Chapter 1: XI (2)

[Throughout Europe, in different soils and under different climates, branches on this tree have repeatedly and suddenly reverted to the two parent species in their flowers and leaves. To behold mingled on the same tree tufts of dingy-red, bright yellow, and purple flowers, borne on branches having widely different leaves and manner of growth, is a surprising sight. The same raceme sometimes bears two kinds of flowers; and I have seen a single flower exactly divided into halves, one side being bright yellow and the other purple; so that one half of the standard-petal was yellow and of larger size, and the other half purple and smaller. In another flower the whole corolla was bright yellow, but exactly half the calyx was purple. In another, one of the dingy-red wing-petals had a narrow bright yellow stripe on it; and lastly, in another flower, one of the stamens, which had become slightly foliaceous, was half yellow and half purple; so that the tendency to segregation of character or reversion affects even single parts and organs. (11/91. For analogous facts see Braun 'Rejuvenescence' in 'Ray Soc. Bot. Mem.' 1853 page 320; and 'Gardener's Chronicle' 1842 page 397; also Braun in 'Sitzungsberichte der Ges. naturforschender Freunde' June 1873 page 63.) The most remarkable fact about this tree is that in its intermediate state, even when growing near both parent-species, it is quite sterile; but when the flowers become pure yellow or pure purple they yield seed. I believe that the pods from the yellow flowers yield a full complement of seed; they certainly yield a larger number. Two seedlings raised by Mr. Herbert from such seed (11/92. 'Journal of Hort. Soc.' volume 2 1847 page 100.) exhibited a purple tinge on the stalks of their flowers; but several seedlings raised by myself resembled in every character the common laburnum, with the exception that some of them had remarkably long racemes: these seedlings were perfectly fertile. That such purity of character and fertility should be suddenly reacquired from so hybridised and sterile a form is an astonishing phenomenon. The branches with purple flowers appear at first sight exactly to resemble those of C. purpureus; but on careful comparison I found that they differed from the pure species in the shoots being thicker, the leaves a little broader, and the flowers slightly shorter, with the corolla and calyx less brightly purple: the basal part of the standard-petal also plainly showed a trace of the yellow stain. So that the flowers, at least in this instance, had not perfectly recovered their true character; and in accordance with this, they were not perfectly fertile, for many of the pods contained no seed, some produced one, and very few contained as many as two seeds; whilst numerous pods on a tree of the pure C. purpureus in my garden contained three, four, and five fine seeds. The pollen, moreover, was very imperfect, a multitude of grains being small and shrivelled; and this is a singular fact; for, as we shall immediately see, the pollen-grains in the dingy-red and sterile flowers on the parent-tree, were, in external appearance, in a much better state, and included very few shrivelled grains. Although the pollen of the reverted purple flowers was in so poor a condition, the ovules were well formed, and the seeds, when mature, germinated freely with me. Mr. Herbert raised plants from seeds of the reverted purple flowers, and they differed a VERY LITTLE from the usual state of C. purpureus. Some which I raised in the same manner did not differ at all, either in the character of their flowers or of the whole bush, from the pure C. purpureus. Prof. Caspary has examined the ovules of the dingy-red and sterile flowers in several plants of C. adami on the Continent (11/93. See 'Transact. of Hort. Congress of Amsterdam' 1865; but I owe most of the following information to Prof. Caspary's letters.) and finds them generally monstrous. In three plants examined by me in England, the ovules were likewise monstrous, the nucleus varying much in shape, and projecting irregularly beyond the proper coats. The pollen grains, on the other hand, judging from their external appearance, were remarkably good, and readily protruded their tubes. By repeatedly counting, under the microscope, the proportional number of bad grains, Prof. Caspary ascertained that only 2.5 per cent were bad, which is a less proportion than in the pollen of three pure species of Cytisus in their cultivated state, viz., C. purpureus, laburnum, and alpinus. Although the pollen of C. adami is thus in appearance good, it does not follow, according to M. Naudin's observation (11/94. 'Nouvelles Archives du Museum' tome 1 page 143.) on Mirabilis, that it would be functionally effective. The fact of the ovules of C. adami being monstrous, and the pollen apparently sound, is all the more remarkable, because it is opposed to what usually occurs not only with most hybrids (11/95. See on this head Naudin ibid page 141.), but with two hybrids in the same genus, namely in C. purpureo-elongatus, and C. alpino-laburnum. In both these hybrids, the ovules, as observed by Prof. Caspary and myself, were well-formed, whilst many of the pollen-grains were ill-formed; in the latter hybrid 20.3 per cent, and in the former no less than 84.8 per cent of the grains were ascertained by Prof. Caspary to be bad. This unusual condition of the male and female reproductive elements in C. adami has been used by Prof. Caspary as an argument against this plant being considered as an ordinary hybrid produced from seed; but we should remember that with hybrids the ovules have not been examined nearly so frequently as the pollen, and they may be much oftener imperfect than is generally supposed. Dr. E. Bornet, of Antibes, informs me (through Mr. J. Traherne Moggridge) that with hybrid Cisti the ovarium is frequently deformed, the ovules being in some cases quite absent, and in other cases incapable of fertilisation.

