed, between the hare and rabbit, called leporides, had been formed in France, and was found capable of propagating {99} itself; but it is now positively affirmed[213] that this is an error. With plants which can be multiplied by buds and cuttings, hybridisation has done wonders, as with many kinds of Roses, Rhododendrons, Pelargoniums, Calceolarias, and Petunias. Nearly all these plants can be propagated by seed; most of them freely; but extremely few or none come true by seed.
Some authors believe that crossing is the chief cause of variability,--that is, of the appearance of absolutely new characters. Some have gone so far as to look at it as the sole cause; but this conclusion is disproved by some of the facts given in the chapter on Bud-variation. The belief that characters not present in either parent or in their ancestors frequently originate from crossing is doubtful; that they occasionally thus arise is probable; but this subject will be more conveniently discussed in a future chapter on the causes of Variability.
A condensed summary of this and of the three following chapters, together with some remarks on Hybridism, will be given in the nineteenth chapter.
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Chapter XVI
CAUSES WHICH INTERFERE WITH THE FREE CROSSING OF VARIETIES--INFLUENCE OF DOMESTICATION ON FERTILITY.
DIFFICULTIES IN JUDGING OF THE FERTILITY OF VARIETIES WHEN CROSSED--VARIOUS CAUSES WHICH KEEP VARIETIES DISTINCT, AS THE PERIOD OF BREEDING AND SEXUAL PREFERENCE--VARIETIES OF WHEAT SAID TO BE STERILE WHEN CROSSED--VARIETIES OF MAIZE, VERBASCUM, HOLLYHOCK, GOURDS, MELONS, AND TOBACCO, RENDERED IN SOME DEGREE MUTUALLY STERILE--DOMESTICATION ELIMINATES THE TENDENCY TO STERILITY NATURAL TO SPECIES WHEN CROSSED--ON THE INCREASED FERTILITY OF UNCROSSED ANIMALS AND PLANTS FROM DOMESTICATION AND CULTIVATION.
The domesticated races of both animals and plants, when crossed, are with extremely few exceptions quite prolific,--in some cases even more so than the purely bred parent-races. The offspring, also, raised from such crosses are likewise, as we shall see in the following chapter, generally more vigorous and fertile than their parents. On the other hand, species when crossed, and their hybrid offspring, are almost invariability in some degree sterile; and here there seems to exist a broad and insuperable distinction between races and species. The importance of this subject as bearing on the origin of species is obvious; and we shall hereafter recur to it.
It is unfortunate how few precise observations have been made on the fertility of mongrel animals and plants during several successive generations. Dr. Broca[214] has remarked that no one has observed whether, for instance, mongrel dogs, bred inter se, are indefinitely fertile; yet, if a shade of infertility be detected by careful observation in the offspring of natural forms when crossed, it is thought that their specific distinction is proved. But so many breeds of sheep, cattle, pigs, dogs, and poultry, have been crossed and recrossed in various ways, that any sterility, if it had existed, would from being injurious {101} almost certainly have been observed. In investigating the fertility of crossed varieties many sources of doubt occur. Whenever the least trace of sterility between two plants, however closely allied, was observed by Kölreuter, and more especially by Gärtner, who counted the exact number of seed in each capsule, the two forms were at once ranked as distinct species; and if this rule be followed, assuredly it will never be proved that varieties when crossed are in any degree sterile. We have formerly seen that certain breeds of dogs do not readily pair together; but no observations have been made whether, when paired, they produce the full number of young, and whether the latter are perfectly fertile inter se; but, supposing that some degree of sterility were found to exist, naturalists would simply infer that these breeds were descended from aboriginally distinct species; and it would be scarcely possible to ascertain whether or not this explanation was the true one.
The Sebright Bantam is much less prolific than any other breed of fowls, and is descended from a cross between two very distinct breeds, recrossed by a third sub-variety. But it would be extremely rash to infer that the loss of fertility was in any manner connected with its crossed origin, for it may with more probability be attributed either to long-continued close interbreeding, or to an innate tendency to sterility correlated with the absence of hackles and sickle tail-feathers.
Before giving the few recorded cases of forms, which must be ranked as varieties, being in some degree sterile when crossed, I may remark that other causes sometimes interfere with varieties freely intercrossing. Thus they may differ too greatly in size, as with some kinds of dogs and fowls: for instance, the editor of the 'Journal of Horticulture, &c.,'[215] says that he can keep Bantams with the larger breeds without much danger of their crossing, but not with the smaller breeds, such as Games, Hamburgs, &c. With plants a difference in the period of flowering serves to keep varieties distinct, as with the various kinds of maize and wheat: thus Colonel Le Couteur[216] remarks, "the Talavera wheat, from flowering much earlier than any other kind, is sure to continue pure." In different parts of {102} the Falkland Islands the cattle are breaking up into herds of different colours; and those on the higher ground, which are generally white, usually breed, as I am informed by Admiral Sulivan, three months earlier than those on the lowlands; and this would manifestly tend to keep the herds from blending.
Certain domestic races seem to prefer breeding with their own kind; and this is a fact of some importance, for it is a step towards that instinctive feeling which helps to keep closely allied species in a state of nature distinct. We have now abundant evidence that, if it were not for this feeling, many more hybrids would be naturally produced than is the case. We have seen in the first chapter that the alco dog of Mexico dislikes dogs of other breeds; and the hairless dog of Paraguay mixes less readily with the European races, than the latter do with each other. In Germany the female Spitz-dog is said to receive the fox more readily than will other dogs; a female Australian Dingo in England attracted the wild male foxes. But these differences in the sexual instinct and attractive power of the various breeds may be wholly due to their descent from distinct species. In Paraguay the horses have much freedom, and an excellent observer[217] believes that the native horses of the same colour and size prefer associating with each other, and that the horses which have been imported from Entre Rios and Banda Oriental into Paraguay likewise prefer associating together. In Circassia six sub-races of the horse are known and have received distinct names; and a native proprietor of rank[218] asserts that horses of three of these races, whilst living a free life, almost always refuse to mingle and cross, and will even attack each other.
It has been observed, in a district stocked with heavy Lincolnshire and light Norfolk sheep, that both kinds, though bred together, when turned out, "in a short time separate to a sheep;" the Lincolnshires drawing off to the rich soil, and the Norfolks to their own dry light soil; and as long as there is plenty of grass, "the two breeds keep themselves as distinct as rooks and pigeons." In this case different habits of {103} life tend to keep the races distinct. On one of the Faroe islands, not more than half a mile in diameter, the half-wild native black sheep are said not to have readily mixed with the imported white sheep. It is a more curious fact that the semi-monstrous ancon sheep of modern origin "have been observed to keep together, separating themselves from the rest of the flock, when put into enclosures with other sheep."[219] With respect to fallow deer, which live in a semi-domesticated condition, Mr. Bennett[220] states that the dark and pale coloured herds, which have long been kept together in the Forest of Dean, in High Meadow Woods, and in the New Forest, have never been known to mingle: the dark-coloured deer, it may be added, are believed to have been first brought by James I. from Norway, on account of their greater hardiness. I imported from the island of Porto Santo two of the feral rabbits, which differ, as described in the fourth chapter, from common rabbits; both proved to be males, and, though they lived during some years in the Zoological Gardens, the superintendent, Mr. Bartlett, in vain endeavoured to make them breed with various tame kinds; but whether this refusal to breed was due to any change in instinct, or simply to their extreme wildness; or whether confinement had rendered them sterile, as often occurs, cannot be told.
Whilst matching for the sake of experiment many of the most distinct breeds of pigeons, it frequently appeared to me that the birds, though faithful to their marriage vow, retained some desire after their own kind. Accordingly I asked Mr. Wicking, who has kept a larger stock of various breeds together than any man in England, whether he thought that they would prefer pairing with their own kind, supposing that there were males and females enough of each; and he without hesitation answered that he was convinced that this was the case. It has often been noticed that the dovecot pigeon seems to have an actual aversion towards the several fancy breeds;[221] yet all have {104} certainly sprung from a common progenitor. The Rev. W. D. Fox informs me that his flocks of white and common Chinese geese kept distinct.
These facts and statements, though some of them are incapable of proof, resting only on the opinion of experienced observers, show that some domestic races are led by different habits of life to keep to a certain extent separate, and that others prefer coupling with their own kind, in the same manner as species in a state of nature, though in a much less degree.
With respect to sterility from the crossing of domestic races, I know of no well-ascertained case with animals. This fact, seeing the great difference in structure between some breeds of pigeons, fowls, pigs, dogs, &c., is extraordinary, in contrast with the sterility of many closely allied natural species when crossed; but we shall hereafter attempt to show that it is not so extraordinary as it at first appears. And it may be well here to recall to mind that the amount of external difference between two species will not safely guide us in foretelling whether or not they will breed together,--some closely allied species when crossed being utterly sterile, and others which are extremely unlike being moderately fertile. I have said that no case of sterility in crossed races rests on satisfactory evidence; but here is one which at first seems trustworthy. Mr. Youatt,[222] and a better authority cannot be quoted, states, that formerly in Lancashire crosses were frequently made between longhorn and shorthorn cattle; the first cross was excellent, but the produce was uncertain; in the third or fourth generation the cows were bad milkers; "in addition to which, there was much uncertainty whether the cows would conceive; and full one-third of the cows among some of these half-breds failed to be in calf." This at first seems a good case; but Mr. Wilkinson states,[223] that a breed derived from this same cross was actually established in another part of England; and if it had failed in fertility, the fact would surely have been noticed. Moreover, supposing that Mr. Youatt had proved his case, it might be argued that the sterility was wholly due to the two parent-breeds being descended from primordially distinct species.
I will give a case with plants, to show how difficult it is to get sufficient evidence. Mr. Sheriff, who has been so successful in the formation of new races of wheat, fertilised the Hopetoun with the Talavera; in the first and second generations the produce was intermediate in character, but in the fourth generation "it was found to consist of many varieties; nine-tenths of the florets proved barren, and many of the seeds seemed shrivelled abortions, void of vitality, and the whole race was evidently verging to extinction."[224] Now, considering how little these {105} varieties of wheat differ in any important character, it seems to me very improbable that the sterility resulted, as Mr. Sheriff thought, from the cross, but from some quite distinct cause. Until such experiments are many times repeated, it would be rash to trust them; but unfortunately they have been rarely tried even once with sufficient care.
Gärtner has recorded a more remarkable and trustworthy case: he fertilised thirteen panicles (and subsequently nine others) on a dwarf maize bearing yellow seed[225] with pollen of a tall maize having red seed; and one head alone produced good seed, only five in number. Though these plants are monoecious, and therefore do not require castration, yet I should have suspected some accident in the manipulation had not Gärtner expressly stated that he had during many years grown these two varieties together, and they did not spontaneously cross; and this, considering that the plants are monoecious and abound with pollen, and are well known generally to cross freely, seems explicable only on the belief that these two varieties are in some degree mutually infertile. The hybrid plants raised from the above five seed were intermediate in structure, extremely variable, and perfectly fertile.[226] No one, I believe, has hitherto suspected that these varieties of maize are distinct species; but had the hybrids been in the least sterile, no doubt Gärtner would at once have so classed them. I may here remark, that with undoubted species there is not necessarily any close relation between the sterility of a first cross and that of the hybrid offspring. Some species can be crossed with facility, but produce utterly sterile hybrids; others can be crossed with extreme difficulty, but the hybrids when produced are moderately fertile. I am not aware, however, of any instance quite like this of the maize with natural species, namely, of a first cross made with difficulty, but yielding perfectly fertile hybrids.
The following case is much more remarkable, and evidently perplexed Gärtner, whose strong wish it was to draw a broad line of distinction between species and varieties. In the genus Verbascum, he made, during eighteen years, a vast number of experiments, and crossed no less than 1085 flowers and counted their seeds. Many of these experiments consisted in crossing white and yellow varieties of both V. lychnitis and V. blattaria with nine other species and their hybrids. That the white and yellow flowered plants of these two species are really varieties, no one has doubted; and Gärtner actually raised in the case of both species one variety from the seed of the other. Now in two of his works[227] he distinctly asserts that crosses between similarly-coloured flowers yield more seed than between dissimilarly-coloured; so that the yellow-flowered variety of either species (and conversely with the white-flowered variety), when crossed with pollen of its own kind, yields more seed than when crossed with that of the white variety; and so it is when differently coloured species are crossed. The general results may be seen in the Table at the {106} end of his volume. In one instance he gives[228] the following details; but I must premise that Gärtner, to avoid exaggerating the degree of sterility in his crosses, always compares the maximum number obtained from a cross with the average number naturally given by the pure mother-plant. The white-variety of V. lychnitis, naturally fertilised by its own pollen, gave from an average of twelve capsules ninety-six good seeds in each; whilst twenty flowers fertilised with pollen from the yellow variety of this same species, gave as the maximum only eighty-nine good seed; so that we have the proportion of 1000 to 908, according to Gärtner's usual scale. I should have thought it possible that so small a difference in fertility might have been accounted for by the evil effects of the necessary castration; but Gärtner shows that the white variety of V. lychnitis, when fertilised first by the white variety of V. blattaria, and then by the yellow variety of this species, yielded seed in the proportion of 622 to 438; and in both these cases castration was performed. Now the sterility which results from the crossing of the differently coloured varieties of the same species, is fully as great as that which occurs in many cases when distinct species are crossed. Unfortunately Gärtner compared the results of the first unions alone, and not the sterility of the two sets of hybrids produced from the white variety of V. lychnitis when fertilised by the white and yellow varieties of V. blattaria, for it is probable that they would have differed in this respect.
Mr. J. Scott has given me the results of a series of experiments on Verbascum, made by him in the Botanic Gardens of Edinburgh. He repeated some of Gärtner's experiments on distinct species, but obtained only fluctuating results; some confirmatory, but the greater number contradictory; nevertheless these seem hardly sufficient to overthrow the conclusions arrived at by Gärtner from experiments tried on a much larger scale. In the second place Mr. Scott experimented on the relative fertility of unions between similarly and dissimilarly-coloured varieties of the same species. Thus he fertilised six flowers of the yellow variety of V. lychnitis by its own pollen, and obtained six capsules, and calling, for the sake of having a standard of comparison, the average number of good seed in each one hundred, he found that this same yellow variety, when fertilised by the white variety, yielded from seven capsules an average of ninety-four seed. On the same principle, the white variety of V. lychnitis by its own pollen (from six capsules), and by the pollen of the yellow variety (eight capsules), yielded seed in the proportion of 100 to 82. The yellow variety of V. thapsus by its own pollen (eight capsules), and by that of the white variety (only two capsules), yielded seed in the proportion of 100 to 94. Lastly, the white variety of V. blattaria by its own pollen (eight capsules), and by that of the yellow variety (five capsules), yielded seed in the proportion of 100 to 79. So that in every case the unions of dissimilarly-coloured varieties of the same species were less fertile than the unions of similarly-coloured varieties; when all the cases are grouped together, the difference of fertility is as 86 to 100. Some additional trials were made, and altogether thirty-six similarly-coloured unions yielded thirty-five good {107} capsules; whilst thirty-five dissimilarly-coloured unions yielded only twenty-six good capsules. Besides the foregoing experiments, the purple V. phoeniceum was crossed by a rose-coloured and a white variety of the same species; these two varieties were also crossed together, and these several unions yielded less seed than V. phoeniceum by its own pollen. Hence it follows from Mr. Scott's experiments, that in the genus Verbascum the similarly and dissimilarly-coloured varieties of the same species behave, when crossed, like closely allied but distinct species.[229]
This remarkable fact of the sexual affinity of similarly-coloured varieties, as observed by Gärtner and Mr. Scott, may not be of very rare occurrence; for the subject has not been attended to by others. The following case is worth giving, partly to show how difficult it is to avoid error. Dr. Herbert[230] has remarked that variously-coloured double varieties of the hollyhock (Althæa rosea) may be raised with certainty by seed from plants growing close together. I have been informed that nurserymen who raise seed for sale do not separate their plants; accordingly I procured seed of eighteen named varieties; of these, eleven varieties produced sixty-two plants all perfectly true to their kind; and seven produced forty-nine plants, half of which were true and half false. Mr. Masters of Canterbury has given me a more striking case; he saved seed from a great bed of twenty-four named varieties planted in closely adjoining rows, and each variety reproduced itself truly with only sometimes a shade of difference in tint. Now in the hollyhock the pollen, which is abundant, is matured and nearly all shed before the stigma of the same flower is ready to receive it;[231] and as bees covered with pollen incessantly fly from plant to plant, it would appear that adjoining varieties could not escape being crossed. As, however, this does not occur, it appeared to me probable that the pollen {108} of each variety was prepotent on its own stigma over that of all other varieties. But Mr. C. Turner of Slough, well known for his success in the cultivation of this plant, informs me that it is the doubleness of the flowers which prevents the bees gaining access to the pollen and stigma; and he finds that it is difficult even to cross them artificially. Whether this explanation will fully account for varieties in close proximity propagating themselves so truly by seed, I do not know.
The following cases are worth giving, as they relate to monoecious forms, which do not require, and consequently have not been injured by, castration. Girou de Buzareingues crossed what he designates three varieties of gourd,[232] and asserts that their mutual fertilisation is less easy in proportion to the difference which they present. I am aware how imperfectly the forms in this group were until recently known; but Sageret,[233] who ranked them according to their mutual fertility, considers the three forms above alluded to as varieties, as does a far higher authority, namely, M. Naudin.[234] Sageret[235] has observed that certain melons have a greater tendency, whatever the cause may be, to keep true than others; and M. Naudin, who has had such immense experience in this group, informs me that he believes that certain varieties intercross more readily than others of the same species; but he has not proved the truth of this conclusion; the frequent abortion of the pollen near Paris being one great difficulty. Nevertheless, he has grown close together, during seven years, certain forms of Citrullus, which, as they could be artificially crossed with perfect facility and produced fertile offspring, are ranked as varieties; but these forms when not artificially crossed kept true. Many other varieties, on the other hand, in the same group cross with such facility, as M. Naudin repeatedly insists, that without being grown far apart they cannot be kept in the least true.
Another case, though somewhat different, may be here given, as it is highly remarkable, and is established on excellent evidence. Kölreuter minutely describes five varieties of the common tobacco,[236] which were reciprocally crossed, and the offspring were intermediate in character and as fertile as their parents: from this fact Kölreuter inferred that they are really varieties; and no one, as far as I can discover, seems to have doubted that such is the case. He also crossed reciprocally these five varieties with N. glutinosa, and they yielded very sterile hybrids; but those raised from the var. perennis, whether used as the father or mother plant, were not so sterile as the hybrids from the four other varieties.[237] So that the sexual {109} capacity of this one variety has certainly been in some degree modified, so as to approach in nature that of N. glutinosa.[238]
These facts with respect to plants show that in some few cases certain varieties have had their sexual powers so far modified, that they cross together less readily and yield less seed than other varieties of the same species. We shall presently see that the sexual functions of most animals and plants are eminently liable to be affected by the conditions of life to which they are exposed; and hereafter we shall briefly discuss the conjoint bearing of this and other facts on the difference in fertility between crossed varieties and crossed species.
Domestication eliminates the tendency to Sterility which is general with Species when crossed.
This hypothesis was first propounded by Pallas,[239] and has been adopted by several authors. I can find hardly any direct facts in its support; but unfortunately no one has compared, in the case of either animals or plants, the fertility of anciently domesticated varieties, when crossed with a distinct species, with that of the wild parent-species when similarly crossed. No one has compared, for instance, the fertility of Gallus bankiva and of the domesticated fowl, when crossed with a distinct species of Gallus or Phasianus; and the {110} experiment would in all cases be surrounded by many difficulties. Dureau de la Malle, who has so closely studied classical literature, states[240] that in the time of the Romans the common mule was produced with more difficulty than at the present day; but whether this statement may be trusted I know not. A much more important, though somewhat different, case is given by M. Groenland,[241] namely, that plants, known from their intermediate character and sterility to be hybrids between Ægilops and wheat, have perpetuated themselves under culture since 1857, with a rapid but varying increase of fertility in each generation. In the fourth generation the plants, still retaining their intermediate character, had become as fertile as common cultivated wheat.
The indirect evidence in favour of the Pallasian doctrine appears to me to be extremely strong. In the earlier chapters I have attempted to show that our various breeds of dogs are descended from several wild species; and this probably is the case with sheep. There can no longer be any doubt that the Zebu or humped Indian ox belongs to a distinct species from European cattle: the latter, moreover, are descended from two or three forms, which may be called either species or wild races, but which co-existed in a state of nature and kept distinct. We have good evidence that our domesticated pigs belong to at least two specific types, S. scrofa and Indica, which probably lived together in a wild state in South-eastern Europe. Now, a widely-extended analogy leads to the belief that if these several allied species, in the wild state or when first reclaimed, had been crossed, they would have exhibited, both in their first unions and in their hybrid offspring, some degree of sterility. Nevertheless the several domesticated races descended from them are now all, as far as can be ascertained, perfectly fertile together. If this reasoning be trustworthy, and it is apparently sound, we must admit the Pallasian doctrine that long-continued domestication tends to eliminate that sterility which is natural to species when crossed in their aboriginal state.
{111}
On increased Fertility from Domestication and Cultivation.
Increased fertility from domestication, without any reference to crossing, may be here briefly considered. This subject bears indirectly on two or three points connected with the modification of organic beings. As Buffon long ago remarked,[242] domestic animals breed oftener in the year and produce more young at a birth than wild animals of the same species; they, also, sometimes breed at an earlier age. The case would hardly have deserved further notice, had not some authors lately attempted to show that fertility increases and decreases in an inverse ratio with the amount of food. This strange doctrine has apparently arisen from individual animals when supplied with an inordinate quantity of food, and from plants of many kinds when grown on excessively rich soil, as on a dunghill, becoming sterile; but to this latter point I shall have occasion presently to return. With hardly an exception, our domesticated animals, which have long been habituated to a regular and copious supply of food, without the labour of searching for it, are more fertile than the corresponding wild animals. It is notorious how frequently cats and dogs breed, and how many young they produce at a birth. The wild rabbit is said generally to breed four times yearly, and to produce from four to eight young; the tame rabbit breeds six or seven times yearly, and produces from four to eleven young. The ferret, though generally so closely confined, is more prolific than its supposed wild prototype. The wild sow is remarkably prolific, for she often breeds twice in the year, and produces from four to eight and sometimes even twelve young at a birth; but the domestic sow regularly breeds twice a year, and would breed oftener if permitted; and a sow that produces less than eight at a birth "is worth little, and the sooner she is fattened for the butcher the better." The amount of food affects the fertility even of the same individual: thus sheep, which on mountains never produce more than one lamb at a birth, when brought {112} down to lowland pastures frequently bear twins. This difference apparently is not due to the cold of the higher land, for sheep and other domestic animals are said to be extremely prolific in Lapland. Hard living, also, retards the period at which animals conceive; for it has been found disadvantageous in the northern islands of Scotland to allow cows to bear calves before they are four years old.[243]
Birds offer still better evidence of increased fertility from domestication: the hen of the wild Gallus bankiva lays from six to ten eggs, a number which would be thought nothing of with the domestic hen. The wild duck lays from five to ten eggs; the tame one in the course of the year from eighty to one hundred. The wild grey-lag goose lays from five to eight eggs; the tame from thirteen to eighteen, and she lays a second time; as Mr. Dixon has remarked, "high-feeding, care, and moderate warmth induce a habit of prolificacy which becomes in some measure hereditary." Whether the semi-domesticated dovecot pigeon is more fertile than the wild rock-pigeon C. livia, I know not; but the more thoroughly domesticated breeds are nearly twice as fertile as dovecots: the latter, however, when caged and highly fed, become equally fertile with house pigeons. The peahen alone of domesticated birds is rather more fertile, according to some accounts, when wild in its native Indian home, than when domesticated in Europe and exposed to our much colder climate.[244]
With respect to plants, no one would expect wheat to tiller more, and each ear to produce more grain, in poor than in rich soil; or to get in poor soil a heavy crop of peas or beans. Seeds vary so much in number {113} that it is difficult to estimate them; but on comparing beds of carrots saved for seed in a nursery garden with wild plants, the former seemed to produce about twice as much seed. Cultivated cabbages yielded thrice as many pods by measure as wild cabbages from the rocks of South Wales. The excess of berries produced by the cultivated Asparagus in comparison with the wild plant is enormous. No doubt many highly cultivated plants, such as pears, pineapples, bananas, sugar-cane, &c., are nearly or quite sterile; and I am inclined to attribute this sterility to excess of food and to other unnatural conditions; but to this subject I shall presently recur.
In some cases, as with the pig, rabbit, &c., and with those plants which are valued for their seed, the direct selection of the more fertile individuals has probably much increased their fertility; and in all cases this may have occurred indirectly, from the better chance of the more numerous offspring produced by the more fertile individuals having survived. But with cats, ferrets, and dogs, and with plants like carrots, cabbages, and asparagus, which are not valued for their prolificacy, selection can have played only a subordinate part; and their increased fertility must be attributed to the more favourable conditions of life under which they have long existed.
* * * * *
{114}
Chapter XVII
ON THE GOOD EFFECTS OF CROSSING, AND ON THE EVIL EFFECTS OF CLOSE INTERBREEDING.
DEFINITION OF CLOSE INTERBREEDING--AUGMENTATION OF MORBID TENDENCIES--GENERAL EVIDENCE ON THE GOOD EFFECTS DERIVED FROM CROSSING, AND ON THE EVIL EFFECTS FROM CLOSE INTERBREEDING--CATTLE, CLOSELY INTERBRED; HALF-WILD CATTLE LONG KEPT IN THE SAME PARKS--SHEEP--FALLOW-DEER--DOGS--RABBITS--PIGS--MAN, ORIGIN OF HIS ABHORRENCE OF INCESTUOUS MARRIAGES--FOWLS--PIGEONS--HIVE-BEES--PLANTS, GENERAL CONSIDERATIONS ON THE BENEFITS DERIVED FROM CROSSING--MELONS, FRUIT-TREES, PEAS, CABBAGES, WHEAT, AND FOREST-TREES--ON THE INCREASED SIZE OF HYBRID PLANTS, NOT EXCLUSIVELY DUE TO THEIR STERILITY--ON CERTAIN PLANTS WHICH EITHER NORMALLY OR ABNORMALLY ARE SELF-IMPOTENT, BUT ARE FERTILE, BOTH ON THE MALE AND FEMALE SIDE, WHEN CROSSED WITH DISTINCT INDIVIDUALS EITHER OF THE SAME OR ANOTHER SPECIES--CONCLUSION.
The gain in constitutional vigour, derived from an occasional cross between individuals of the same variety, but belonging to distinct families, or between distinct varieties, has not been so largely or so frequently discussed, as have the evil effects of too close interbreeding. But the former point is the more important of the two, inasmuch as the evidence is more decisive. The evil results from close interbreeding are difficult to detect, for they accumulate slowly, and differ much in degree with different species; whilst the good effects which almost invariably follow a cross are from the first manifest. It should, however, be clearly understood that the advantage of close interbreeding, as far as the retention of character is concerned, is indisputable, and often outweighs the evil of a slight loss of constitutional vigour. In relation to the subject of domestication, the whole question is of some importance, as too close interbreeding interferes with the improvement of old races, and especially with the formation of new ones. It is important as indirectly bearing on Hybridism; and perhaps on the extinction of species, when any form has become so rare that only a few individuals {115} remain within a confined area. It bears in an important manner on the influence of free intercrossing, in obliterating individual differences, and thus giving uniformity of character to the individuals of the same race or species; for if additional vigour and fertility be thus gained, the crossed offspring will multiply and prevail, and the ultimate result will be far greater than otherwise would have occurred. Lastly, the question is of high interest, as bearing on mankind. Hence I shall discuss this subject at full length. As the facts which prove the evil effects of close interbreeding are more copious, though less decisive, than those on the good effects of crossing, I shall, under each group of beings, begin with the former.
There is no difficulty in defining what is meant by a cross; but this is by no means easy in regard to "breeding in and in" or "too close interbreeding," because, as we shall see, different species of animals are differently affected by the same degree of interbreeding. The pairing of a father and daughter, or mother and son, or brothers and sisters, if carried on during several generations, is the closest possible form of interbreeding. But some good judges, for instance Sir J. Sebright, believe that the pairing of a brother and sister is closer than that of parents and children; for when the father is matched with his daughter he crosses, as is said, with only half his own blood. The consequences of close interbreeding carried on for too long a time, are, as is generally believed, loss of size, constitutional vigour, and fertility, sometimes accompanied by a tendency to malformation. Manifest evil does not usually follow from pairing the nearest relations for two, three, or even four generations; but several causes interfere with our detecting the evil--such as the deterioration being very gradual, and the difficulty of distinguishing between such direct evil and the inevitable augmentation of any morbid tendencies which may be latent or apparent in the related parents. On the other hand, the benefit from a cross, even when there has not been any very close interbreeding, is almost invariably at once conspicuous. There is reason to believe, and this was the opinion of that most experienced observer Sir J. Sebright,[245] that the evil effects of close interbreeding may be checked by the related individuals {116} being separated during a few generations and exposed to different conditions of life.
That evil directly follows from any degree of close interbreeding has been denied by many persons; but rarely by any practical breeder; and never, as far as I know, by one who has largely bred animals which propagate their kind quickly. Many physiologists attribute the evil exclusively to the combination and consequent increase of morbid tendencies common to both parents: that this is an active source of mischief there can be no doubt. It is unfortunately too notorious that men and various domestic animals endowed with a wretched constitution, and with a strong hereditary disposition to disease, if not actually ill, are fully capable of procreating their kind. Close interbreeding, on the other hand, induces sterility; and this indicates something quite distinct from the augmentation of morbid tendencies common to both parents. The evidence immediately to be given convinces me that it is a great law of nature, that all organic beings profit from an occasional cross with individuals not closely related to them in blood; and that, on the other hand, long-continued close interbreeding is injurious.
Various general considerations have had much influence in leading me to this conclusion; but the reader will probably rely more on special facts and opinions. The authority of experienced observers, even when they do not advance the grounds of their belief, is of some little value. Now almost all men who have bred many kinds of animals and have written on the subject, such as Sir J. Sebright, Andrew Knight, &c.,[246] have expressed the strongest conviction on the impossibility of long-continued close interbreeding. Those who have compiled works on agriculture, and have associated much with breeders, such as the sagacious Youatt, Low, &c., have strongly declared their opinion to the same effect. Prosper Lucas, trusting largely to French authorities, has come to a similar conclusion. The distinguished German agriculturist Hermann von Nathusius, who has written the most able treatise on this subject which I have met with, concurs; and as I shall have to quote from {117} this treatise, I may state that Nathusius is not only intimately acquainted with works on agriculture in all languages, and knows the pedigrees of our British breeds better than most Englishmen, but has imported many of our improved animals, and is himself an experienced breeder.
Evidence of the evil effects of close interbreeding can most readily be acquired in the case of animals, such as fowls, pigeons, &c., which propagate quickly, and, from being kept in the same place, are exposed to the same conditions. Now I have inquired of very many breeders of these birds, and I have hitherto not met with a single man who was not thoroughly convinced that an occasional cross with another strain of the same sub-variety was absolutely necessary. Most breeders of highly-improved or fancy birds value their own strain, and are most unwilling, at the risk, in their opinion, of deterioration, to make a cross. The purchase of a first-rate bird of another strain is expensive, and exchanges are troublesome; yet all breeders, as far as I can hear, excepting those who keep large stocks at different places for the sake of crossing, are driven after a time to take this step.
Another general consideration which has had great influence on my mind is, that with all hermaphrodite animals and plants, which it might have been thought would have perpetually fertilised themselves, and thus have been subjected for long ages to the closest interbreeding, there is no single species, as far as I can discover, in which the structure ensures self-fertilisation. On the contrary, there are in a multitude of cases, as briefly stated in the fifteenth chapter, manifest adaptations which favour or inevitably lead to an occasional cross between one hermaphrodite and another of the same species; and these adaptive structures are utterly purposeless, as far as we can see, for any other end.
With Cattle there can be no doubt that extremely close interbreeding may be long carried on, advantageously with respect to external characters and with no manifestly apparent evil as far as constitution is concerned. The same remark is applicable to sheep. Whether these animals have gradually been rendered less susceptible than others to this evil, in order to permit them to live in herds,--a habit which leads the old and vigorous males to expel all intruders, and in consequence often to pair with their own daughters, I will not pretend to decide. The case of Bakewell's Long-horns, which were closely interbred for a long period, has often been {118} quoted; yet Youatt says[247] the breed "had acquired a delicacy of constitution inconsistent with common management," and "the propagation of the species was not always certain." But the Shorthorns offer the most striking case of close interbreeding; for instance, the famous bull Favourite (who was himself the offspring of a half-brother and sister from Foljambe) was matched with his own daughter, granddaughter, and great-granddaughter; so that the produce of this last union, or the great-great-granddaughter, had 15-16ths, or 93.75 per cent. of the blood of Favourite in her veins. This cow was matched with the bull Wellington, having 62.5 per cent. of Favourite blood in his veins, and produced Clarissa; Clarissa was matched with the bull Lancaster, having 68.75 of the same blood, and she yielded valuable offspring.[248] Nevertheless Collings, who reared these animals, and was a strong advocate for close breeding, once crossed his stock with a Galloway, and the cows from this cross realised the highest prices. Bates's herd was esteemed the most celebrated in the world. For thirteen years he bred most closely in and in; but during the next seventeen years, though he had the most exalted notion of the value of his own stock, he thrice infused fresh blood into his herd: it is said that he did this, not to improve the form of his animals, but on account of their lessened fertility. Mr. Bates's own view, as given by a celebrated breeder,[249] was, that "to breed in and in from a bad stock was ruin and devastation; yet that the practice may be safely followed within certain limits when the parents so related are descended from first-rate animals." We thus see that there has been extremely close interbreeding with Shorthorns; but Nathusius, after the most careful study of their pedigrees, says that he can find no instance of a breeder who has strictly followed this practice during his whole life. From this study and his own experience, he concludes that close interbreeding is necessary to ennoble the stock; but that in effecting this the greatest care is necessary, on account of the tendency to infertility and weakness. It may be added, that another high authority[250] asserts that many more calves are born cripples from Shorthorns than from other and less closely interbred races of cattle.
Although by carefully selecting the best animals (as Nature effectually does by the law of battle) close interbreeding may be long carried on with cattle, yet the good effects of a cross between almost any two breeds is at once shown by the greater size and vigour of the offspring; as Mr. Spooner writes to me, "crossing distinct breeds certainly improves cattle for the butcher." Such crossed animals are of course of no value to the breeder; but they have been raised during many years in several {119} parts of England to be slaughtered;[251] and their merit is now so fully recognised, that at fat-cattle shows a separate class has been formed for their reception. The best fat ox at the great show at Islington in 1862 was a crossed animal.
The half-wild cattle, which have been kept in British parks probably for 400 or 500 years, or even for a longer period, have been advanced by Culley and others as a case of long-continued interbreeding within the limits of the same herd without any consequent injury. With respect to the cattle at Chillingham, the late Lord Tankerville owned that they were bad breeders.[252] The agent, Mr. Hardy, estimates (in a letter to me, dated May, 1861) that in the herd of about fifty the average number annually slaughtered, killed by fighting, and dying, is about ten, or one in five. As the herd is kept up to nearly the same average number, the annual rate of increase must be likewise about one in five. The bulls, I may add, engage in furious battles, of which battles the present Lord Tankerville has given me a graphic description, so that there will always be rigorous selection of the most vigorous males. I procured in 1855 from Mr. D. Gardner, agent to the Duke of Hamilton, the following account of the wild cattle kept in the Duke's park in Lanarkshire, which is about 200 acres in extent. The number of cattle varies from sixty-five to eighty; and the number annually killed (I presume by all causes) is from eight to ten; so that the annual rate of increase can hardly be more than one in six. Now in South America, where the herds are half-wild, and therefore offer a nearly fair standard of comparison, according to Azara the natural increase of the cattle on an estancia is from one-third to one-fourth of the total number, or one in between three and four; and this, no doubt, applies exclusively to adult animals fit for consumption. Hence the half-wild British cattle which have long interbred within the limits of the same herd are relatively far less fertile. Although in an unenclosed country like Paraguay there must be some crossing between the different herds, yet even there the inhabitants believe that the occasional introduction of animals from distant localities is necessary to prevent "degeneration in size and diminution of fertility."[253] The decrease in size from ancient times in the Chillingham and Hamilton cattle must have been prodigious, for Professor Rütimeyer has shown that they are almost certainly the descendants of the gigantic Bos primigenius. No doubt this decrease in size may be largely attributed to less favourable conditions of life; yet animals roaming over large parks, and fed during severe winters, can hardly be considered as placed under very unfavourable conditions.