Several theories have been propounded to account for the origin of C. adami, and for the transformations which it undergoes. The whole case has been attributed by some authors to bud-variation; but considering the wide difference between C. laburnum and purpureus, both of which are natural species, and considering the sterility of the intermediate form, this view may be summarily rejected. We shall presently see that, with hybrid plants, two embryos differing in their characters may be developed within the same seed and cohere; and it has been supposed that C. adami thus originated. Many botanists maintain that C. adami is a hybrid produced in the common way by seed, and that it has reverted by buds to its two parent-forms. Negative results are not of much value; but Reisseck, Caspary, and myself, tried in vain to cross C. laburnum and purpureus; when I fertilised the former with pollen of the latter, I had the nearest approach to success, for pods were formed, but in sixteen days after the withering of the flowers, they fell off. Nevertheless, the belief that C. adami is a spontaneously produced hybrid between these two species is supported by the fact that such hybrids have arisen in this genus. In a bed of seedlings from C. elongatus, which grew near to C. purpureus, and was probably fertilised by it through the agency of insects (for these, as I know by experiment, play an important part in the fertilisation of the laburnum), the sterile hybrid C. purpureo-elongatus appeared. (11/96. Braun in 'Bot. Mem. Ray. Soc.' 1853 page 23.) Thus, also, Waterer's laburnum, the C. alpino-laburnum (11/97. This hybrid has never been described. It is exactly intermediate in foliage, time of flowering, dark striae at the base of the standard petal, hairiness of the ovarium, and in almost every other character, between C. laburnum and alpinus; but it approaches the former species more nearly in colour, and exceeds it in the length of the racemes. We have before seen that 20.3 per cent of its pollen-grains are ill-formed and worthless. My plant, though growing not above thirty or forty yards from both parent-species, during some seasons yielded no good seeds; but in 1866 it was unusually fertile, and its long racemes produced from one to occasionally even four pods. Many of the pods contained no good seeds, but generally they contained a single apparently good seed, sometimes two, and in one case three seeds. Some of these seeds germinated, and I raised two trees from them; one resembles the present form; the other has a remarkable dwarf character with small leaves, but has not yet flowered.) spontaneously appeared, as I am informed by Mr. Waterer, in a bed of seedlings.

On the other hand, we have a clear and distinct account given to Poiteau (11/98. 'Annales de la Soc. de l'Hort. de Paris' tome 7 1830 page 93.) by M. Adam, who raised the plant, showing that C. adami is not an ordinary hybrid; but is what may be called a graft-hybrid, that is, one produced from the united cellular tissue of two distinct species. M. Adam inserted in the usual manner a shield of the bark of C. purpureus into a stock of C. laburnum; and the bud lay dormant, as often happens, for a year; the shield then produced many buds and shoots, one of which grew more upright and vigorous with larger leaves than the shoots of C. purpureus, and was consequently propagated. Now it deserves especial notice that these plants were sold by M. Adam, as a variety of C. purpureus, before they had flowered; and the account was published by Poiteau after the plants had flowered, but before they had exhibited their remarkable tendency to revert into the two parent species. So that there was no conceivable motive for falsification, and it is difficult to see how there could have been any error. (11/99. An account was given in the 'Gardener's Chronicle' 1857 pages 382, 400, of a common laburnum on which grafts of C. purpureus had been inserted, and which gradually assumed the character of C. adami; but I have little doubt that C. adami had been sold to the purchaser, who was not a botanist, in the place of C. purpureus. I have ascertained that this occurred in another instance.) If we admit as true M. Adam's account, we must admit the extraordinary fact that two distinct species can unite by their cellular tissue, and subsequently produce a plant bearing leaves and sterile flowers intermediate in character between the scion and stock, and producing buds liable to reversion; in short, resembling in every important respect a hybrid formed in the ordinary way by seminal reproduction.]