With Sheep there has often been long-continued interbreeding within the limits of the same flock; but whether the nearest relations have been matched so frequently as in the case of Shorthorn cattle, I do not know. The Messrs. Brown during fifty years have never infused fresh blood into their excellent flock of Leicesters. Since 1810 Mr. Barford has acted on the same principle with the Foscote flock. He asserts that half a century {120} of experience has convinced him that when two nearly related animals are quite sound in constitution, in-and-in breeding does not induce degeneracy; but he adds that he "does not pride himself on breeding from the nearest affinities." In France the Naz flock has been bred for sixty years without the introduction of a single strange ram.[254] Nevertheless, most great breeders of sheep have protested against close interbreeding prolonged for too great a length of time.[255] The most celebrated of recent breeders, Jonas Webb, kept five separate families to work on, thus "retaining the requisite distance of relationship between the sexes."[256]
Although by the aid of careful selection the near interbreeding of sheep may be long continued without any manifest evil, yet it has often been the practice with farmers to cross distinct breeds to obtain animals for the butcher, which plainly shows that good is derived from this practice. Mr. Spooner sums up his excellent Essay on Crossing by asserting that there is a direct pecuniary advantage in judicious cross-breeding, especially when the male is larger than the female. A former celebrated breeder, Lord Somerville, distinctly states that his half-breeds from Ryelands and Spanish sheep were larger animals than either the pure Ryelands or pure Spanish sheep.[257]
As some of our British parks are ancient, it occurred to me that there must have been long-continued close interbreeding with the fallow deer (Cervus dama) kept in them; but on inquiry I find that it is a common practice to infuse new blood by procuring bucks from other parks. Mr. Shirley,[258] who has carefully studied the management of deer, admits that in some parks there has been no admixture of foreign blood from a time beyond the memory of man. But he concludes "that in the end the constant breeding in-and-in is sure to tell to the disadvantage of the whole herd, though it may take a very long time to prove it; moreover, when we find, as is very constantly the case, that the introduction of fresh blood has been of the very greatest use to deer, both by improving their size and appearance, and particularly by being of service in removing the taint of 'rickback,' if not of other diseases, to which deer are sometimes subject when the blood has not been changed, there can, I think, be no doubt but that a judicious cross with a good stock is of the greatest consequence, and is indeed essential, sooner or later, to the prosperity of every well-ordered park."
Mr. Meynell's famous foxhounds have been adduced, as showing that no ill effects follow from close interbreeding; and Sir J. Sebright ascertained from him that he frequently bred from father and daughter, mother and {121} son, and sometimes even from brothers and sisters. Sir J. Sebright, however, declares,[259] that by breeding in-and-in, by which he means matching brothers and sisters, he has actually seen strong spaniels become weak and diminutive lapdogs. The Rev. W. D. Fox has communicated to me the case of a small lot of bloodhounds, long kept in the same family, which had become very bad breeders, and nearly all had a bony enlargement in the tail. A single cross with a distinct strain of bloodhounds restored their fertility, and drove away the tendency to malformation in the tail. I have heard the particulars of another case with bloodhounds, in which the female had to be held to the male. Considering how rapid is the natural increase of the dog, it is difficult to understand the high price of most highly improved breeds, which almost implies long-continued close interbreeding, except on the belief that this process lessens fertility and increases liability to distemper and other diseases. A high authority, Mr. Scrope, attributes the rarity and deterioration in size of the Scotch deerhound (the few individuals now existing throughout the country being all related) in large part to close interbreeding.
With all highly-bred animals there is more or less difficulty in getting them to procreate quickly, and all suffer much from delicacy of constitution; but I do not pretend that these effects ought to be wholly attributed to close interbreeding. A great judge of rabbits[260] says, "the long-eared does are often too highly bred or forced in their youth to be of much value as breeders, often turning out barren or bad mothers." Again: "Very long-eared bucks will also sometimes prove barren." These highly-bred rabbits often desert their young, so that it is necessary to have nurse-rabbits.
With Pigs there is more unanimity amongst breeders on the evil effects of close interbreeding than, perhaps, with any other large animal. Mr. Druce, a great and successful breeder of the Improved Oxfordshires (a crossed race), writes, "without a change of boars of a different tribe, but of the same breed, constitution cannot be preserved." Mr. Fisher Hobbs, the raiser of the celebrated Improved Essex breed, divided his stock into three separate families, by which means he maintained the breed for more than twenty years, "by judicious selection from the three distinct families."[261] Lord Western was the first importer of a Neapolitan boar and sow. "From this pair he bred in-and-in, until the breed was in danger of becoming extinct, a sure result (as Mr. Sidney remarks) of in-and-in breeding." Lord Western then crossed his Neapolitan pigs with the old Essex, and made the first great step towards the Improved Essex breed. Here is a more interesting case. Mr. J. Wright, well known as a breeder, crossed[262] the same boar with the daughter, granddaughter, and great-granddaughter, and so on for seven generations. The result was, that in many instances the offspring failed to breed; in others they produced few that lived; and of the latter many were idiotic, without sense {122} even to suck, and when attempting to move could not walk straight. Now it deserves especial notice, that the two last sows produced by this long course of interbreeding were sent to other boars, and they bore several litters of healthy pigs. The best sow in external appearance produced during the whole seven generations was one in the last stage of descent; but the litter consisted of this one sow. She would not breed to her sire, yet bred at the first trial to a stranger in blood. So that, in Mr. Wright's case, long-continued and extremely close interbreeding did not affect the external form or merit of the young; but with many of them the general constitution and mental powers, and especially the reproductive functions, were seriously affected.
Nathusius gives[263] an analogous and even more striking case: he imported from England a pregnant sow of the large Yorkshire breed, and bred the product closely in-and-in for three generations: the result was unfavourable, as the young were weak in constitution, with impaired fertility. One of the latest sows, which he esteemed a good animal, produced, when paired with her own uncle (who was known to be productive with sows of other breeds), a litter of six, and a second time a litter of only five weak young pigs. He then paired this sow with a boar of a small black breed, which he had likewise imported from England, and which boar, when matched with sows of his own breed, produced from seven to nine young: now, the sow of the large breed, which was so unproductive when paired with her own uncle, yielded to the small black boar, in the first litter twenty-one, and in the second litter eighteen young pigs; so that in one year she produced thirty-nine fine young animals!
As in the case of several other animals already mentioned, even when no injury is perceptible from moderately close interbreeding, yet, to quote the words of Mr. Coate, a most successful breeder (who five times won the annual gold medal of the Smithfield Club Show for the best pen of pigs), "Crosses answer well for profit to the farmer, as you get more constitution and quicker growth; but for me, who sell a great number of pigs for breeding purposes, I find it will not do, as it requires many years to get anything like purity of blood again."[264]
Before passing on to Birds, I ought to refer to man, though I am unwilling to enter on this subject, as it is surrounded by natural prejudices. It has moreover been discussed by various authors under many points of view.[265] Mr. Tylor[266] has shown {123} that with widely different races, in the most distant quarters of the world, marriages between relations--even between distant relations--have been strictly prohibited. A few exceptional cases can be specified, especially with royal families; and these have been enlarged on in a learned article[267] by Mr. W. Adam, and formerly in 1828 by Hofacker. Mr. Tylor is inclined to believe that the almost universal prohibition of closely-related marriages has arisen from their evil effects having been observed, and he ingeniously explains some apparent anomalies in the prohibition not extending equally to the relations on both the male and female side. He admits, however, that other causes, such as the extension of friendly alliances, may have come into play. Mr. W. Adam, on the other hand, concludes that related marriages are prohibited and viewed with repugnance from the confusion which would thus arise in the descent of property, and from other still more recondite reasons; but I cannot accept this view, seeing that the savages of Australia and South America,[268] who have no property to bequeath or fine moral feelings to confuse, hold the crime of incest in abhorrence.
It would be interesting to know, if it could be ascertained, as throwing light on this question with respect to man, what occurs with the higher anthropomorphous apes--whether the young males and females soon wander away from their parents, or whether the old males become jealous of their sons and expel them, or whether any inherited instinctive feeling, from being beneficial, has been generated, leading the young males and females of the same family to prefer pairing with distinct families, and to dislike pairing with each other. A considerable body of evidence has already been advanced, showing that the offspring from parents which are not related are more vigorous and fertile than those from parents which are closely related; hence any slight feeling, arising from the sexual excitement of novelty or other cause, which led to the former rather than to the latter unions, would be augmented through natural selection, and thus might become instinctive; for those individuals which had an innate preference of this kind would increase in number. It seems more probable, that degraded savages should {124} thus unconsciously have acquired their dislike and even abhorrence of incestuous marriages, rather than that they should have discovered by reasoning and observation the evil results. The abhorrence occasionally failing is no valid argument against the feeling being instinctive, for any instinct may occasionally fail or become vitiated, as sometimes occurs with parental love and the social sympathies. In the case of man, the question whether evil follows from close interbreeding will probably never be answered by direct evidence, as he propagates his kind so slowly and cannot be subjected to experiment; but the almost universal practice of all races at all times of avoiding closely-related marriages is an argument of considerable weight; and whatever conclusion we arrive at in regard to the higher animals may be safely extended to man.
Turning now to Birds: in the case of the Fowl a whole array of authorities could be given against too close interbreeding. Sir J. Sebright positively asserts that he made many trials, and that his fowls, when thus treated, became long in the legs, small in the body, and bad breeders.[269] He produced the famous Sebright Bantams by complicated crosses, and by breeding in-and-in; and since his time there has been much close interbreeding with these Bantams; and they are now notoriously bad breeders. I have seen Silver Bantams, directly descended from his stock, which had become almost as barren as hybrids; for not a single chicken had been that year hatched from two full nests of eggs. Mr. Hewitt says that with these Bantams the sterility of the male stands, with rare exceptions, in the closest relation with their loss of certain secondary male characters: he adds, "I have noticed, as a general rule, that even the slightest deviation from feminine character in the tail of the male Sebright--say the elongation by only half an inch of the two principal tail-feathers--brings with it improved probability of increased fertility."[270]
Mr. Wright states[271] that Mr. Clark, "whose fighting-cocks were so notorious, continued to breed from his own kind till they lost their disposition to fight, but stood to be cut up without making any resistance, and were so reduced in size as to be under those weights required for the best prizes; but on obtaining a cross from Mr. Leighton, they again resumed their former courage and weight." It should be borne in mind that game-cocks before they fought were always weighed, so that nothing was left to the imagination about any reduction or increase of {125} weight. Mr. Clark does not seem to have bred from brothers and sisters, which is the most injurious kind of union; and he found, after repeated trials, that there was a greater reduction in weight in the young from a father paired with his daughter, than from a mother with her son. I may add that Mr. Eyton, of Eyton, the well-known ornithologist, who is a large breeder of Grey Dorkings, informs me that they certainly diminish in size, and become less prolific, unless a cross with another strain is occasionally obtained. So it is with Malays, according to Mr. Hewitt, as far as size is concerned.[272]
An experienced writer[273] remarks that the same amateur, as is well known, seldom long maintains the superiority of his birds; and this, he adds, undoubtedly is due to all his stock "being of the same blood;" hence it is indispensable that he should occasionally procure a bird of another strain. But this is not necessary with those who keep a stock of fowls at different stations. Thus, Mr. Ballance, who has bred Malays for thirty years, and has won more prizes with these birds than any other fancier in England, says that breeding in-and-in does not necessarily cause deterioration; "but all depends upon how this is managed." "My plan has been to keep about five or six distinct runs, and to rear about two hundred or three hundred chickens each year, and select the best birds from each run for crossing. I thus secure sufficient crossing to prevent deterioration."[274]
We thus see that there is almost complete unanimity with poultry-breeders that, when fowls are kept at the same place, evil quickly follows from interbreeding carried on to an extent which would be disregarded in the case of most quadrupeds. On the other hand, it is a generally received opinion that cross-bred chickens are the hardiest and most easily reared.[275] Mr. Tegetmeier, who has carefully attended to poultry of all breeds, says[276] that Dorking hens, allowed to run with Houdan or Crevecoeur cocks, "produce in the early spring chickens that for size, hardihood, early maturity, and fitness for the market, surpass those of any pure breed that we have ever raised." Mr. Hewitt gives it as a general rule with fowls, that crossing the breed increases their size. He makes this remark after stating that hybrids from the pheasant and fowl are considerably larger than either progenitor: so again, hybrids from the male golden pheasant and hen common pheasant "are of far larger size than either parent-bird."[277] To this subject of the increased size of hybrids I shall presently return.
With Pigeons, breeders are unanimous, as previously stated, that it is absolutely indispensable, notwithstanding the trouble and expense thus caused, occasionally to cross their much-prized birds with individuals of another strain, but belonging, of course, to the same variety. It deserves {126} notice that, when large size is one of the desired characters, as with pouters,[278] the evil effects of close interbreeding are much sooner perceived than when small birds, such as short-faced tumblers, are valued. The extreme delicacy of the high fancy breeds, such as these tumblers and improved English carriers, is remarkable; they are liable to many diseases, and often die in the egg or during the first moult; and their eggs have generally to be hatched under foster-mothers. Although these highly-prized birds have invariably been subjected to much close interbreeding, yet their extreme delicacy of constitution cannot perhaps be thus fully explained. Mr. Yarrell informed me that Sir J. Sebright continued closely interbreeding some owl-pigeons, until from their extreme sterility he as nearly as possible lost the whole family. Mr. Brent[279] tried to raise a breed of trumpeters, by crossing a common pigeon, and recrossing the daughter, granddaughter, great-granddaughter, and great-great-granddaughter, with the same male trumpeter, until he obtained a bird with 15/16ths of trumpeter's blood; but then the experiment failed, for "breeding so close stopped reproduction." The experienced Neumeister[280] also asserts that the offspring from dovecotes and various other breeds are "generally very fertile and hardy birds:" so again, MM. Boitard and Corbié,[281] after forty-five years' experience, recommend persons to cross their breeds for amusement; for, if they fail to make interesting birds, they will succeed under an economical point of view, "as it is found that mongrels are more fertile than pigeons of pure race."
I will refer only to one other animal, namely, the Hive-bee, because a distinguished entomologist has advanced this as a case of inevitable close interbreeding. As the hive is tenanted by a single female, it might have been thought that her male and female offspring would always have bred together, more especially as bees of different hives are hostile to each other; a strange worker being almost always attacked when trying to enter another hive. But Mr. Tegetmeier has shown[282] that this instinct does not apply to drones, which are permitted to enter any hive; so that there is no à priori improbability of a queen receiving a foreign drone. The fact of the union invariably and necessarily taking place on the wing, during the queen's nuptial flight, seems to be a special provision against continued interbreeding. However this may be, experience has shown, since the introduction of the yellow-banded Ligurian race into Germany and England, that bees freely cross: Mr. Woodbury, who introduced Ligurian bees into Devonshire, found during a single season that three stocks, at distances of from one to two miles from his hives, were crossed by his drones. In one case the Ligurian drones must have flown over the city of Exeter, and over several intermediate hives. On another occasion several common black queens were crossed by Ligurian drones at a distance of from one to three and a half miles.[283]
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Plants.
When a single plant of a new species is introduced into any country, if propagated by seed, many individuals will soon be raised, so that if the proper insects be present there will be crossing. With newly-introduced trees or other plants not propagated by seed we are not here concerned. With old-established plants it is an almost universal practice occasionally to make exchanges of seed, by which means individuals which have been exposed to different conditions of life,--and this, as we have seen, diminishes the evil from close interbreeding,--will occasionally be introduced into each district.
Experiments have not been tried on the effects of fertilising flowers with their own pollen during several generations. But we shall presently see that certain plants, either normally or abnormally, are more or less sterile, even in the first generation, when fertilised by their own pollen. Although nothing is directly known on the evil effects of long-continued close interbreeding with plants, the converse proposition that great good is derived from crossing is well established.
With respect to the crossing of individuals belonging to the same sub-variety, Gärtner, whose accuracy and experience exceeded that of all other hybridisers, states[284] that he has many times observed good effects from this step, especially with exotic genera, of which the fertility is somewhat impaired, such as Passiflora, Lobelia, and Fuchsia. Herbert also says,[285] "I am inclined to think that I have derived advantage from impregnating the flower from which I wished to obtain seed with pollen from another individual of the same variety, or at least from another flower, rather than with its own." Again, Professor Lecoq asserts that he has ascertained that crossed offspring are more vigorous and robust than their parents.[286]
General statements of this kind, however, can seldom be fully trusted; consequently I have begun a series of experiments, which, if they continue to give the same results as hitherto, will for ever settle the question of the good effects of crossing two distinct plants of the same variety, and of the evil effects of self-fertilisation. A clear light will thus also be thrown on the fact that flowers are invariably constructed so as to permit, or favour, or necessitate the union of two individuals. We shall clearly understand why monoecious and dioecious,--why dimorphic and trimorphic plants exist, and many other such cases. The plan which I have followed in my experiments is to grow plants in the same pot, or in pots of the same size, or close together in the open ground; to carefully exclude insects; and then to fertilise some of the flowers with pollen from the same flower, and others on the same plant with pollen from a distinct but adjoining plant. In many, but not all, of these experiments, the crossed plants yielded much more seed than the self-fertilised plants; and I have never seen the {128} reversed case. The self-fertilised and crossed seeds thus obtained were allowed to germinate in the same glass vessel on damp sand; and as the seeds successively germinated, they were planted in pairs on opposite sides of the same pot, with a superficial partition between them, and were placed so as to be equally exposed to the light. In other cases the self-fertilised and crossed seeds were simply sown on opposite sides of the same small pot. I have, in short, followed different plans, but in every case have taken all the precautions which I could think of, so that the two lots should be equally favoured. Now, I have carefully observed the growth of plants raised from crossed and self-fertilised seed, from their germination to maturity, in species of the following genera, namely, Brassica, Lathyrus, Lupinus, Lobelia, Lactuca, Dianthus, Myosotis, Petunia, Linaria, Calceolaria, Mimulus, and Ipomoea, and the difference in their powers of growth, and of withstanding in certain cases unfavourable conditions, was most manifest and strongly marked. It is of importance that the two lots of seed should be sown or planted on opposite sides of the same pot, so that the seedlings may struggle against each other; for if sown separately in ample and good soil, there is often but little difference in their growth.