I will therefore give all the facts which I have been able to collect on the formation of hybrids between distinct species or varieties, without the intervention of the sexual organs. For if, as I am now convinced, this is possible, it is a most important fact, which will sooner or later change the views held by physiologists with respect to sexual reproduction. A sufficient body of facts will afterwards be adduced, showing that the segregation or separation of the characters of the two parent-forms by bud- variation, as in the case of Cytisus adami, is not an unusual though a striking phenomenon. We shall further see that a whole bud may thus revert, or only half, or some smaller segment.

[The famous bizzarria Orange offers a strictly parallel case to that of Cytisus adami. The gardener who in 1644 in Florence raised this tree, declared that it was a seedling which had been grafted; and after the graft had perished, the stock sprouted and produced the bizzarria. Gallesio, who carefully examined several living specimens and compared them with the description given by the original describer, P. Nato (11/100. Gallesio 'Gli Agrumi dei Giard. Bot. Agrar. di. Firenze' 1839 page 11. In his 'Traite du Citrus' 1811 page 146, he speaks as if the compound fruit consisted in part of a lemon, but this apparently was a mistake.), states that the tree produces at the same time leaves, flowers, and fruit identical with the bitter orange and with the citron of Florence, and likewise compound fruit, with the two kinds either blended together, both externally and internally, or segregated in various ways. This tree can be propagated by cuttings, and retains its diversified character. The so-called trifacial orange of Alexandria and Smyrna (11/101. 'Gardener's Chron.' 1855 page 628. See also Prof. Caspary in 'Transact. Hort. Congress of Amsterdam' 1865.) resembles in its general nature the bizzarria, and differs only in the orange being of the sweet kind; this and the citron are blended together in the same fruit, or are separately produced on the same tree; nothing is known of its origin. In regard to the bizzarria, many authors believe that it is a graft-hybrid; Gallesio, on the other hand, thinks that it is an ordinary hybrid, with the habit of partially reverting by buds to the two parent- forms; and we have seen that the species in this genus often cross spontaneously.

It is notorious that when the variegated Jessamine is budded on the common kind, the stock sometimes produces buds bearing variegated leaves: Mr. Rivers, as he informs me, has seen instances of this. The same thing occurs with the Oleander. (11/102. Gartner 'Bastarderzeugung' s. 611 gives many references on this subject.) Mr. Rivers, on the authority of a trustworthy friend, states that some buds of a golden-variegated ash, which were inserted into common ashes, all died except one; but the ash-stocks were affected (11/103. A nearly similar account was given by Brabley in 1724 in his 'Treatise on Husbandry' volume 1 page 199.) and produced, both above and below the points of insertion of the plates of bark bearing the dead buds, shoots which bore variegated leaves. Mr. J. Anderson Henry has communicated to me a nearly similar case: Mr. Brown, of Perth, observed many years ago, in a Highland glen, an ash-tree with yellow leaves; and buds taken from this tree were inserted into common ashes, which in consequence were affected, and produced the Blotched Breadalbane Ash. This variety has been propagated, and has preserved its character during the last fifty years. Weeping ashes, also, were budded on the affected stocks, and became similarly variegated. It has been repeatedly proved that several species of Abutilon, on which the variegated A. thompsonii has been grafted, become variegated. (11/104. Morren 'Bull. de l'Acad. R. des Sciences de Belgique' 2de series tome 28 1869 page 434. Also Magnus 'Gesellschaft naturforschender Freunde, Berlin' February 21, 1871 page 13; ibid June 21, 1870 and October 17, 1871. Also 'Bot. Zeitung' February 24, 1871.)

Many authors consider variegation as the result of disease; and the foregoing cases may be looked at as the direct result of the inoculation of a disease or some weakness. This has been almost proved to be the case by Morren in the excellent paper just referred to, who shows that even a leaf inserted by its footstalk into the bark of the stock is sufficient to communicate variegation to it, though the leaf soon perishes. Even fully formed leaves on the stock of Abutilon are sometimes affected by the graft and become variegated. Variegation is much influenced, as we shall hereafter see, by the nature of the soil in which the plants are grown; and it does not seem improbable that whatever change in the sap or tissues certain soils induce, whether or not called a disease, might spread from the inserted piece of bark to the stock. But a change of this kind cannot be considered to be of the nature of a graft-hybrid.

There is a variety of the hazel with dark-purple leaves, like those of the copper-beech: no one has attributed this colour to disease, and it apparently is only an exaggeration of a tint which may often be seen on the leaves of the common hazel. When this variety is grafted on the common hazel (11/105. Loudon's 'Arboretum' volume 4 page 2595.), it sometimes colours, as has been asserted, the leaves below the graft; although negative evidence is not of much value, I may add that Mr. Rivers, who has possessed hundreds of such grafted trees, has never seen an instance.