I will briefly describe the two most striking cases as yet observed by me. Six crossed and six self-fertilised seeds of Ipomoea purpurea, from plants treated in the manner above described, were planted as soon as they had germinated, in pairs on opposite sides of two pots, and rods of equal thickness were given them to twine up. Five of the crossed plants grew from the first more quickly than the opposed self-fertilised plants; the sixth, however, was weakly and was for a time beaten, but at last its sounder constitution prevailed and it shot ahead of its antagonist. As soon as each crossed plant reached the top of its seven-foot rod its fellow was measured, and the result was that, when the crossed plants were seven feet high, the self-fertilised had attained the average height of only five feet four and a half inches. The crossed plants flowered a little before, and more profusely than the self-fertilised plants. On opposite sides of another small pot a large number of crossed and self-fertilised seeds were sown, so that they had to struggle for bare existence; a single rod was given to each lot: here again the crossed plants showed from the first their advantage; they never quite reached the summit of the seven-foot rod, but relatively to the self-fertilised plants their average height was as seven feet to five feet two inches. The experiment was repeated in the two following generations with plants raised from the self-fertilised and crossed plants, treated in exactly the same manner, and with nearly the same result. In the second generation, the crossed plants, which were again crossed, produced 121 seed-capsules, whilst the self-fertilised plants, again self-fertilised, produced only 84 capsules.
Some flowers of the Mimulus luteus were fertilised with their own pollen, and others were crossed with pollen from distinct plants growing in the same pot. The seeds after germinating were thickly planted on opposite sides of a pot. The seedlings were at first equal in height; but when the young crossed plants were exactly half an inch, the {129} self-fertilised plants were only a quarter of an inch high. But this inequality did not continue, for, when the crossed plants were four and a half inches high, the self-fertilised were three inches; and they retained the same relative difference till their growth was complete. The crossed plants looked far more vigorous than the uncrossed, and flowered before them; they produced also a far greater number of flowers, which yielded capsules (judging, however, from only a few) containing more seeds. As in the former case, the experiment was repeated in the same manner during the next two generations, and with exactly the same result. Had I not watched these plants of the Mimulus and Ipomoea during their whole growth, I could not have believed it possible, that a difference apparently so slight, as that of the pollen being taken from the same flower, and from a distinct plant growing in the same small pot, could have made so wonderful a difference in the growth and vigour of the plants thus produced. This, under a physiological point of view, is a most remarkable phenomenon.
* * * * *
With respect to the benefit derived from crossing distinct varieties, plenty of evidence has been published. Sageret[287] repeatedly speaks in strong terms of the vigour of melons raised by crossing different varieties, and adds that they are more easily fertilised than common melons, and produce numerous good seed. Here follows the evidence of an English gardener:[288] "I have this summer met with better success in my cultivation of melons, in an unprotected state, from the seeds of hybrids (i.e. mongrels) obtained by cross impregnation, than with old varieties. The offspring of three different hybridisations (one more especially, of which the parents were the two most dissimilar varieties I could select) each yielded more ample and finer produce than any one of between twenty and thirty established varieties."
Andrew Knight[289] believed that his seedlings from crossed varieties of the apple exhibited increased vigour and luxuriance; and M. Chevreul[290] alludes to the extreme vigour of some of the crossed fruit-trees raised by Sageret.
By crossing reciprocally the tallest and shortest peas, Knight[291] says, "I had in this experiment a striking instance of the stimulative effects of crossing the breeds; for the smallest variety, whose height rarely exceeded two feet, was increased to six feet; whilst the height of the large and luxuriant kind was very little diminished." Mr. Laxton gave me seed-peas produced from crosses between four distinct kinds; and the plants thus raised were extraordinarily vigorous, being in each case from one to two or three feet taller than the parent-forms growing close alongside them.
{130}
Wiegmann[292] made many crosses between several varieties of cabbage; and he speaks with astonishment of the vigour and height of the mongrels, which excited the amazement of all the gardeners who beheld them. Mr. Chaundy raised a great number of mongrels by planting together six distinct varieties of cabbage. These mongrels displayed an infinite diversity of character; "But the most remarkable circumstance was, that, while all the other cabbages and borecoles in the nursery were destroyed by a severe winter, these hybrids were little injured, and supplied the kitchen when there was no other cabbage to be had."
Mr. Maund exhibited before the Royal Agricultural Society[293] specimens of crossed wheat, together with their parent varieties; and the editor states that they were intermediate in character, "united with that greater vigour of growth, which it appears, in the vegetable as in the animal world, is the result of a first cross." Knight also crossed several varieties of wheat,[294] and he says "that in the years 1795 and 1796, when almost the whole crop of corn in the island was blighted, the varieties thus obtained, and these only, escaped in this neighbourhood, though sown in several different soils and situations."
Here is a remarkable case: M. Clotzsch[295] crossed Pinus sylvestris and nigricans, Quercus robur and pedunculata, Alnus glutinosa and incana, Ulmus campestris and effusa; and the cross-fertilised seeds, as well as seeds of the pure parent-trees, were all sown at the same time and in the same place. The result was, that after an interval of eight years, the hybrids were one-third taller than the pure trees!
* * * * *
The facts above given refer to undoubted varieties, excepting the trees crossed by Clotzsch, which are ranked by various botanists as strongly-marked races, sub-species, or species. That true hybrids raised from entirely distinct species, though they lose in fertility, often gain in size and constitutional vigour, is certain. It would be superfluous to quote any facts; for all experimenters, Kölreuter, Gärtner, Herbert, Sageret, Lecoq, and Naudin, have been struck with the wonderful vigour, height, size, tenacity of life, precocity, and hardiness of their hybrid productions. Gärtner[296] sums up his conviction on this head in the strongest terms. Kölreuter[297] gives numerous precise measurements of the weight and height of his hybrids in comparison with measurements of both parent-forms; and speaks with astonishment of their "statura portentosa," their "ambitus vastissimus ac altitudo valde conspicua." Some exceptions to the rule in the case of very sterile hybrids have, however, been noticed by Gärtner and {131} Herbert; but the most striking exceptions are given by Max Wichura,[298] who found that hybrid willows were generally tender in constitution, dwarf, and short-lived.
Kölreuter explains the vast increase in the size of the roots, stems, &c., of his hybrids, as the result of a sort of compensation due to their sterility, in the same way as many emasculated animals are larger than the perfect males. This view seems at first sight extremely probable, and has been accepted by various authors;[299] but Gärtner[300] has well remarked that there is much difficulty in fully admitting it; for with many hybrids there is no parallelism between the degree of their sterility and their increased size and vigour. The most striking instances of luxuriant growth have been observed with hybrids which were not sterile in any extreme degree. In the genus Mirabilis, certain hybrids are unusually fertile, and their extraordinary luxuriance of growth, together with their enormous roots,[301] have been transmitted to their progeny. The increased size of the hybrids produced between the fowl and pheasant, and between the distinct species of pheasants, has been already noticed. The result in all cases is probably in part due to the saving of nutriment and vital force through the sexual organs not acting, or acting imperfectly, but more especially to the general law of good being derived from a cross. For it deserves especial attention that mongrel animals and plants, which are so far from being sterile that their fertility is often actually augmented, have, as previously shown, their size, hardiness, and constitutional vigour generally increased. It is not a little remarkable that an accession of vigour and size should thus arise under the opposite contingencies of increased and diminished fertility.
It is a perfectly well ascertained fact[302] that hybrids will invariably breed more readily with either pure parent, and not rarely with a distinct species, than with each other. Herbert is inclined to explain even this fact by the advantage derived from a cross; but Gärtner more justly accounts for it by the pollen of the hybrid, and probably its ovules, being in some degree vitiated, whereas the pollen and ovules of both pure parents and of any third species are sound. Nevertheless there are some well-ascertained and remarkable facts, which, as we shall immediately see, show that the act of crossing in itself undoubtedly tends to increase or re-establish the fertility of hybrids.
On certain Hermaphrodite Plants which, either normally or abnormally, require to be fertilised by pollen from a distinct individual or species.
The facts now to be given differ from those hitherto detailed, as the self-sterility does not here result from long-continued, {132} close interbreeding. These facts are, however, connected with our present subject, because a cross with a distinct individual is shown to be either necessary or advantageous. Dimorphic and trimorphic plants, though they are hermaphrodites, must be reciprocally crossed, one set of forms by the other, in order to be fully fertile, and in some cases to be fertile in any degree. But I should not have noticed these plants, had it not been for the following cases given by Dr. Hildebrand:[303]--
Primula sinensis is a reciprocally dimorphic species: Dr. Hildebrand fertilised twenty-eight flowers of both forms, each by pollen of the other form, and obtained the full number of capsules containing on an average 42.7 seed per capsule; here we have complete and normal fertility. He then fertilised forty-two flowers of both forms with pollen of the same form, but taken from a distinct plant, and all produced capsules containing on an average only 19.6 seed. Lastly, and here we come to our more immediate point, he fertilised forty-eight flowers of both forms with pollen of the same form, taken from the same flower, and now he obtained only thirty-two capsules, and these contained on an average 18.6 seed, or one less per capsule than in the former case. So that, with these illegitimate unions, the act of impregnation is less assured, and the fertility slightly less, when the pollen and ovules belong to the same flower, than when belonging to two distinct individuals of the same form. Dr. Hildebrand has recently made analogous experiments on the long-styled form of Oxalis rosea, with the same result.[304]
It has recently been discovered that certain plants, whilst growing in their native country under natural conditions, cannot be fertilised with pollen from the same plant. They are sometimes so utterly self-impotent, that, though they can readily be fertilised by the pollen of a distinct species or even distinct genus, yet, wonderful as the fact is, they never produce a single seed by their own pollen. In some cases, moreover, the plant's own pollen and stigma mutually act on each other in a deleterious manner. Most of the facts to be given relate to Orchids, but I will commence with a plant belonging to a widely different family.
Sixty-three flowers of Corydalis cava, borne on distinct plants, were fertilised by Dr. Hildebrand[305] with pollen from other plants of the same species; and fifty-eight capsules were obtained, including on an average {133} 4.5 seed in each. He then fertilised sixteen flowers produced by the same raceme, one with another, but obtained only three capsules, one of which alone contained any good seeds, namely, two in number. Lastly, he fertilised twenty-seven flowers, each with its own pollen; he left also fifty-seven flowers to be spontaneously fertilised, and this would certainly have ensued if it had been possible, for the anthers not only touch the stigma, but the pollen-tubes were seen by Dr. Hildebrand to penetrate it; nevertheless these eighty-four flowers did not produce a single seed-capsule! This whole case is highly instructive, as it shows how widely different the action of the same pollen is, according as it is placed on the stigma of the same flower, or on that of another flower on the same raceme, or on that of a distinct plant.
With exotic Orchids several analogous cases have been observed, chiefly by Mr. John Scott.[306] Oncidium sphacelatum has effective pollen, for with it Mr. Scott fertilised two distinct species; its ovules are likewise capable of impregnation, for they were readily fertilised by the pollen of O. divaricatum; nevertheless, between one and two hundred flowers fertilised by their own pollen did not produce a single capsule, though the stigmas were penetrated by the pollen-tubes. Mr. Robinson Munro, of the Royal Botanic Gardens of Edinburgh, also informs me (1864) that a hundred and twenty flowers of this same species were fertilised by him with their own pollen, and did not produce a capsule, but eight flowers fertilised by the pollen of O. divaricatum produced four fine capsules: again, between two and three hundred flowers of O. divaricatum, fertilised by their own pollen, did not set a capsule, but twelve flowers fertilised by O. flexuosum produced eight fine capsules: so that here we have three utterly self-impotent species, with their male and female organs perfect, as shown by their mutual fertilisation. In these cases fertilisation was effected only by the aid of a distinct species. But, as we shall presently see, distinct plants, raised from seed, of Oncidium flexuosum, and probably of the other species, would have been perfectly capable of fertilising each other, for this is the natural process. Again, Mr. Scott found that the pollen of a plant of O. microchilum was good, for with it he fertilised two distinct species; he found its ovules good, for they could be fertilised by the pollen of one of these species, and by the pollen of a distinct plant of O. microchilum; but they could not be fertilised by pollen of the same plant, though the pollen-tubes penetrated the stigma. An analogous case has been recorded by M. Rivière,[307] with two plants of O. Cavendishianum, which were both self-sterile, but reciprocally fertilised each other. All these cases refer to the genus Oncidium, but Mr. Scott found that Maxillaria atro-rubens was "totally insusceptible of fertilisation with its own pollen," but fertilised, and was fertilised by, a widely distinct species, viz. M. squalens.
As these orchids had grown under unnatural conditions, in {134} hot-houses, I concluded without hesitation that their self-sterility was due to this cause. But Fritz Müller informs me that at Desterro, in Brazil, he fertilised above one hundred flowers of the above-mentioned Oncidium flexuosum, which is there endemic, with its own pollen, and with that taken from distinct plants; all the former were sterile, whilst those fertilised by pollen from any other plant of the same species were fertile. During the first three days there was no difference in the action of the two kinds of pollen: that placed on the stigma of the same plant separated in the usual manner into grains, and emitted tubes which penetrated the column, and the stigmatic chamber shut itself; but the flowers alone which had been fertilised by pollen taken from a distinct plant produced seed-capsules. On a subsequent occasion these experiments were repeated on a large scale with the same result. Fritz Müller found that four other endemic species of Oncidium were in like manner utterly sterile with their own pollen, but fertile with that from any other plant: some of them likewise produced seed-capsules when impregnated with pollen of widely distinct genera, such as Leptotes, Cyrtopodium, and Rodriguezia! Oncidium crispum, however, differs from the foregoing species in varying much in its self-sterility; some plants producing fine pods with their own pollen, others failing to do so; in two or three instances, Fritz Müller observed that the pods produced by pollen taken from a distinct flower on the same plant, were larger than those produced by the flower's own pollen. In Epidendrum cinnabarinum, an orchid belonging to another division of the family, fine pods were produced by the plant's own pollen, but they contained by weight only about half as much seed as the capsules which had been fertilized by pollen from a distinct plant, and in one instance from a distinct species; moreover, a very large proportion, and in some cases nearly all the seed produced by the plant's own pollen, was embryonless and worthless. Some self-fertilized capsules of a Maxillaria were in a similar state.
Another observation made by Fritz Müller is highly remarkable, namely, that with various orchids the plant's own pollen not only fails to impregnate the flower, but acts on the stigma, and is acted on, in an injurious or poisonous manner. This is shown by the surface of the stigma in contact with the pollen, and by the pollen itself, becoming in from three to five days dark brown, and then decaying. The discolouration and decay are not caused by parasitic cryptogams, which were observed by Fritz Müller in only a single instance. These changes are well shown by placing on the same stigma, at the same time, the plant's own pollen and that from a distinct plant of the same species, or of another species, or even of another and widely remote genus. Thus, on the stigma of Oncidium flexuosum, the plant's own pollen and that from a distinct plant were placed side by side, and in five days' time the latter was perfectly fresh, whilst the plant's own pollen was brown. On the other hand, when the pollen of a distinct plant of the Oncidium flexuosum, and of the Epidendrum zebra (nov. spec.?), were placed together on the same stigma, they behaved in exactly the same manner, the grains separating, emitting tubes, and penetrating the stigma, so that the two {135} pollen-masses, after an interval of eleven days, could not be distinguished except by the difference of their caudicles, which, of course, undergo no change. Fritz Müller has, moreover, made a large number of crosses between orchids belonging to distinct species and genera, and he finds that in all cases when the flowers are not fertilised their footstalks first begin to wither; and the withering slowly spreads upwards until the germens fall off, after an interval of one or two weeks, and in one instance of between six and seven weeks; but even in this latter case, and in most other cases, the pollen and stigma remained in appearance fresh. Occasionally, however, the pollen becomes brownish, generally on the external surface, and not in contact with the stigma, as is invariably the case when the plant's own pollen is applied.
Fritz Müller observed the poisonous action of the plant's own pollen in the above-mentioned Oncidium flexuosum, O. unicorne, pubes (?), and in two other unnamed species. Also in two species of Rodriguezia, in two of Notylia, in one of Burlingtonia, and of a fourth genus in the same group. In all these cases, except the last, it was proved that the flowers were, as might have been expected, fertile with pollen from a distinct plant of the same species. Numerous flowers of one species of Notylia were fertilized with pollen from the same raceme; in two days' time they all withered, the germens began to shrink, the pollen-masses became dark brown, and not one pollen-grain emitted a tube. So that in this orchid the injurious action of the plant's own pollen is more rapid than with Oncidium flexuosum. Eight other flowers on the same raceme were fertilized with pollen from a distinct plant of the same species: two of these were dissected, and their stigmas were found to be penetrated by numberless pollen-tubes; and the germens of the other six flowers became well developed. On a subsequent occasion many other flowers were fertilized with their own pollen, and all fell off dead in a few days; whilst some flowers on the same raceme which had been left simply unfertilised adhered and long remained fresh. We have seen that in cross-unions between extremely distinct orchids the pollen long remains undecayed; but Notylia behaved in this respect differently; for when its pollen was placed on the stigma of Oncidium flexuosum, both the stigma and pollen quickly became dark brown, in the same manner as if the plant's own pollen had been applied.
Fritz Müller suggests that, as in all these cases the plant's own pollen is not only impotent (thus effectually preventing self-fertilization), but likewise prevents, as was ascertained in the case of the Notylia and Oncidium flexuosum, the action of subsequently applied pollen from a distinct individual, it would be an advantage to the plant to have its own pollen rendered more and more deleterious; for the germens would thus quickly be killed, and, dropping off, there would be no further waste in nourishing a part which ultimately could be of no avail. Fritz Müller's discovery that a plant's own pollen and stigma in some cases act on each other as if mutually poisonous, is certainly most remarkable.
We now come to cases closely analogous with those just {136} given, but different, inasmuch as individual plants alone of the species are self-impotent. This self-impotence does not depend on the pollen or ovules being in a state unfit for fertilisation, for both have been found effective in union with other plants of the same or of a distinct species. The fact of these plants having spontaneously acquired so peculiar a constitution, that they can be fertilised more readily by the pollen of a distinct species than by their own, is remarkable. These abnormal cases, as well as the foregoing normal cases, in which certain orchids, for instance, can be much more easily fertilised by the pollen of a distinct species than by their own, are exactly the reverse of what occurs with all ordinary species. For in these latter the two sexual elements of the same individual plant are capable of freely acting on each other; but are so constituted that they are more or less impotent when brought into union with the sexual elements of a distinct species, and produce more or less sterile hybrids. It would appear that the pollen or ovules, or both, of the individual plants which are in this abnormal state, have been affected in some strange manner by the conditions to which they themselves or their parents have been exposed; but whilst thus rendered self-sterile, they have retained the capacity common to most species of partially fertilizing and being partially fertilized by allied forms. However this may be, the subject, to a certain extent, is related to our general conclusion that good is derived from the act of crossing.
Gärtner experimented on two plants of Lobelia fulgens, brought from separate places, and found[308] that their pollen was good, for he fertilised with it L. cardinalis and syphilitica; their ovules were likewise good, for they were fertilised by the pollen of these same two species; but these two plants of L. fulgens could not be fertilised by their own pollen, as can generally be effected with perfect ease with this species. Again, the pollen of a plant of Verbascum nigrum grown in a pot was found by Gärtner[309] capable of fertilising V. lychnitis and V. Austriacum; the ovules could be fertilised by the pollen of V. thapsus; but the flowers could not be fertilised by their own pollen. Kölreuter, also,[310] gives the case of three {137} garden plants of Verbascum phoeniceum, which bore during two years many flowers; these he successfully fertilised by the pollen of no less than four distinct species, but they produced not a seed with their own apparently good pollen; subsequently these same plants, and others raised from seed, assumed a strangely fluctuating condition, being temporarily sterile on the male or female side, or on both sides, and sometimes fertile on both sides; but two of the plants were perfectly fertile throughout the summer.
It appears[311] that certain flowers on certain plants of Lilium candidum can be fertilised more easily by pollen from a distinct individual than by their own. So, again, with the varieties of the potato. Tinzmann,[312] who made many trials with this plant, says that pollen from another variety sometimes "exerts a powerful influence, and I have found sorts of potatoes which would not bear seed from impregnation with the pollen of their own flowers, would bear it when impregnated with other pollen." It does not, however, appear to have been proved that the pollen which failed to act on the flower's own stigma was in itself good.
In the genus Passiflora it has long been known that several species do not produce fruit, unless fertilised by pollen taken from distinct species: thus, Mr. Mowbray[313] found that he could not get fruit from P. alata and racemosa except by reciprocally fertilising them with each other's pollen. Similar facts have been observed in Germany and France;[314] and I have received two authentic accounts of P. quadrangularis, which never produced fruit with its own pollen, but would do so freely when fertilised in one case with the pollen of P. coerulea, and in another case with that of P. edulis. So again, with respect to P. laurifolia, a cultivator of much experience has recently remarked[315] that the flowers "must be fertilised with the pollen of P. coerulea, or of some other common kind, as their own pollen will not fertilise them." But the fullest details on this subject have been given by Mr. Scott:[316] plants of Passiflora racemosa, coerulea, and alata flowered profusely during many years in the Botanic Gardens of Edinburgh, and, though repeatedly fertilised by Mr. Scott and by others with their own pollen, never produced any seed; yet this occurred at once with all three species when they were crossed together in various ways. But in the case of P. coerulea, three plants, two of which grew in the Botanic Gardens, were all rendered fertile, merely by impregnating the one with pollen of the other. The same result was attained in the same manner with P. alata, but only with one plant out of three. As so many self-sterile species have been mentioned, it may be stated that in the case of P. gracilis, which is an annual, the flowers are nearly as fertile with their own pollen as with that from a distinct plant; thus sixteen flowers {138} spontaneously self-fertilised produced fruit, each containing on an average 21.3 seed, whilst fruit from fourteen crossed flowers contained 24.1 seed.
Returning to P. alata, I have received (1866) some interesting details from Mr. Robinson Munro. Three plants, including one in England, have already been mentioned which were inveterately self-sterile, and Mr. Munro informs me of several others which, after repeated trials during many years, have been found in the same predicament. At some other places, however, this species fruits readily when fertilised with its own pollen. At Taymouth Castle there is a plant which was formerly grafted by Mr. Donaldson on a distinct species, name unknown, and ever since the operation it has produced fruit in abundance by its own pollen; so that this small and unnatural change in the state of this plant has restored its self-fertility! Some of the seedlings from the Taymouth Castle plant were found to be not only sterile with their own pollen, but with each other's pollen, and with the pollen of distinct species. Pollen from the Taymouth plant failed to fertilise certain plants of the same species, but was successful on one plant in the Edinburgh Botanic Gardens. Seedlings were raised from this latter union, and some of their flowers were fertilised by Mr. Munro with their own pollen; but they were found to be as self-impotent as the mother-plant had always proved, except when fertilised by the grafted Taymouth plant, and except, as we shall see, when fertilised by her own seedlings. For Mr. Munro fertilised eighteen flowers on the self-impotent mother-plant with pollen from these her own self-impotent seedlings, and obtained, remarkable as the fact is, eighteen fine capsules full of excellent seed! I have met with no case in regard to plants which shows so well as this of P. alata, on what small and mysterious causes complete fertility or complete sterility depends.
The facts hitherto given relate to the much-lessened or completely destroyed fertility of pure species when impregnated with their own pollen, in comparison with their fertility when impregnated by distinct individuals or distinct species; but closely analogous facts have been observed with hybrids.
Herbert states[317] that having in flower at the same time nine hybrid Hippeastrums, of complicated origin, descended from several species, he found that "almost every flower touched with pollen from another cross produced seed abundantly, and those which were touched with their own pollen either failed entirely, or formed slowly a pod of inferior size, with fewer seeds." In the 'Horticultural Journal' he adds that, "the admission of the pollen of another cross-bred Hippeastrum (however complicated the cross) to any one flower of the number, is almost sure to check the fructification of the others." In a letter written to me in 1839, Dr. Herbert says that he had already tried these experiments during five consecutive years, and he subsequently repeated them, with the same invariable result. {139} He was thus led to make an analogous trial on a pure species, namely, on the Hippeastrum aulicum, which he had lately imported from Brazil: this bulb produced four flowers, three of which were fertilised by their own pollen, and the fourth by the pollen of a triple cross between H. bulbulosum, reginæ, and vittatum; the result was, that "the ovaries of the three first flowers soon ceased to grow, and after a few days perished entirely: whereas the pod impregnated by the hybrid made vigorous and rapid progress to maturity, and bore good seed, which vegetated freely." This is, indeed, as Herbert remarks, "a strange truth," but not so strange as it then appeared.
As a confirmation of these statements, I may add that Mr. M. Mayes,[318] after much experience in crossing the species of Amaryllis (Hippeastrum), says, "neither the species nor the hybrids will, we are well aware, produce seed so abundantly from their own pollen as from that of others." So, again, Mr. Bidwell, in New South Wales,[319] asserts that Amaryllis belladonna bears many more seeds when fertilised by the pollen of Brunswigia (Amaryllis of some authors) Josephinæ or of B. multiflora, than when fertilised by its own pollen. Mr. Beaton dusted four flowers of a Cyrtanthus with their own pollen, and four with the pollen of Vallota (Amaryllis) purpurea; on the seventh day "those which received their own pollen slackened their growth, and ultimately perished; those which were crossed with the Vallota held on."[320] These latter cases, however, relate to uncrossed species, like those before given with respect to Passiflora, Orchids, &c., and are here referred to only because the plants belong to the same group of Amaryllidaceæ.
In the experiments on the hybrid Hippeastrums, if Herbert had found that the pollen of two or three kinds alone had been more efficient on certain kinds than their own pollen, it might have been argued that these, from their mixed parentage, had a closer mutual affinity than the others; but this explanation is inadmissible, for the trials were made reciprocally backwards and forwards on nine different hybrids; and a cross, whichever way taken, always proved highly beneficial. I can add a striking and analogous case from experiments made by the Rev. A. Rawson, of Bromley Common, with some complex hybrids of Gladiolus. This skilful horticulturist possessed a number of French varieties, differing from each other only in the colour and size of the flowers, all descended from Gandavensis, a well-known old hybrid, said to be descended from G. Natalensis by the pollen of G. oppositiflorus.[321] Mr. Rawson, after repeated trials, found that none of the varieties would set seed with their own pollen, although {140} taken from distinct plants of the same variety, which had, of course, been propagated by bulbs, but that they all seeded freely with pollen from any other variety. To give two examples: Ophir did not produce a capsule with its own pollen, but when fertilised with that of Janire, Brenchleyensis, Vulcain, and Linné, it produced ten fine capsules; but the pollen of Ophir was good, for when Linné was fertilised by it seven capsules were produced. This later variety, on the other hand, was utterly barren with its own pollen, which we have seen was perfectly efficient on Ophir. Altogether, Mr. Rawson, in the year 1861, fertilised twenty-six flowers borne by four varieties with pollen taken from other varieties, and every single flower produced a fine seed-capsule; whereas fifty-two flowers on the same plants, fertilised at the same time with their own pollen, did not yield a single seed-capsule. Mr. Rawson fertilised, in some cases, the alternate flowers, and in other cases all those down one side of the spike, with pollen of other varieties, and the remaining flowers with their own pollen; I saw these plants when the capsules were nearly mature, and their curious arrangement at once brought full conviction to the mind that an immense advantage had been derived from crossing these hybrids.
Lastly, I have heard from Dr. E. Bornet, of Antibes, who has made numerous experiments in crossing the species of Cistus, but as not yet published the results, that, when any of these hybrids are fertile, they may be said to be, in regard to function, dioecious; "for the flowers are always sterile when the pistil is fertilised by pollen taken from the same flower or from flowers on the same plant. But they are often fertile if pollen be employed from a distinct individual of the same hybrid nature, or from a hybrid made by a reciprocal cross."
Conclusion.--The facts just given, which show that certain plants are self-sterile, although both sexual elements are in a fit state for reproduction when united with distinct individuals of the same or other species, appear at first sight opposed to all analogy. The sexual elements of the same flower have become, as already remarked, differentiated in relation to each other, almost like those of two distinct species.
With respect to the species which, whilst living under their natural conditions, have their reproductive organs in this peculiar state, we may conclude that it has been naturally acquired for the sake of effectually preventing self-fertilisation. The case is closely analous with dimorphic and trimorphic plants, which can be fully fertilised only by plants belong to the opposite form, and not, as in the foregoing cases, in differently by any other plant. Some of these dimorphic plants are completely sterile with pollen taken from the same plant or from the same {141} form. It is interesting to observe the graduated series from plants which, when fertilised by their own pollen, yield the full number of seed, but with the seedlings a little dwarfed in stature--to plants which when self-fertilised yield few seeds--to those with yield none--and, lastly, to those in which the plant's own pollen and stigma act on each other like poison. This peculiar state of the reproductive organs, when occurring in certain individuals alone, is evidently abnormal; and as it chiefly affects exotic plants, or indigenous plants cultivated in pots, we may attribute it to some change in the conditions of life, acting on the plants themselves or on their parents. The self-impotent Passiflora alata, which recovered its self-fertility after having been grafted on a distinct stock, shows how small a change is sufficient to act powerfully on the reproductive system. The possibility of a plant becoming under culture self-impotent is interesting as throwing light on the occurrence of this same condition in natural species. A cultivated plant in this state generally remains so during its whole life; and from this fact we may infer that the state is probably congenital.
Kölreuter, however, has described some plants of Verbascum which varied in this respect even during the same season. As in all the normal cases, and in many, probably in most, of the abnormal cases, any two self-impotent plants can reciprocally fertilize each other, we may infer that a very slight difference in the nature of their sexual elements suffices to give fertility; but in other instances, as with some Passifloras and the hybrid Gladioli, a greater degree of differentiation appears to be necessary, for with these plants fertility is gained only by the union of distinct species, or of hybrids of distinct parentage. These facts all point to the same general conclusion, namely, that good is derived from a cross between individuals, which either innately, or from exposure to dissimilar conditions, have come to differ in sexual constitution.
Exotic animals confined in menageries are sometimes in nearly the same state as the above-described self-impotent plants; for, as we shall see in the following chapter, certain monkeys, the larger carnivora, several finches, geese, and pheasants, cross together, quite as freely as, or even more freely than, the individuals of the same species breed together. Cases will, {142} also, be given of sexual incompatibility between certain male and female domesticated animals, which, nevertheless, are fertile when matched with any other individual of the same kind.
In the early part of this chapter it was shown that the crossing of distinct forms, whether closely or distantly allied, gives increased size and constitutional vigour, and, except in the case of crossed species, increased fertility, to the offspring. The evidence rests on the universal testimony of breeders (for it should be observed that I am not here speaking of the evil results of close interbreeding), and is practically exemplified in the higher value of cross-bred animals for immediate consumption. The good results of crossing have also been demonstrated, in the case of some animals and of numerous plants, by actual weight and measurement. Although animals of pure blood will obviously be deteriorated by crossing, as far as their characteristic qualities are concerned, there seems to be no exception to the rule that advantages of the kind just mentioned are thus gained, even when there has not been any previous close interbreeding. The rule applies to all animals, even to cattle and sheep, which can long resist breeding in-and-in between the nearest blood-relations. It applies to individuals of the same sub-variety but of distinct families, to varieties or races, to sub-species, as well as to quite distinct species.
In this latter case, however, whilst size, vigour, precocity, and hardiness are, with rare exceptions, gained, fertility, in a greater or less degree, is lost; but the gain cannot be exclusively attributed to the principle of compensation; for there is no close parallelism between the increased size and vigour of the offspring and their sterility. Moreover it has been clearly proved that mongrels which are perfectly fertile gain these same advantages as well as sterile hybrids.
The evil consequences of long-continued close interbreeding are not so easily recognised as the good effects from crossing, for the deterioration is gradual. Nevertheless it is the general opinion of those who have had most experience, especially with animals which propagate quickly, that evil does inevitably follow sooner or later, but at different rates with different animals. No doubt a false belief may widely prevail like a superstition; yet it is difficult to suppose that so many acute and original {143} observers have all been deceived at the expense of much cost and trouble. A male animal may sometimes be paired with his daughter, granddaughter, and so on, even for seven generations, without any manifest bad result; but the experiment has never been tried of matching brothers and sisters, which is considered the closest form of interbreeding, for an equal number of generations. There is good reason to believe that by keeping the members of the same family in distinct bodies, especially if exposed to somewhat different conditions of life, and by occasionally crossing these families, the evil results may be much diminished, or quite eliminated. These results are loss of constitutional vigour, size, and fertility; but there is no necessary deterioration in the general form of the body, or in other good qualities. We have seen that with pigs first-rate animals have been produced after long-continued close interbreeding, though they had become extremely infertile when paired with their near relations. The loss of fertility, when it occurs, seems never to be absolute, but only relative to animals of the same blood; so that this sterility is to a certain extent analogous with that of self-impotent plants which cannot be fertilised by their own pollen, but are perfectly fertile with pollen of any other plant of the same species. The fact of infertility of this peculiar nature being one of the results of long-continued interbreeding, shows that interbreeding does not act merely by combining and augmenting various morbid tendencies common to both parents; for animals with such tendencies, if not at the time actually ill, can generally propagate their kind. Although offspring descended from the nearest blood-relations are not necessarily deteriorated in structure, yet some authors[322] believe that they are eminently liable to malformations; and this is not improbable, as everything which lessens the vital powers acts in this manner. Instances of this kind have been recorded in the case of pigs, bloodhounds, and some other animals.
Finally, when we consider the various facts now given which plainly show that good follows from crossing, and less plainly {144} that evil follows from close interbreeding, and when we bear in mind that throughout the whole organic world elaborate provision has been made for the occasional union of distinct individuals, the existence of a great law of nature is, if not proved, at least rendered in the highest degree probable; namely, that the crossing of animals and plants which are not closely related to beach other is highly beneficial or even necessary, and that interbreeding prolonged during many generations is highly injurious.
* * * * *
{145}
Chapter XVIII
ON THE ADVANTAGES AND DISADVANTAGES OF CHANGED CONDITIONS OF LIFE: STERILITY FROM VARIOUS CAUSES.
ON THE GOOD DERIVED FROM SLIGHT CHANGES IN THE CONDITIONS OF LIFE--STERILITY FROM CHANGED CONDITIONS, IN ANIMALS, IN THEIR NATIVE COUNTRY AND IN MENAGERIES--MAMMALS, BIRDS, AND INSECTS--LOSS OF SECONDARY SEXUAL CHARACTERS AND OF INSTINCTS--CAUSES OF STERILITY--STERILITY OF DOMESTICATED ANIMALS FROM CHANGED CONDITIONS--SEXUAL INCOMPATIBILITY OF INDIVIDUAL ANIMALS--STERILITY OF PLANTS FROM CHANGED CONDITIONS OF LIFE--CONTABESCENCE OF THE ANTHERS--MONSTROSITIES AS A CAUSE OF STERILITY--DOUBLE FLOWERS--SEEDLESS FRUIT--STERILITY FROM THE EXCESSIVE DEVELOPMENT OF THE ORGANS OF VEGETATION--FROM LONG-CONTINUED PROPAGATION BY BUDS--INCIPIENT STERILITY THE PRIMARY CAUSE OF DOUBLE FLOWERS AND SEEDLESS FRUIT.