Gartner (11/106. 'Bastarderzeugung' s. 619.) quotes two separate accounts of branches of dark and white-fruited vines which had been united in various ways, such as being split longitudinally, and then joined, etc.; and these branches produced distinct bunches of grapes of the two colours, and other bunches with berries, either striped, or of an intermediate and new tint. Even the leaves in one case were variegated. These facts are the more remarkable because Andrew Knight never succeeded in raising variegated grapes by fertilising white kinds by pollen of dark kinds; though, as we have seen, he obtained seedlings with variegated fruits and leaves, by fertilising a white variety by the already variegated dark Aleppo grape. Gartner attributes the above-quoted cases merely to bud-variation; but it is a strange coincidence that the branches which had been grafted in a peculiar manner should alone thus have varied; and H. Adorne de Tscharner positively asserts that he produced the described result more than once, and could do so at will, by splitting and uniting the branches in the manner described by him.

I should not have quoted the following case had not the author of 'Des Jacinthes' (11/107. Amsterdam 1768 page 124.) impressed me with the belief not only of his extensive knowledge, but of his truthfulness: he says that bulbs of blue and red hyacinths may be cut in two, and that they will grow together and throw up a united stem (and this I have myself seen) with flowers of the two colours on the opposite sides. But the remarkable point is, that flowers are sometimes produced with the two colours blended together, which makes the case closely analogous with that of the blended colours of the grapes on the united vine branches.

In the case of roses it is supposed that several graft-hybrids have been formed, but there is much doubt about these cases, owing to the frequency of ordinary bud-variations. The most trustworthy instance known to me is one, recorded by Mr. Poynter (11/108. 'Gardener's Chron.' 1860 page 672 with a woodcut.) who assures me in a letter of the entire accuracy of the statement. Rosa devoniensis had been budded some years previously on a white Banksian rose; and from the much enlarged point of junction, whence the Devoniensis and Banksian still continued to grow, a third branch issued, which was neither pure Banksian nor pure Devoniensis, but partook of the character of both; the flowers resembled, but were superior in character to those of the variety called Lamarque (one of the Noisettes), while the shoots were similar in their manner of growth to those of the Banksian rose, with the exception that the longer and more robust shoots were furnished with prickles. This rose was exhibited before the Floral Committee of the Horticultural Society of London. Dr. Lindley examined it and concluded that it had certainly been produced by the mingling of R. banksiae with some rose like R. devoniensis, "for while it was very greatly increased in vigour and in size of all the parts, the leaves were half-way between a Banksian and Tea-scented rose." It appears that rose-growers were previously aware that the Banksian rose sometimes affects other roses. As Mr. Poynter's new variety is intermediate in its fruit and foliage between the stock and scion, and as it arose from the point of junction between the two, it is very improbable that it owes its origin to mere bud-variation, independently of the mutual influence of the stock and scion.

Lastly, with respect to potatoes. Mr. R. Trail stated in 1867 before the Botanical Society of Edinburgh (and has since given me fuller information), that several years ago he cut about sixty blue and white potatoes into halves through the eyes or buds, and then carefully joined them, destroying at the same time the other eyes. Some of these united tubers produced white, and others blue tubers; some, however, produced tubers partly white and partly blue; and the tubers from about four or five were regularly mottled with the two colours. In these latter cases we may conclude that a stem had been formed by the union of the bisected buds, that is, by graft- hybridisation.

In the 'Botanische Zeitung' (May 16, 1868), Professor Hildebrand gives an account with a coloured figure, of his experiments on two varieties which were found during the same season to be constant in character, namely, a somewhat elongated rough-skinned red potato and a rounded smooth white one. He inserted buds reciprocally into both kinds, destroying the other buds. He thus raised two plants, and each of these produced a tuber intermediate in character between the two parent-forms. That from the red bud grafted into the white tuber, was at one end red and rough, as the whole tuber ought to have been if not affected; in the middle it was smooth with red stripes, and at the other end smooth and altogether white like that of the stock.

Mr. Taylor, who had received several accounts of potatoes having been grafted by wedge-shaped pieces of one variety inserted into another, though sceptical on the subject, made twenty-four experiments which he described in detail before the Horticultural Society. (11/109. 'Gardener's Chronicle' 1869 page 220.) He thus raised many new varieties, some like the graft or like the stock; others having an intermediate character. Several persons witnessed the digging up of the tubers from these graft-hybrids; and one of them, Mr. Jameson, a large dealer in potatoes, writes thus, "They were such a mixed lot, as I have never before or since seen. They were of all colours and shapes, some very ugly and some very handsome." Another witness says "some were round, some kidney, pink-eyed kidney, piebald, and mottled red and purple, of all shapes and sizes." Some of these varieties have been found valuable, and have been extensively propagated. Mr. Jameson took away a large piebald potato which he cut into five sets and propagated; these yielded round, white, red, and piebald potatoes.

Mr. Fitzpatrick followed a different plan (11/110. 'Gard. Chronicle' 1869 page 335.); he grafted together not the tubers but the young stems of varieties producing black, white, and red potatoes. The tubers borne by three of these twin or united plants were coloured in an extraordinary manner; one was almost exactly half black and half white, so that some persons on seeing it thought that two potatoes had been divided and rejoined; other tubers were half red and half white, or curiously mottled with red and white, or with red and black, according to the colours of the graft and stock.

The testimony of Mr. Fenn is of much value, as he is "a well known potato- grower" who has raised many new varieties by crossing different kinds in the ordinary manner. He considers it "demonstrated" that new, intermediate varieties can be produced by grafting the tubers, though he doubts whether such will prove valuable. (11/111. 'Gardener's Chronicle' 1869 page 1018 with remarks by Dr. Masters on the adhesion of the united wedges. See also ibid 1870 pages 1277, 1283.) He made many trials and laid the results, exhibiting specimens, before the Horticultural Society. Not only were the tubers affected, some being smooth and white at one end and rough and red at the other, but the stems and leaves were modified in their manner of growth, colour and precocity. Some of these graft-hybrids after being propagated for three years still showed in their haulms their new character, different from that of the kind from which the eyes had been taken. Mr. Fenn gave twelve of the tubers of the third generation to Mr. Alex. Dean, who grew them, and was thus converted into a believer in graft- hybridisation, having previously been a complete sceptic. For comparison he planted the pure parent-forms alongside the twelve tubers; and found that many of the plants from the latter (11/112. 'Gard. Chronicle' 1871 page 837.) were intermediate between the two parent-forms in precocity, in the tallness, uprightness, jointing, and robustness of the stems, and in the size and colour of the leaves.

Another experimentalist, Mr. Rintoul, grafted no less than fifty-nine tubers, which differed in shape (some being kidneys) in smoothness and colour (11/113. 'Gardener's Chronicle' 1870 page 1506.) and many of the plants thus raised "were intermediate in the tubers as well as in the haulms." He describes the more striking cases.

In 1871 I received a letter from Mr. Merrick, of Boston, U.S.A., who states that, "Mr. Fearing Burr, a very careful experimenter and author of a much valued book, 'The Garden Vegetables of America' has succeeded in producing distinctly mottled and most curious potatoes--evidently graft-hybrids, by inserting eyes from blue or red potatoes into the substance of white ones, after removing the eyes of the latter. I have seen the potatoes, and they are very curious."

We will now turn to the experiments made in Germany, since the publication of Prof. Hildebrand's paper. Herr Magnus relates (11/114. 'Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin' October 17, 1871.) the results of numerous trials made by Herren Reuter and Lindemuth, both attached to the Royal Gardens of Berlin. They inserted the eyes of red potatoes into white ones, and vice versa. Many different forms partaking of the characters of the inserted bud and of the stock were thus obtained; for instance, some of the tubers were white with red eyes.

Herr Magnus also exhibited in the following year before the same Society (November 19, 1872), the produce of grafts between black, white, and red potatoes, made by Dr. Neubert. These were made by uniting not the tubers but the young stems, as was done by Mr. Fitzpatrick. The result was remarkable, inasmuch as all the tubers thus produced were intermediate in character, though in a variable degree. Those between the black and the white or the red were the most striking in appearance. Some from between the white and red had one half of one colour and the other half of the other colour.

At the next meeting of the society Herr Magnus communicated the results of Dr. Heimann's experiments in grafting together the tubers of red Saxon, blue, and elongated white potatoes. The eyes were removed by a cylindrical instrument, and inserted into corresponding holes in other varieties. The plants thus produced yielded a great number of tubers, which were intermediate between the two parent-forms in shape, and in the colour both of the flesh and skin.

Herr Reuter experimented (11/115. Ibid November 17, 1874. See also excellent remarks by Herr Magnus.) by inserting wedges of the elongated White Mexican potato into a Black Kidney potato. Both sorts are known to be very constant, and differ much not only in form and colour, but in the eyes of the Black Kidney being deeply sunk, whereas those of the White Mexican are superficial and of a different shape. The tubers produced by these hybrids were intermediate in colour and form; and some which resembled in form the graft, i.e. the Mexican, had eyes deeply sunk and of the same shape as in the stock or Black Kidney.]