On the Good derived from slight Changes in the Conditions of Life.--In considering whether any facts were known which might throw light on the conclusion arrived at in the last chapter, namely, that benefits ensue from crossing, and that it is a law of nature that all organic beings should occasionally cross, it appeared to me probable that the good derived from slight changes in the conditions of life, from being an analogous phenomenon, might serve this purpose. No two individuals, and still less no two varieties, are absolutely alike in constitution and structure; and when the germ of one is fertilised by the male element of another, we may believe that it is acted on in a somewhat similar manner as an individual when exposed to slightly changed conditions. Now, every one must have observed the remarkable influence on convalescents of a change of residence, and no medical man doubts the truth of this fact. Small farmers who hold but little land are convinced that their cattle derive great benefit from a change of pasture. In the case of plants, the evidence is strong that a great advantage is derived from exchanging seeds, tubers, bulbs, and cuttings from one soil or place to another as different as possible. {146}
The belief that plants are thus benefited, whether or not well founded, has been firmly maintained from the time of Columella, who wrote shortly after the Christian era, to the present day; and it now prevails in England, France, and Germany.[323] A sagacious observer, Bradley, writing in 1724,[324] says, "When we once become Masters of a good Sort of Seed, we should at least put it into Two or Three Hands, where the Soils and Situations are as different as possible; and every Year the Parties should change with one another; by which Means, I find the Goodness of the Seed will be maintained for several Years. For Want of this Use many Farmers have failed in their Crops and been great Losers." He then gives his own practical experience on this head. A modern writer[325] asserts, "Nothing can be more clearly established in agriculture than that the continual growth of any one variety in the same district makes it liable to deterioration either in quality or quantity." Another writer states that he sowed close together in the same field two lots of wheat-seed, the product of the same original stock, one of which had been grown on the same land, and the other at a distance, and the difference in favour of the crop from the latter seed was remarkable. A gentleman in Surrey who has long made it his business to raise wheat to sell for seed, and who has constantly realised in the market higher prices than others, assures me that he finds it indispensable continually to change his seed; and that for this purpose he keeps two farms differing much in soil and elevation.
With respect to the tubers of the potato, I find that at the present day the practice of exchanging sets is almost everywhere followed. The great growers of potatoes in Lancashire formerly used to get tubers from Scotland, but they found that "a change from the moss-lands, and vice versâ, was generally sufficient." In former times in France the crop of potatoes in the Vosges had become reduced in the course of fifty or sixty years in the proportion from 120-150 to 30-40 bushels; and the famous Oberlin attributed the surprising good which he effected in large part to changing the sets.[326]
A well-known practical gardener, Mr. Robson[327] positively states that he has himself witnessed decided advantage from obtaining bulbs of the onion, tubers of the potato, and various seeds, all of the same kind, from different soils and distant parts of England. He further states that with {147} plants propagated by cuttings, as with the Pelargonium, and especially the Dahlia, manifest advantage is derived from getting plans of the same variety, which have been cultivated in another place; or, "where the extent of the place allows, to take cuttings from one description of soil to plant on another, so as to afford the change that seems so necessary to the well-being of the plants." He maintains that after a time an exchange of this nature is "forced on the grower, whether he be prepared for it or not." Similar remarks have been made by another excellent gardener, Mr. Fish, namely, that cuttings of the same variety of Calceolaria, which he obtained from a neighbour, "showed much greater vigour than some of his own that were treated in exactly the same manner," and he attributed this solely to his own plants having become "to a certain extent worn out or tired of their quarters." Something of this kind apparently occurs in grafting and budding fruit-trees; for, according to Mr. Abbey, grafts or buds generally take on a distinct variety or even species, or on a stock previously grafted, with greater facility than on stocks raised from seeds of the variety which is to be grafted; and he believes this cannot be altogether explained by the stocks in question being better adapted to the soil and climate of the place. It should, however, be added, that varieties grafted or budded on very distinct kinds, though they may take more readily and grow at first more vigorously than when grafted on closely allied stocks, afterwards often become unhealthy.
I have studied M. Tessier's careful and elaborate experiments,[328] made to disprove the common belief that good is derived from a change of seed; and he certainly shows that the same seed may with care be cultivated on the same farm (it is not stated whether on exactly the same soil) for ten consecutive years without loss. Another excellent observer, Colonel Le Couteur,[329] has come to the same conclusion; but then he expressly adds, if the same seed be used, "that which is grown on land manured from the mixen one year becomes seed for land prepared with lime, and that again becomes seed for land dressed with ashes, then for land dressed with mixed manure, and so on." But this in effect is a systematic exchange of seed, within the limits of the same farm.
On the whole the belief, which has long been held by many skilful cultivators, that good follows from exchanging seed, tubers, &c., seems to be fairly well founded. Considering the small size of most seeds, it seems hardly credible that the advantage thus derived can be due to the seeds obtaining in one soil some chemical element deficient in the other soil. As plants after once germinating naturally become fixed to the same spot, it might have been anticipated that they would show the good effects of a change more plainly than animals, which continually wander about; and this apparently is the {148} case. Life depending on, or consisting in, an incessant play of the most complex forces, it would appear that their action is in some way stimulated by slight changes in the circumstances to which each organism is exposed. All forces throughout nature, as Mr. Herbert Spencer[330] remarks, tend towards an equilibrium, and for the life of each being it is necessary that this tendency should be checked. If these views and the foregoing facts can be trusted, they probably throw light, on the one hand, on the good effects of crossing the breed, for the germ will be thus slightly modified or acted on by new forces; and on the other hand, on the evil effects of close interbreeding prolonged during many generations, during which the germ will be acted on by a male having almost identically the same constitution.
Sterility from changed Conditions of Life.
I will now attempt to show that animals and plants, when removed from their natural conditions, are often rendered in some degree infertile or completely barren; and this occurs even when the conditions have not been greatly changed. This conclusion is not necessarily opposed to that at which we have just arrived, namely, that lesser changes of other kinds are advantageous to organic beings. Our present subject is of some importance, from having an intimate connexion with the causes of variability. Indirectly it perhaps bears on the sterility of species when crossed: for as, on the one hand, slight changes in the conditions of life are favourable to plants and animals, and the crossing of varieties adds to the size, vigour, and fertility of their offspring; so, on the other hand, certain other changes in the conditions of life cause sterility; and as this likewise ensues from crossing much-modified forms or species, we have a parallel and double series of facts, which apparently stand in close relation to each other.
It is notorious that many animals, though perfectly tamed, {149} refuse to breed in captivity. Isidore Geoffroy St. Hilaire[331] consequently has drawn a broad distinction between tamed animals which will not breed under captivity, and truly domesticated animals which breed freely--generally more freely, as shown in the sixteenth chapter, than in a state of nature. It is possible and generally easy to tame most animals; but experience has shown that it is difficult to get them to breed regularly, or even at all. I shall discuss this subject in detail; but will give only those cases which seem most illustrative. My materials are derived from notices scattered through various works, and especially from a Report, drawn up for me by the kindness of the officers of the Zoological Society of London, which has especial value, as it records all the cases, during nine years from 1838-46, in which the animals were seen to couple but produced no offspring, as well as the cases in which they never, as far as known, coupled. This MS. Report I have corrected by the annual Reports subsequently published. Many facts are given on the breeding of the animals in that magnificent work, 'Gleanings from the Menageries of Knowsley Hall,' by Dr. Gray. I made, also, particular inquiries from the experienced keeper of the birds in the old Surrey Zoological Gardens. I should premise that a slight change in the treatment of animals sometimes makes a great difference in their fertility; and it is probable that the results observed in different menageries would differ. Indeed some animals in our Zoological Gardens have become more productive since the year 1846. It is, also, manifest from F. Cuvier's account of the Jardin des Plantes,[332] that the animals formerly bred much less freely there than with us; for instance, in the Duck tribe, which is highly prolific, only one species had at that period produced young.
The most remarkable cases, however, are afforded by animals kept in their native country, which, though perfectly tamed, quite healthy, and allowed some freedom, are absolutely incapable of breeding. Rengger,[333] who in Paraguay particularly attended to this subject, specifies six quadrupeds in this condition; and he mentions two or three others which most rarely {150} breed. Mr. Bates, in his admirable work on the Amazons, strongly insists on similar cases;[334] and he remarks, that the fact of thoroughly tamed native mammals and birds not breeding when kept by the Indians, cannot be wholly accounted for by their negligence or indifference, for the turkey is valued by them, and the fowl has been adopted by the remotest tribes. In almost every part of the world--for instance, in the interior of Africa, and in several of the Polynesian islands--the natives are extremely fond of taming the indigenous quadrupeds and birds; but they rarely or never succeed in getting them to breed.
The most notorious case of an animal not breeding in captivity is that of the elephant. Elephants are kept in large numbers in their native Indian home, live to old age, and are vigorous enough for the severest labour; yet, with one or two exceptions, they have never been known even to couple, though both males and females have their proper periodical seasons. If, however, we proceed a little eastward to Ava, we hear from Mr. Crawfurd[335] that their "breeding in the domestic state, or at least in the half-domestic state in which the female elephants are generally kept, is of every-day occurrence;" and Mr. Crawfurd informs me that he believes that the difference must be attributed solely to the females being allowed to roam the forests with some degree of freedom. The captive rhinoceros, on the other hand, seems from Bishop Heber's account[336] to breed in India far more readily than the elephant. Four wild species of the horse genus have bred in Europe, though here exposed to a great change in their natural habits of life; but the species have generally been crossed one with another. Most of the members of the pig family breed readily in our menageries: even the Red River hog (Potamochoerus penicillatus), from the sweltering plains of West Africa, has bred twice in the Zoological Gardens. Here also the Peccary (Dicotyles torquatus) has bred several times; but another species, the D. labiatus, though rendered so tame as to be half-domesticated, breeds so rarely in its native country of Paraguay, that according to Rengger[337] the fact requires confirmation. Mr. Bates remarks that the tapir, though often kept tame in Amazonia by the Indians, never breeds.
Ruminants generally breed quite freely in England, though brought from widely different climates, as may be seen in the Annual Reports of the Zoological Gardens, and in the Gleanings from Lord Derby's menagerie.
The Carnivora, with the exception of the Plantigrade division, generally breed (though with capricious exceptions) almost as freely as ruminants. Many species of Felidæ have bred in various menageries, although imported from various climates and closely confined. Mr. Bartlett, the present superintendent of the Zoological Gardens,[338] remarks that the lion appears to breed more frequently and to bring forth more young at a birth than any other species of the family. He adds that the tiger has rarely bred; {151} "but there are several well-authenticated instances of the female tiger breeding with the lion." Strange as the fact may appear, many animals under confinement unite with distinct species and produce hybrids quite as freely as, or even more freely than, with their own species. On inquiring from Dr. Falconer and others, it appears that the tiger when confined in India does not breed, though it has been known to couple. The cheetah (Felis jubata) has never been known by Mr. Bartlett to breed in England, but it has bred at Frankfort; nor does it breed in India, where it is kept in large numbers for hunting; but no pains would be taken to make them breed, as only those animals which have hunted for themselves in a state of nature are serviceable and worth training.[339] According to Rengger, two species of wild cats in Paraguay, though thoroughly tamed, have never bred. Although so many of the Felidæ breed readily in the Zoological Gardens, yet conception by no means always follows union: in the nine-year Report, various species are specified which were observed to couple seventy-three times, and no doubt this must have passed many times unnoticed; yet from the seventy-three unions only fifteen births ensued. The Carnivora in the Zoological Gardens were formerly less freely exposed to the air and cold than at present, and this change of treatment, as I was assured by the former superintendent, Mr. Miller, greatly increased their fertility. Mr. Bartlett, and there cannot be a more capable judge, says, "it is remarkable that lions breed more freely in travelling collections than in the Zoological Gardens; probably the constant excitement and irritation produced by moving from place to place, or change of air, may have considerable influence in the matter."
Many members of the Dog family breed readily when confined. The Dhole is one of the most untameable animals in India, yet a pair kept there by Dr. Falconer produced young. Foxes, on the other hand, rarely breed, and I have never heard of such an occurrence with the European fox: the silver fox of North America (Canis argentatus), however, has bred several times in the Zoological Gardens. Even the otter has bred there. Every one knows how readily the semi-domesticated ferret breeds, though shut up in miserably small cages; but other species of Viverra and Paradoxurus absolutely refuse to breed in the Zoological Gardens. The Genetta has bred both here and in the Jardin des Plantes, and produced hybrids. The Herpestes fasciatus has likewise bred; but I was formerly assured that the H. griseus, though many were kept in the Gardens, never bred.
The Plantigrade Carnivora breed under confinement much less freely, without our being able to assign any reason, than other members of the group. In the nine-year Report it is stated that the bears had been seen in the Zoological Gardens to couple freely, but previously to 1848 had most rarely conceived. In the Reports published since this date three species have produced young (hybrids in one case), and, wonderful to relate, the white Polar bear has produced young. The badger (Meles taxus) has bred several times in the Gardens; but I have not heard of this {152} occurring elsewhere in England, and the event must be very rare, for an instance in Germany has been thought worth recording.[340] In Paraguay the native Nasua, though kept in pairs during many years and perfectly tamed, has never been known, according to Rengger, to breed or show any sexual passion; nor, as I hear from Mr. Bates, does this animal, or the Cercoleptes, breed in the region of the Amazons. Two other plantigrade genera, Procyon and Gulo, though often kept tame in Paraguay, never breed there. In the Zoological Gardens species of Nasua and Procyon have been seen to couple; but they did not produce young.
As domesticated rabbits, guinea-pigs, and white mice breed so abundantly when closely confined under various climates, it might have been thought that most other members of the Rodent order would have bred in captivity, but this is not the case. It deserves notice, as showing how the capacity to breed sometimes goes by affinity, that the one native rodent of Paraguay, which there breeds freely and has yielded successive generations, is the Cavia aperea; and this animal is so closely allied to the guinea-pig, that it has been erroneously thought to be the parent-form.[341] In the Zoological Gardens, some rodents have coupled, but have never produced young; some have neither coupled nor bred; but a few have bred, as the porcupine more than once, the Barbary mouse, lemming, chinchilla, and the agouti (Dasyprocta aguti), several times. This latter animal has also produced young in Paraguay, though they were born dead and ill-formed; but in Amazonia, according to Mr. Bates, it never breeds, though often kept tame about the houses. Nor does the paca (Coelogenys paca) breed there. The common hare when confined has, I believe, never bred in Europe;[342] though, according to a recent statement, it has crossed with the rabbit. I have never heard of the dormouse breeding in confinement. But squirrels offer a more curious case: with one exception, no species has ever bred in the Zoological Gardens, yet as many as fourteen individuals of S. palmarum were kept together during several years. The S. cinerea has been seen to couple, but it did not produce young; nor has this species, when rendered extremely tame in its native country, North America, been ever known to breed.[343] At Lord Derby's menagerie squirrels of many kinds were kept in numbers, but Mr. Thompson, the superintendent, told me that none had ever bred there, or elsewhere as far as he knew. I have never heard of the English squirrel breeding in confinement. But the species which has bred more than once in the Zoological Gardens is the one which perhaps might have been least expected, namely, the flying squirrel (Sciuropterus volucella): it has, also, bred several times {153} near Birmingham; but the female never produced more than two young at a birth, whereas in its native American home she bears from three to six young.[344]
Monkeys, in the nine-year Report from the Zoological Gardens, are stated to unite most freely, but during this period, though many individuals were kept, there were only seven births. I have heard of one American monkey alone, the Ouistiti, breeding in Europe.[345] A Macacus, according to Flourens, bred in Paris; and more than one species of this genus has produced young in London, especially the Macacus rhesus, which everywhere shows a special capacity to breed under confinement. Hybrids have been produced both in Paris and London from this same genus. The Arabian baboon, or Cynocephalus hamadryas,[346] and a Cercopithecus have bred in the Zoological Gardens, and the latter species at the Duke of Northumberland's. Several members of the family of Lemurs have produced hybrids in the Zoological Gardens. It is much more remarkable that monkeys very rarely breed when confined in their native country; thus the Cay (Cebus azaræ) is frequently and completely tamed in Paraguay, but Rengger[347] says that it breeds so rarely, that he never saw more than two females which had produced young. A similar observation has been made with respect to the monkeys which are frequently tamed by the aborigines in Brazil.[348] In the region of the Amazons, these animals are so often kept in a tame state, that Mr. Bates in walking through the streets of Parà counted thirteen species; but, as he asserts, they have never been known to breed in captivity.[349]
Birds.
Birds offer in some respects better evidence than quadrupeds, from their breeding more rapidly and being kept in greater numbers. We have seen that carnivorous animals are more fertile under confinement than most other mammals. The reverse holds good with carnivorous birds. It is said[350] that as many as eighteen species have been used in Europe for hawking, and several others in Persia and India;[351] they have been kept in their native country in the finest condition, and have been flown during six, eight, or nine years;[352] yet there is no record of their having ever produced young. As these birds were formerly caught whilst young, at great expense, being imported from Iceland, Norway, and Sweden, there can {154} be little doubt that, if possible, they would have been propagated. In the Jardin des Plantes, no bird of prey has been known to couple.[353] No hawk, vulture, or owl has ever produced fertile eggs in the Zoological Gardens, or in the old Surrey Gardens, with the exception, in the former place on one occasion, of a condor and a kite (Milvus niger). Yet several species, namely, the Aquila fusca, Haliætus leucocephalus, Falco tinnunculus, F. subbuteo, and Buteo vulgaris, have been seen to couple in the Zoological Gardens. Mr. Morris[354] mentions as a unique fact that a kestrel (Falco tinnunculus) bred in an aviary. The one kind of owl which has been known to couple in the Zoological Gardens was the Eagle Owl (Bubo maximus); and this species shows a special inclination to breed in captivity; for a pair at Arundel Castle, kept more nearly in a state of nature "than ever fell to the lot of an animal deprived of its liberty,"[355] actually reared their young. Mr. Gurney has given another instance of this same owl breeding in confinement; and he records the case of a second species of owl, the Strix passerina, breeding in captivity.[356]
Of the smaller graminivorous birds, many kinds have been kept tame in their native countries, and have lived long; yet, as the highest authority on cage-birds[357] remarks, their propagation is "uncommonly difficult." The canary-bird shows that there is no inherent difficulty in these birds breeding freely in confinement; and Audubon says[358] that the Fringilla (Spiza) ciris of North America breeds as perfectly as the canary. The difficulty with the many finches which have been kept in confinement is all the more remarkable as more than a dozen species could be named which have yielded hybrids with the canary; but hardly any of these, with the exception of the siskin (Fringilla spinus), have reproduced their own kind. Even the bullfinch (Loxia pyrrhula) has bred as frequently with the canary, though belonging to a distinct genus, as with its own species.[359] With respect to the skylark (Alauda arvensis), I have heard of birds living for seven years in an aviary, which never produced young; and a great London bird-fancier assured me that he had never known an instance of their breeding; nevertheless one case has been recorded.[360] In the nine-year Report from the Zoological Society, twenty-four incessorial species are enumerated which had not bred, and of these only four were known to have coupled.