Any one who will attentively consider the abstract now given, of the experiments made by many observers in several countries, will, I think, be convinced that by grafting two varieties of the potato together in various ways, hybridised plants can be produced. It should be observed that several of the experimentalists are scientific horticulturists, and some of them potato-growers on a large scale, who, though beforehand sceptical, have been fully convinced of the possibility, even of the ease, of making graft- hybrids. The only way of escaping from this conclusion is to attribute all the many recorded cases to simple bud-variation. Undoubtedly the potato, as we have seen in this chapter, does sometimes, though not often, vary by buds; but it should be especially noted that it is experienced potato- growers, whose business it is to look out for new varieties, who have expressed unbounded astonishment at the number of new forms produced by graft-hybridisation. It may be argued that it is merely the operation of grafting, and not the union of two kinds, which causes so extraordinary an amount of bud-variation; but this objection is at once answered by the fact that potatoes are habitually propagated by the tubers being cut into pieces, and the sole difference in the case of graft-hybrids is that either a half or a smaller segment or a cylinder is placed in close opposition with the tissue of another variety. Moreover, in two cases, the young stems were grafted together, and the plants thus united yielded the same results as when the tubers were united. It is an argument of the greatest weight that when varieties are produced by simple bud-variation, they frequently present quite new characters; whereas in all the numerous cases above given, as Herr Magnus likewise insists, the graft-hybrids are intermediate in character between the two forms employed. That such a result should follow if the one kind did not affect the other is incredible.

Characters of all kinds are affected by graft hybridisation, in whatever way the grafting may have been effected. The plants thus raised yield tubers which partake of the widely different colours, form, state of surface, position and shape of the eye of the parents; and according to two careful observers they are also intermediate in certain constitutional peculiarities. But we should bear in mind that in all the varieties of the potato, the tubers differ much more than any other part.

The potato affords the best evidence of the possibility of the formation of graft-hybrids, but we must not overlook the account given of the origin of the famous Cytisus adami by M. Adam, who had no conceivable motive for deception, and the exactly parallel account of the origin of the Bizzarria orange, namely by graft-hybridisation. Nor must the cases be undervalued in which different varieties or species of vines, hyacinths and roses, have been grafted together, and have yielded intermediate forms. It is evident that graft-hybrids can be made much more easily with some plants, as the potato, than with others, for instance our common fruit trees; for these latter have been grafted by the million during many centuries, and though the graft is often slightly affected, it is very doubtful whether this may not be accounted for, merely by a more or less free supply of nutriment. Nevertheless, the cases above given seem to me to prove that under certain unknown conditions graft-hybridisation can be effected.

Herr Magnus asserts with much truth that graft-hybrids resemble in all respects seminal hybrids, including their great diversity of character. There is, however, a partial exception, inasmuch as the characters of the two parent forms are not often homogeneously blended together in graft- hybrids. They much more commonly appear in a segregated condition,--that is, in segments either at first, or subsequently through reversion. It would seem that the reproductive elements are not so completely blended by grafting as by sexual generation. But segregation of this kind occurs by no means rarely, as will be immediately shown, in seminal hybrids. Finally it must, I think, be admitted that we learn from the foregoing cases a highly important physiological fact, namely, that the elements that go to the production of a new being, are not necessarily formed by the male and female organs. They are present in the cellular tissue in such a state that they can unite without the aid of the sexual organs, and thus give rise to a new bud partaking of the characters of the two parent-forms.

ON THE SEGREGATION OF THE PARENTAL CHARACTERS IN SEMINAL HYBRIDS BY BUD- VARIATION.

I will now give a sufficient number of cases to show that segregation of this kind, namely, by buds, may occur in ordinary hybrids raised from seed.

[Hybrids were raised by Gartner between Tropaeolum minus and majus (11/116. 'Bastarderzeugung' s. 549. It is, however, doubtful whether these plants should be ranked as species or varieties.) which at first produced flowers intermediate in size, colour, and structure between their two parents; but later in the season some of these plants produced flowers in all respects like those of the mother-form, mingled with flowers still retaining the usual intermediate condition. A hybrid Cereus between C. speciosissimus and phyllanthus (11/117. Gartner ibid s. 550.) plants which are widely different in appearance, produced for the first three years angular, five- sided stems, and then some flat stems like those of C. phyllanthus. Kolreuter also gives cases of hybrid Lobelias and Verbascums, which at first produced flowers of one colour, and later in the season, flowers of a different colour. (11/118. 'Journal de Physique' tome 23 1873 page 100. 'Act. Acad. St. Petersburgh' 1781 part 1 page 249.) Naudin (11/119. 'Nouvelles Archives du Museum' tome 1 page 49.) raised forty hybrids from Datura laevis fertilised by D. stramonium; and three of these hybrids produced many capsules, of which a half, or quarter, or lesser segment was smooth and of small size, like the capsule of the pure D. laevis, the remaining part being spinose and of larger size, like the capsule of the pure D. stramonium: from one of these composite capsules, plants perfectly resembling both parent-forms were raised.

Turning now to varieties. A SEEDLING apple, conjectured to be of crossed parentage, has been described in France (11/120. L'Hermes January 14, 1837 quoted in Loudon's 'Gardener's Mag.' volume 13 page 230.) which bears fruit with one half larger than the other, of a red colour, acid taste, and peculiar odour; the other side being greenish-yellow and very sweet: it is said scarcely ever to include perfectly developed seed. I suppose that this is not the same tree as that which Gaudichaud (11/121. 'Comptes Rendus' tome 34 1852 page 746.) exhibited before the French institute, bearing on the same branch two distinct kinds of apples, one a reinette rouge, and the other like a reinette canada jaunatre: this double-bearing variety can be propagated by grafts, and continues to produce both kinds; its origin is unknown. The Rev. J.D. La Touche sent me a coloured drawing of an apple which he brought from Canada, of which half, surrounding and including the whole of the calyx and the insertion of the foot-stalk, is green, the other half being brown and of the nature of the pomme gris apple, with the line of separation between the two halves exactly defined. The tree was a grafted one, and Mr. La Touche thinks that the branches which bore this curious apple sprung from the point of junction of the graft and stock: had this fact been ascertained, the case would probably have come into the class of graft-hybrids already given. But the branch may have sprung from the stock, which no doubt was a seedling.

Prof. H. Lecoq, who has made a great number of crossings between the differently coloured varieties of Mirabilis jalapa (11/122. 'Geograph. Bot. de l'Europe' tome 3 1854 page 405; and 'De la Fecondation' 1862 page 302.) finds that in the seedlings the colours rarely combine, but form distinct stripes; or half the flower is of one colour and half of a different colour. Some varieties regularly bear flowers striped with yellow, white, and red; but plants of such varieties occasionally produce on the same root branches with uniformly coloured flowers of all three tints, and other branches with half-and-half coloured flowers, and others with marbled flowers. Gallesio (11/123. 'Traite du Citrus' 1811 page 45.) crossed reciprocally white and red carnations, and the seedlings were striped; but some of the striped plants also bore entirely white and entirely red flowers. Some of these plants produced one year red flowers alone, and in the following year striped flowers; or conversely, some plants, after having borne for two or three years striped flowers, would revert and bear exclusively red flowers. It may be worth mentioning that I fertilised the PURPLE SWEET-PEA (Lathyrus odoratus) with pollen from the light-coloured PAINTED LADY: seedlings raised from the same pod were not intermediate in character, but perfectly resembled either parent. Later in the summer, the plants which had at first borne flowers identical with those of the PAINTED LADY, produced flowers streaked and blotched with purple; showing in these darker marks a tendency to reversion to the mother-variety. Andrew Knight (11/124. 'Transact. Linn. Soc.' volume 9 page 268.) fertilised two white grapes with pollen of the Aleppo grape, which is darkly variegated both in its leaves and fruit. The result was that the young seedlings were not at first variegated, but all became variegated during the succeeding summer; besides this, many produced on the same plant bunches of grapes which were all black, or all white, or lead-coloured striped with white, or white dotted with minute black stripes; and grapes of all these shades could frequently be found on the same foot-stalk.

I will append a very curious case, not of bud-variation, but of two cohering embryos, different in character and contained within the same seed. A distinguished botanist, Mr. G.H. Thwaites (11/125. 'Annals and Mag. of Nat. Hist.' March 1848.) states that a seed from Fuchsia coccinea fertilised by F. fulgens, contained two embryos, and was "a true vegetable twin." The two plants produced from the two embryos were "extremely different in appearance and character," though both resembled other hybrids of the same parentage produced at the same time. These twin plants "were closely coherent, below the two pairs of cotyledon-leaves, into a single cylindrical stem, so that they had subsequently the appearance of being branches on one trunk." Had the two united stems grown up to their full height, instead of dying, a curiously mixed hybrid would have been produced. A mongrel melon described by Sageret (11/126. 'Pomologie Physiolog.' 1830 page 126.) may perhaps have thus originated; for the two main branches, which arose from two cotyledon-buds, produced very different fruit,--on the one branch like that of the paternal variety, and on the other branch like to a certain extent that of the maternal variety, the melon of China.]