Parrots are singularly long-lived birds; and Humboldt mentions the curious fact of a parrot in South America, which spoke the language of {155} an extinct Indian tribe, so that this bird preserved the sole relic of a lost language. Even in this country there is reason to believe[361] that parrots have lived to the age of nearly one hundred years; yet, though many have been kept in Europe, they breed so rarely that the event has been thought worth recording in the gravest publications.[362] According to Bechstein[363] the African Psittacus erithacus breeds oftener than any other species: the P. macoa occasionally lays fertile eggs, but rarely succeeds in hatching them; this bird, however, has the instinct of incubation sometimes so strongly developed, that it will hatch the eggs of fowls or pigeons. In the Zoological Gardens and in the old Surrey Gardens some few species have coupled, but, with the exception of three species of parrakeets, none have bred. It is a much more remarkable fact that in Guiana parrots of two kinds, as I am informed by Sir E. Schomburgk, are often taken from the nests by the Indians and reared in large numbers; they are so tame that they fly freely about the houses, and come when called to be fed, like pigeons; yet he has never heard of a single instance of their breeding.[364] In Jamaica, a resident naturalist, Mr. R. Hill,[365] says, "no birds more readily submit to human dependence than the parrot-tribe, but no instance of a parrot breeding in this tame life has been known yet." Mr. Hill specifies a number of other native birds kept tame in the West Indies, which never breed in this state.
The great pigeon family offers a striking contrast with parrots: in the nine-year Report thirteen species are recorded as having bred, and, what is more noticeable, only two were seen to couple without any result. Since the above date every annual Report gives many cases of various pigeons breeding. The two magnificent crowned pigeons (Goura coronata and Victoriæ) produced hybrids; nevertheless, of the former species more than a dozen birds were kept, as I am informed by Mr. Crawfurd, in a park at Penang, under a perfectly well-adapted climate, but never once bred. The Columba migratoria in its native country, North America, invariably lays two eggs, but in Lord Derby's menagerie never more than one. The same fact has been observed with the C. leucocephala.[366]
Gallinaceous birds of many genera likewise show an eminent capacity for breeding under captivity. This is particularly the case with pheasants; yet our English species seldom lays more than ten eggs in confinement; whilst from eighteen to twenty is the usual number in the wild state.[367] With the Gallinaceæ, as with all other orders, there are marked and {156} inexplicable exceptions in regard to the fertility of certain species and genera under confinement. Although many trials have been made with the common partridge, it has rarely bred, even when reared in large aviaries; and the hen will never hatch her own eggs.[368] The American tribe of Guans or Cracidæ are tamed with remarkable ease, but are very shy breeders in this country;[369] but with care various species were formerly made to breed rather freely in Holland.[370] Birds of this tribe are often kept in a perfectly tamed condition in their native country by the Indians, but they never breed.[371] It might have been expected that grouse from their habits of life would not have bred in captivity, more especially as they are said soon to languish and die.[372] But many cases are recorded of their breeding: the capercailzie (Tetrao urogallus) has bred in the Zoological Gardens; it breeds without much difficulty when confined in Norway, and in Russia five successive generations have been reared: Tetrao tetrix has likewise bred in Norway; T. Scoticus in Ireland; T. umbellus at Lord Derby's; and T. cupido in North America.
It is scarcely possible to imagine a greater change in habits than that which the members of the ostrich family must suffer, when cooped up in small enclosures under a temperate climate, after freely roaming over desert and tropical plains or entangled forests. Yet almost all the kinds, even the mooruk (Casuarius Bennettii) from New Ireland, has frequently produced young in the various European menageries. The African ostrich, though perfectly healthy and living long in the South of France, never lays more than from twelve to fifteen eggs, though in its native country it lays from twenty-five to thirty.[373] Here we have another instance of fertility impaired, but not lost, under confinement, as with the flying squirrel, the hen-pheasant, and two species of American pigeons.
Most Waders can be tamed, as the Rev. E. S. Dixon informs me, with remarkable facility; but several of them are short-lived under confinement, so that their sterility in this state is not surprising. The cranes breed more readily than other genera: Grus montigresia has bred several times in Paris and in the Zoological Gardens, as has G. cinerea at the latter place, and G. antigone at Calcutta. Of other members of this great order, Tetrapteryx paradisea has bred at Knowsley, a Porphyrio in Sicily, and the Gallinula chloropus in the Zoological Gardens. On the other hand, several {157} birds belonging to this order will not breed in their native country, Jamaica; and the Psophia, though often kept by the Indians of Guiana about their houses, "is seldom or never known to breed."[374]
No birds breed with such complete facility under confinement as the members of the great Duck family; yet, considering their aquatic and wandering habits, and the nature of their food, this could not have been anticipated. Even some time ago above two dozen species had bred in the Zoological Gardens; and M. Selys-Longchamps has recorded the production of hybrids from forty-four different members of the family; and to these Professor Newton has added a few more cases.[375] "There is not," says Mr. Dixon,[376] "in the wide world, a goose which is not in the strict sense of the word domesticable;" that is, capable of breeding under confinement; but this statement is probably too bold. The capacity to breed sometimes varies in individuals of the same species; thus Audubon[377] kept for more than eight years some wild geese (Anser Canadensis), but they would not mate; whilst other individuals of the same species produced young during the second year. I know of but one instance in the whole family of a species which absolutely refuses to breed in captivity, namely, the Dendrocygna viduata, although, according to Sir R. Schomburgk,[378] it is easily tamed, and is frequently kept by the Indians of Guiana. Lastly, with respect to Gulls, though many have been kept in the Zoological Gardens and in the old Surrey Gardens, no instance was known before the year 1848 of their coupling or breeding; but since that period the herring gull (Larus argentatus) has bred many times in the Zoological Gardens and at Knowsley.
There is reason to believe that insects are affected by confinement like the higher animals. It is well known that the Sphingidæ rarely breed when thus treated. An entomologist[379] in Paris kept twenty-five specimens of Saturnia pyri, but did not succeed in getting a single fertile egg. A number of females of Orthosia munda and of Mamestra suasa reared in confinement were unattractive to the males.[380] Mr. Newport kept nearly a hundred individuals of two species of Vanessa, but not one paired; this, however, might have been due to their habit of coupling on the wing.[381] Mr. Atkinson could never succeed in India in making the Tarroo silk-moth breed in confinement.[382] It appears that a number of moths, especially the Sphingidæ, when hatched in the autumn out of their proper season, {158} are completely barren; but this latter case is still involved in some obscurity.[383]
Independently of the fact of many animals under confinement not coupling, or, if they couple, not producing young, there is evidence of another kind, that their sexual functions are thus disturbed. For many cases have been recorded of the loss by male birds when confined of their characteristic plumage. Thus the common linnet (Linota cannabina) when caged does not acquire the fine crimson colour on its breast, and one of the buntings (Emberiza passerina) loses the black on its head. A Pyrrhula and an Oriolus have been observed to assume the quiet plumage of the hen-bird; and the Falco albidus returned to the dress of an earlier age.[384] Mr. Thomson, the superintendent of the Knowsley menagerie, informed me that he had often observed analogous facts. The horns of a male deer (Cervus Canadensis) during the voyage from America were badly developed; but subsequently in Paris perfect horns were produced.
When conception takes place under confinement, the young are often born dead, or die soon, or are ill-formed. This frequently occurs in the Zoological Gardens, and, according to Rengger, with native animals confined in Paraguay. The mother's milk often fails. We may also attribute to the disturbance of the sexual functions the frequent occurrence of that monstrous instinct which leads the mother to devour her own offspring,--a mysterious case of perversion, as it at first appears.
Sufficient evidence has now been advanced to prove that animals when first confined are eminently liable to suffer in their reproductive systems. We feel at first naturally inclined to attribute the result to loss of health, or at least to loss of vigour; but this view can hardly be admitted when we reflect how healthy, long-lived, and vigorous many animals are under {159} captivity, such as parrots, and hawks when used for hawking, chetahs when used for hunting, and elephants. The reproductive organs themselves are not diseased; and the diseases, from which animals in menageries usually perish, are not those which in any way affect their fertility. No domestic animal is more subject too disease than the sheep, yet it is remarkably prolific. The failure of animals to breed under confinement has been sometimes attributed exclusively to a failure in their sexual instincts: this may occasionally come into play, but there is no obvious reason why this instinct should be especially liable to be affected with perfectly tamed animals, except indeed indirectly through the reproductive system itself being disturbed. Moreover, numerous cases have been given of various animals which couple freely under confinement, but never conceive; or, if they conceive and produce young, these are fewer in number than is natural to the species. In the vegetable kingdom instinct of course can play no part; and we shall presently see that plants when removed from their natural conditions are affected in nearly the same manner as animals. Change of climate cannot be the cause of the loss of fertility, for, whilst many animals imported into Europe from extremely different climates breed freely, many others when confined in their native land are completely sterile. Change of food cannot be the chief cause; for ostriches, ducks, and many other animals, which must have undergone a great change in this respect, breed freely. Carnivorous birds when confined are extremely sterile; whilst most carnivorous mammals, except plantigrades, are moderately fertile. Nor can the amount of food be the cause; for a sufficient supply will certainly be given to valuable animals; and there is no reason to suppose that much more food would be given to them, than to our choice domestic productions which retain their full fertility. Lastly, we may infer from the case of the elephant, chetah, various hawks, and of many animals which are allowed to lead an almost free life in their native land, that want of exercise is not the sole cause.
It would appear that any change in the habits of life, whatever these habits may be, if great enough, tends to affect in an inexplicable manner the powers of reproduction. The result {160} depends more on the constitution of the species than on the nature of the change; for certain whole groups are affected more than others; but exceptions always occur, for some species in the most fertile groups refuse to breed, and some in the most sterile groups breed freely. Those animals which usually breed freely under confinement, rarely breed, as I was assured, in the Zoological Gardens, within a year or two after their first importation. When an animal which is generally sterile under confinement happens to breed, the young apparently do not inherit this power; for had this been the case, various quadrupeds and birds, which are valuable for exhibition, would have become common. Dr. Broca even affirms[385] that many animals in the Jardin des Plantes, after having produced young for three or four successive generations, become sterile; but this may be the result of too close interbreeding. It is a remarkable circumstance that many mammals and birds have produced hybrids under confinement quite as readily as, or even more readily than, they have procreated their own kind. Of this fact many instances have been given;[386] and we are thus reminded of those plants which when cultivated refuse to be fertilised by their own pollen, but can easily be fertilised by that of a distinct species. Finally, we must conclude, limited as the conclusion is, that changed conditions of life have an especial power of acting injuriously on the reproductive system. The whole case is quite peculiar, for these organs, though not diseased, are thus rendered incapable of performing their proper functions, or perform them imperfectly.
Sterility of Domesticated Animals from changed conditions.--With respect to domesticated animals, as their domestication mainly depends on the accident of their breeding freely under captivity, we ought not to expect that their reproductive system would be affected by any moderate degree of change. Those orders of quadrupeds and birds, of which the wild species breed most readily in our menageries, have afforded us the greatest number of domesticated productions. Savages in most parts of the world are fond of taming animals;[387] and if any of these regularly produced {161} young, and were at the same time useful, they would be at once domesticated. If, when their masters migrated into other countries, they were in addition found capable of withstanding various climates, they would be still more valuable; and it appears that the animals which breed readily in captivity can generally withstand different climates. Some few domesticated animals, such as the reindeer and camel, offer an exception to this rule. Many of our domesticated animals can bear with undiminished fertility the most unnatural conditions; for instance, rabbits, guinea-pigs, and ferrets breed in miserably confined hutches. Few European dogs of any kind withstand without degeneration the climate of India; but as long as they survive, they retain, as I hear from Mr. Falconer, their fertility; so it is, according to Dr. Daniell, with English dogs taken to Sierra Leone. The fowl, a native of the hot jungles of India, becomes more fertile than its parent-stock in every quarter of the world, until we advance as far north as Greenland and Northern Siberia, where this bird will not breed. Both fowls and pigeons, which I received during the autumn direct from Sierra Leone, were at once ready to couple.[388] I have, also, seen pigeons breeding as freely as the common kinds within a year after their importation from the Upper Nile. The guinea-fowl, an aboriginal of the hot and dry deserts of Africa, whilst living under our damp and cool climate, produces a large supply of eggs.
Nevertheless, our domesticated animals under new conditions occasionally show signs of lessened fertility. Roulin asserts that in the hot valleys of the equatorial Cordillera sheep are not fully fecund;[389] and according to Lord Somerville,[390] the merino-sheep which he imported from Spain were not at first perfectly fertile. It is said[391] that mares brought up on dry food in the stable, and turned out to grass, do not at first breed. The peahen, as we have seen, is said not to lay so many eggs in England as in India. It was long before the canary-bird was fully fertile, and even now first-rate breeding birds are not common.[392] In the hot and dry province of Delhi, the eggs of the turkey, as I hear from Dr. Falconer, though placed under a hen, are extremely liable to fail. According to Roulin, geese taken within a recent period to the lofty plateau of Bogota, at first laid seldom, and then only a few eggs; of these scarcely a fourth were hatched, and half the young birds died: in the second generation they were more fertile; and when Roulin wrote they were becoming as {162} fertile as our geese in Europe. In the Philippine Archipelago the goose, it is asserted, will not breed or even lay eggs.[393] A more curious case is that of the fowl, which, according to Roulin, when first introduced would not breed at Cusco in Bolivia, but subsequently became quite fertile; and the English Game fowl, lately introduced, had not as yet arrived a its full fertility, for to raise two or three chickens from a nest of eggs was thought fortunate. In Europe close confinement has a marked effect on the fertility of the fowl: it has been found in France that with fowls allowed considerable freedom only twenty per cent. of the eggs failed; when allowed less freedom forty per cent. failed; and in close confinement sixty out of the hundred were not hatched.[394] So we see that unnatural and changed conditions of life produce some effect on the fertility of our most thoroughly domesticated animals, in the same manner, though in a far less degree, as with captive wild animals.
It is by no means rare to find certain males and females which will not breed together, though both are known to be perfectly fertile with other males and females. We have no reason to suppose that this is caused by these animals having been subjected to any change in their habits of life; therefore such cases are hardly related to our present subject. The cause apparently lies in an innate sexual incompatibility of the pair which are matched. Several instances have been communicated to me by Mr. W. C. Spooner (well known for his essay on Cross-breeding), by Mr. Eyton of Eyton, by Mr. Wicksted and othe breeders, and especially by Mr. Waring of Chelsfield, in relation to horses, cattle, pigs, foxhounds, other dogs, and pigeons.[395] In these cases, females, which either previously or subsequently were proved to be fertile, failed to breed with certain males, with whom it was particularly desired to match them. A change in the constitution of the female may sometimes have occurred before she was put to the second male; but in other cases this explanation is hardly tenable, for a female, known not to be barren, has been unsuccessfully paired seven or eight times with the same male likewise known to be perfectly fertile. With cart-mares, which sometimes will not breed with stallions of pure blood, but subsequently have bred with cart-stallions, Mr. Spooner is inclined to attribute the failure to the lesser sexual power of the race-horse. But I have heard from the greatest breeder of race-horses at the present day, through Mr. Waring, that "it frequently occurs with a mare to be put several times during one or two seasons to a particular stallion of acknowledged power, and yet prove barren; the mare afterwards breeding at once with some other horse." These facts are worth recording, as they show, like so many previous facts, on what slight constitutional differences the fertility of an animal often depends.
{163}
Sterility of Plants from changed Conditions of Life, and from other causes.
In the vegetable kingdom cases of sterility frequently occur, analogous with those previously given in the animal kingdom. But the subject is obscured by several circumstances, presently to be discussed, namely, the contabescence of the anthers, as Gärtner has named a certain affection--monstrosities--doubleness of the flower--much-enlarged fruit--and long-continued or excessive propagation by buds.
It is notorious that many plants in our gardens and hot-houses, though preserved in the most perfect health, rarely or never produce seed. I do not allude to plants which run to leaves, from being kept too damp, or too warm, or too much manured; for these do not produce the reproductive individual or flower, and the case may be wholly different. Nor do I allude to fruit not ripening from want of heat, or rotting from too much moisture. But many exotic plants, with their ovules and pollen appearing perfectly sound, will not set any seed. The sterility in many cases, as I know from my own observation, is simply due to the absence of the proper insects for carrying the pollen to the stigma. But after excluding the several cases just specified, there are many plants in which the reproductive system has been seriously affected by the altered conditions of life to which they have been subjected.
It would be tedious to enter on many details. Linnæus long ago observed[396] that Alpine plants, although naturally laded with seed, produce either few or none when cultivated in gardens. But exceptions often occur: the Draba sylvestris, one of our most thoroughly Alpine plants, multiplies itself by seed in Mr. H. C. Watson's garden, near London; and Kerner, who has particularly attended to the cultivation of Alpine plants, found that various kinds, when cultivated, spontaneously sowed themselves.[397] Many plants which naturally grow in peat-earth are entirely sterile in our gardens. I have noticed the same fact with several liliaceous plants, which nevertheless grew vigorously.
Too much manure renders some kinds utterly sterile, as I have myself observed. The tendency to sterility from this cause runs in families; thus, according to Gärtner,[398] it is hardly possible to give too much manure to most Gramineæ, Cruciferæ, and Leguminosæ, whilst succulent and bulbous-rooted plants are easily affected. Extreme poverty of soil is less {164} apt to induce sterility; but dwarfed plants of Trifolium minus and repens, growing on a lawn often mown and never manured, did not produce any seed. The temperature of the soil, and the season at which plants are watered, often have a marked effect on their fertility, as was observed by Kölreuter in the case of Mirabilis.[399] Mr. Scott in the Botanic Gardens of Edinburgh observed that Oncidium divaricatum would not set seed when grown in a basket in which it throve, but was capable of fertilisation in a pot where it was a little damper. Pelargonium fulgidum, for many years after its introduction, seeded freely; it then became sterile; now it is fertile[400] if kept in a dry stove during the winter. Other varieties of pelargonium are sterile and others fertile without our being able to assign any cause. Very slight changes in the position of a plant, whether planted on a bank or at its base, sometimes make all the difference in its producing seed. Temperature apparently has a much more powerful influence on the fertility of plants than on that of animals. Nevertheless it is wonderful what changes some few plants will withstand with undiminished fertility: thus the Zephyranthes candida, a native of the moderately warm banks of the Plata, sows itself in the hot dry country near Lima, and in Yorkshire resists the severest frosts, and I have seen seeds gathered from pods which had been covered with snow during three weeks.[401] Berberis Wallichii, from the hot Khasia range in India, is uninjured by our sharpest frosts, and ripens its fruit under our cool summers. Nevertheless I presume we must attribute to change of climate the sterility of many foreign plants; thus the Persian and Chinese lilacs (Syringa Persica and Chinensis), though perfectly hardly, never here produce a seed; the common lilac (S. vulgaris) seeds with us moderately well, but in parts of Germany the capsules never contain seed.[402]
Some of the cases, given in the last chapter, of self-impotent plants, which are fertile both on the male and female side when united with distinct individuals or species, might have been here introduced; for as this peculiar form of sterility generally occurs with exotic plants or with endemic plants cultivated in pots, and as it disappeared in the Passiflora alata when grafted, we may conclude that in these cases it is the result of the treatment to which the plants or their parents have been exposed.
The liability of plants to be affected in their fertility by slightly changed conditions is the more remarkable, as the pollen when once in process of formation is not easily injured; a plant may be transplanted, or a branch with flower-buds be cut off and placed in water, and the pollen will be matured. Pollen, also, when once mature, may be kept for weeks or even months.[403] The female organs are more sensitive, for Gärtner[404] found that dicotyledonous plants, when carefully removed so that they did not in the least flag, could seldom be fertilised; this occurred even with potted {165} plants if the roots had grown out of the hole at the bottom. In some few cases, however, as with Digitalis, transplantation did not prevent fertilisation; and according to the testimony of Mawz, Brassica rapa, when pulled up by its roots and placed in water, ripened its seed. Flower-stems of several monocotyledonous plants when cut off and placed in water likewise produce seed. But in these cases I presume that the flowers had been already fertilised, for Herbert[405] found with the Crocus that the plants might be removed or mutilated after the act of fertilisation, and would still perfect their seeds; but that, if transplanted before being fertilised, the application of pollen was powerless.
Plants which have been long cultivated can generally endure with undiminished fertility various and great changes; but not in most cases so great a change of climate as domesticated animals. It is remarkable that many plants under these circumstances are so much affected that the proportions and the nature of their chemical ingredients are modified, yet their fertility is unimpaired. Thus, as Dr. Falconer informs me, there is a great difference in the character of the fibre in hemp, in the quantity of oil in the seed of the Linum, in the proportion of narcotin to morphine in the poppy, in gluten to starch in wheat, when these plants are cultivated on the plains and on the mountains of India; nevertheless, they all remain fully fertile.
Contabescence.--Gärtner has designated by this term a peculiar condition of the anthers in certain plants, in which they are shrivelled, or become brown and tough, and contain no good pollen. When in this state they exactly resemble the anthers of the most sterile hybrids. Gärtner,[406] in his discussion on this subject, has shown that plants of many orders are occasionally thus affected; but the Caryophyllaceæ and Liliaceæ suffer most, and to these orders, I think, the Ericaceæ may be added. Contabescence varies in degree, but on the same plant all the flowers are generally affected to nearly the same extent. The anthers are affected at a very early period in the flower-bud, and remain in the same state (with one recorded exception) during the life of the plant. The affection cannot be cured by any change of treatment, and is propagated by layers, cuttings, &c., and perhaps even by seed. In contabescent plants the female organs are seldom affected, or merely become precocious in their development. The cause of this affection is doubtful, and is different in different cases. Until I read Gärtner's discussion I attributed it, as apparently did Herbert, to the unnatural treatment of the plants; but its permanence under changed conditions, and the female organs not being affected, seem incompatible with this view. The fact of several endemic plants becoming contabescent in our gardens seems, at first sight, equally incompatible with this view; but Kölreuter believes that this is the result of their transplantation. The contabescent plants of Dianthus and Verbascum, found wild by Wiegmann, grew on a dry and sterile bank. The fact that exotic {166} plants are eminently liable to this affection also seems to show that it is in some manner caused by their unnatural treatment. In some instances, as with Silene, Gärtner's view seems the most probable, namely, that it is caused by an inherent tendency in the species to become dioecious. I can add another cause, namely, the illegitimate unions of reciprocally dimorphic or trimorphic plants, for I have observed seedlings of three species of Primula and of Lythrum salicaria, which had been raised from plants illegitimately fertilised by their own-form pollen, with some or all their anthers in a contabescent state. There is perhaps an additional cause, namely, self-fertilisation; for many plants of Dianthus and Lobelia, which had been raised from self-fertilised seeds, had their anthers in this state; but these instances are not conclusive, as both genera are liable from other causes to this affection.
Cases of an opposite nature likewise occur, namely, plants with the female organs struck with sterility, whilst the male organs remain perfect. Dianthus Japonicus, a Passiflora, and Nicotiana, have been described by Gärtner[407] as being in this unusual condition.
Monstrosities as a cause of Sterility.--Great deviations of structure, even when the reproductive organs themselves are not seriously affected, sometimes cause plants to become sterile. But in other cases plants may become monstrous to an extreme degree and yet retain their full fertility. Gallesio, who certainly had great experience,[408] often attributes sterility to this cause; but it may be suspected that in some of his cases sterility was the cause, and not the result, of the monstrous growths. The curious St. Valery apple, although it bears fruit, rarely produces seed. The wonderfully anomalous flowers of Begonia frigida, formerly described, though they appear fit for fructification, are sterile.[409] Species of Primulæ, in which the calyx is brightly coloured, are said[410] to be often sterile, though I have known them to be fertile. On the other hand, Verlot gives several cases of proliferous flowers which can be propagated by seed. This was the case with a poppy, which had become monopetalous by the union of its petals.[411] Another extraordinary poppy, with the stamens replaced by numerous small supplementary capsules, likewise reproduces itself by seed. This has also occurred with a plant of Saxifraga geum, in which a series of adventitious carpels, bearing ovules on their margins, had been developed between the stamens and the normal carpels.[412] Lastly, with respect to peloric flowers, which depart wonderfully from the natural structure,--those of Linaria vulgaris seem generally to be more or less sterile, whilst those before described of Antirrhinum majus, when artificially fertilised with their own pollen, are perfectly {167} fertile, though sterile when left to themselves, for bees are unable to crawl into the narrow tubular flower. The peloric flowers of Corydalis solida, according to Godron,[413] are barren; whilst those of Gloxinia are well known to yield plenty of seed. In our greenhouse Pelargoniums, the central flower of the truss is often peloric, and Mr. Masters informs me that he tried in vain during several years to get seed from these flowers. I likewise made many vain attempts, but sometimes succeeded in fertilising them with pollen from a normal flower of another variety; and conversely I several times fertilised ordinary flowers with peloric pollen. Only once I succeeded in raising a plant from a peloric flower fertilised by pollen from a peloric flower borne by another variety; but the plant, it may be added, presented nothing particular in its structure. Hence we may conclude that no general rule can be laid down; but any great deviation from the normal structure, even when the reproductive organs themselves are not seriously affected, certainly often leads to sexual impotence.
Double Flowers.--When the stamens are converted into petals, the plant becomes on the male side sterile; when both stamens and pistils are thus changed, the plant becomes completely barren. Symmetrical flowers having numerous stamens and petals are the most liable to become double, as perhaps follows from all multiple organs being the most subject to variability. But flowers furnished with only a few stamens, and others which are asymmetrical in structure, sometimes become double, as we see with the double gorse or Ulex, Petunia, and Antirrhinum. The Compositæ bear what are called double flowers by the abnormal development of the corolla of their central florets. Doubleness is sometimes connected with prolification,[414] or the continued growth of the axis of the flower. Doubleness is strongly inherited. No one has produced, as Lindley remarks,[415] double flowers by promoting the perfect health of the plant. On the contrary, unnatural conditions of life favour their production. There is some reason to believe that seeds kept during many years, and seeds believed to be imperfectly fertilised, yield double flowers more freely than fresh and perfectly fertilised seed.[416] Long-continued cultivation in rich soil seems to be the commonest exciting cause. A double narcissus and a double Anthemis nobilis, transplanted into very poor soil, have been observed to become single;[417] and I have seen a completely double white primrose rendered permanently single by being divided and transplanted whilst in full flower. It has been observed by Professor Morren that doubleness of the flowers and variegation of the leaves are antagonistic states; but so many exceptions to the rule have lately been recorded,[418] that, though general, it cannot be looked at as invariable. {168} Variegation seems generally to result from a feeble or atrophied condition of the plant, and a large proportion of the seedlings raised from parents both of which are variegated usually perish at an early age; hence we may perhaps infer that doubleness, which is the antagonistic state, commonly arises from a plethoric condition. On the other hand, extremely poor soil sometimes, though rarely, appears to cause doubleness: I formerly described[419] some completely double, bud-like, flowers produced in large numbers by stunted wild plants of Gentiana amarella growing on a poor chalky bank. I have also noticed a distinct tendency to doubleness in the flowers of a Ranunculus, Horse-chesnut, and Bladder-nut (Ranunculus repens, Æsculus pavia, and Staphylea), growing under very unfavourable conditions. Professor Lehman[420] found several wild plants growing near a hot spring with double flowers. With respect to the cause of doubleness, which arises, as we see, under widely different circumstances, I shall presently attempt to show that the most probable view is that unnatural conditions first give a tendency to sterility, and that then, on the principle of compensation, as the reproductive organs do not perform their proper functions, they either become developed into petals, or additional petals are formed. This view has lately been supported by Mr. Laxton,[421] who advances the case of some common peas, which, after long-continued heavy rain, flowered a second time, and produced double flowers.
Seedless Fruit.--Many of our most valuable fruits, although consisting in a homological sense of widely different organs, are either quite sterile, or produce extremely few seeds. This is notoriously the case with our best pears, grapes, and figs, with the pine-apple, banana, bread-fruit, pomegranate, azarole, date-palms, and some members of the orange-tribe. Poorer varieties of these same fruits either habitually or occasionally yield seed.[422] Most horticulturists look at the great size and anomalous development of the fruit as the cause, and sterility as the result; but the opposite view, as we shall presently see, is more probable.
Sterility from the excessive development of the Organs of Growth or Vegetation.--Plants which from any cause grow too luxuriantly, and produce leaves, stems, runners, suckers, tubers, bulbs, &c., in excess, sometimes do not flower, or if they flower do not yield seed. To make European vegetables under the hot climate of India yield seed, it is necessary to check their growth; and, when one-third grown, they are taken up, and their stems and {169} tap-roots are cut or mutilated.[423] So it is with hybrids; for instance, Prof. Lecoq[424] had three plants of Mirabilis, which, though they grew luxuriantly and flowered, were quite sterile; but after beating one with a stick until a few branches alone were left, these at once yielded good seed. The sugar-cane, which grows vigorously and produces a large supply of succulent stems, never, according to various observers, bears seed in the West Indies, Malaga, India, Cochin China, or the Malay Archipelago.[425] Plants which produce a large number of tubers are apt to be sterile, as occurs, to a certain extent, with the common potato; and Mr. Fortune informs me that the sweet potato (Convolvulus batatas) in China never, as far as he has seen, yields seed. Dr. Royle remarks[426] that in India the Agave vivipara, when grown in rich soil, invariably produces bulbs, but no seeds; whilst a poor soil and dry climate leads to an opposite result. In China, according to Mr. Fortune, an extraordinary number of little bulbs are developed in the axils of the leaves of the yam, and this plant does not bear seed. Whether in these cases, as in those of double flowers and seedless fruit, sexual sterility from changed conditions of life is the primary cause which leads to the excessive development of the organs of vegetation, is doubtful; though some evidence might be advanced in favour of this view. It is perhaps a more probable view that plants which propagate themselves largely by one method, namely by buds, have not sufficient vital power or organised matter for the other method of sexual generation.
Several distinguished botanists and good practical judges believe that long-continued propagation by cuttings, runners, tubers, bulbs, &c., independently of any excessive development of these parts, is the cause of many plants failing to produce flowers and of others failing to produce fertile flowers,--it is as if they had lost the habit of sexual generation.[427] That many plants when thus propagated are sterile there can be no doubt, but whether the long continuance of this form of propagation is the actual cause of their sterility, I will not venture, from the want of sufficient evidence, to express an opinion.
That plants may be propagated for long periods by buds, without the aid of sexual generation, we may safely infer from this being the case with many plants which must have long survived in a state of nature. As I have had occasion before to allude to this subject, I will here give such cases as I have collected. Many alpine plants ascend mountains beyond the height at which they can produce seed.[428] Certain species of {170} Poa and Festuca, when growing on mountain-pastures, propagate themselves, as I hear from Mr. Bentham, almost exclusively by bulblets. Kalm gives a more curious instance[429] of several American trees, which grow so plentifully in marshes or in thick woods, that they are certainly well adapted for these stations, yet scarcely ever produce seeds; but when accidentally growing on the outside of the marsh or wood, are loaded with seed. The common ivy is found in Northern Sweden and Russia, but flowers and fruits only in the southern provinces. The Acorus calamus extends over a large portion of the globe, but so rarely perfects its fruit that this has been seen but by few botanists.[430] The Hypericum calycinum, which propagates itself so freely in our shrubberies by rhizomas and is naturalised in Ireland, blossoms profusely, but sets no seed; nor did it set any when fertilised in my garden by pollen from plants growing at a distance. The Lysimachia nummularia, which is furnished with long runners, so seldom produces seed-capsules, that Prof. Decaisne,[431] who has especially attended to this plant, has never seen it in fruit. The Carex rigida often fails to perfect its seed in Scotland, Lapland, Greenland, Germany, and New Hampshire in the United States.[432] The periwinkle (Vinca minor), which spreads largely by runners, is said scarcely ever to produce fruit in England;[433] but this plant requires insect-aid for its fertilisation, and the proper insects may be absent or rare. The Jussiæa grandiflora has become naturalised in Southern France, and has spread by its rhizomas so extensively as to impede the navigation of the waters, but never produces fertile seed.[434] The horse-radish (Cochlearia armoracia) spreads pertinaciously and is naturalised in various parts of Europe; though it bears flowers, these rarely produce capsules: Professor Caspary also informs me that he has watched this plant since 1851, but has never seen its fruit; nor is this surprising, as he finds scarcely a grain of good pollen. The common little Ranunculus ficaria rarely, and some say never, bears seed in England, France, or Switzerland; but in 1863 I observed seeds on several plants growing near my house. According to M. Chatin, there are two forms of this Ranunculus; and it is the bulbiferous form which does not yield seed from producing no pollen.[435] Other cases {171} analogous with the foregoing could be given; for instance, some kinds of mosses and lichens have never been seen to fructify in France.
Some of these endemic and naturalised plants are probably rendered sterile from excessive multiplication by buds, and their consequent incapacity to produce and nourish seed. But the sterility of others more probably depends on the peculiar conditions under which they live, as in the case of the ivy in the northern parts of Europe, and of the trees in the swamps of the United States; yet these plants must be in some respects eminently well adapted for the stations which they occupy, for they hold their places against a host of competitors.
Finally, when we reflect on the sterility which accompanies the doubling of flowers,--the excessive development of fruit,--and a great increase in the organs of vegetation, we must bear in mind that the whole effect has seldom been caused at once. An incipient tendency is observed, and continued selection completes the work, as is known to be the case with our double flowers and best fruits. The view which seems the most probable, and which connects together all the foregoing facts and brings them within our present subject, is, that changed and unnatural conditions of life first give a tendency to sterility; and in consequence of this, the organs of reproduction being no longer able fully to perform their proper functions, a supply of organised matter, not required for the development of the seed, flows either into these same organs and renders them foliaceous, or into the fruit, stems, tubers, &c., increasing their size and succulency. But I am far from wishing to deny that there exists, independently of any incipient sterility, an antagonism between the two forms of reproduction, namely, by seed and by buds, when either is carried to an extreme degree. That incipient sterility plays an important part in the doubling of flowers, and in the other cases just specified, I infer chiefly from the following facts. When fertility is lost from a wholly different cause, namely, from hybridism, there is a strong tendency, as Gärtner[436] affirms, for flowers to become double, and this tendency is inherited. Moreover it is notorious that with hybrids the male organs become sterile before the female organs, and with double flowers the stamens first become {172} foliaceous. This latter fact is well shown by the male flowers of dioecious plants, which, according to Gallesio,[437] first become double. Again, Gärtner[438] often insists that the flowers of even utterly sterile hybrids, which do not produce any seed, generally yield perfect capsules or fruit,--a fact which has likewise been repeatedly observed by Naudin with the Cucurbitaceæ; so that the production of fruit by plants rendered sterile through any other and distinct cause is intelligible. Kölreuter has also expressed his unbounded astonishment at the size and development of the tubers in certain hybrids; and all experimentalists[439] have remarked on the strong tendency in hybrids to increase by roots, runners, and suckers. Seeing that hybrid plants, which from their nature are more or less sterile, thus tend to produce double flowers; that they have the parts including the seed, that is the fruit, perfectly developed, even when containing no seed; that they sometimes yield gigantic roots; that they almost invariably tend to increase largely by suckers and other such means;--seeing this, and knowing, from the many facts given in the earlier parts of this chapter, that almost all organic beings when exposed to unnatural conditions tend to become more or less sterile, it seems much the most probable view that with cultivated plants sterility is the exciting cause, and double flowers, rich seedless fruit, and in some cases largely-developed organs of vegetation, &c., are the indirect results--these results having been in most cases largely increased through continued selection by man.
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