In most of these cases of crossed varieties, and in some of the cases of crossed species, the colours proper to both parents appeared in the seedlings, as soon as they first flowered, in the form of stripes or larger segments, or as whole flowers or fruit of different kinds borne on the same plant; and in this case the appearance of the two colours cannot strictly be said to be due to reversion, but to some incapacity of fusion. When, however, the later flowers or fruit produced during the same season, or during a succeeding year or generation, become striped or half-and-half, etc., the segregation of the two colours is strictly a case of reversion by bud-variation. Whether all the many recorded cases of striped flowers and fruit are due to previous hybridisation and reversion is by no means clear, for instance with peaches and nectarines, moss-roses, etc. In a future chapter I shall show that, with animals of crossed parentage, the same individual has been known to change its character during growth, and to revert to one of its parents which it did not at first resemble. Finally, from the various facts now given, there can be no doubt that the same individual plant, whether a hybrid or a mongrel, sometimes returns in its leaves, flowers, and fruit, either wholly or by segments, to both parent- forms.

ON THE DIRECT OR IMMEDIATE ACTION OF THE MALE ELEMENT ON THE MOTHER FORM.

Another remarkable class of facts must be here considered, firstly, because they have a high physiological importance, and secondly, because they have been supposed to account for some cases of bud-variation. I refer to the direct action of the male element, not in the ordinary way on the ovules, but on certain parts of the female plant, or in case of animals on the subsequent progeny of the female by a second male. I may premise that with plants the ovarium and the coats of the ovules are obviously parts of the female, and it could not have been anticipated that they would have been affected by the pollen of a foreign variety or species, although the development of the embryo, inside the embryonic sack, inside the ovule and ovarium, of course, depends on the male element.

[Even as long ago as 1729 it was observed (11/127. 'Philosophical Transact.' volume 43 1744-45 page 525.) that white and blue varieties of the Pea, when planted near each other, mutually crossed, no doubt through the agency of bees, and in the autumn blue and white peas were found within the same pods. Wiegmann made an exactly similar observation in the present century. The same result has followed several times when a variety with peas of one colour has been artificially crossed by a differently-coloured variety. (11/128. Mr. Goss 'Transact. Hort. Soc.' volume 5 page 234: and Gartner 'Bastarderzeugung' 1849 ss. 81 and 499.) These statements led Gartner, who was highly sceptical on the subject, carefully to try a long series of experiments: he selected the most constant varieties, and the result conclusively showed that the colour of the skin of the pea is modified when pollen of a differently coloured variety is used. This conclusion has since been confirmed by experiments made by the Rev. J.M. Berkeley. (11/129. 'Gardener's Chronicle' 1854 page 404.)

Mr. Laxton of Stamford, whilst making experiments on peas for the express purpose of ascertaining the influence of foreign pollen on the mother- plant, has recently (11/130. Ibid 1866 page 900.) observed an important additional fact. He fertilised the Tall Sugar-pea, which bears very thin green pods, becoming brownish-white when dry, with pollen of the Purple- podded pea, which, as its name expresses, has dark-purple pods with very thick skin, becoming pale reddish purple when dry. Mr. Laxton has cultivated the tall sugar-pea during twenty years, and has never seen or heard of it producing a purple pod: nevertheless, a flower fertilised by pollen from the purple-pod yielded a pod clouded with purplish-red which Mr. Laxton kindly gave to me. A space of about two inches in length towards the extremity of the pod, and a smaller space near the stalk, were thus coloured. On comparing the colour with that of the purple pod, both pods having been first dried and then soaked in water, it was found to be identically the same; and in both the colour was confined to the cells lying immediately beneath the outer skin of the pod. The valves of the crossed pod were also decidedly thicker and stronger than those of the pods of the mother-plant, but this may possibly have been an accidental circumstance, for I know not how far their thickness is a variable character in the Tall Sugar-pea.

The peas of the Tall Sugar-pea, when dry, are pale greenish-brown, thickly covered with dots of dark purple so minute as to be visible only through a lens, and Mr. Laxton has never seen or heard of this variety producing a purple pea; but in the crossed pod one of the peas was of a uniform beautiful violet-purple tint, and a second was irregularly clouded with pale purple. The colour lies in the outer of the two coats which surround the pea. As the peas of the purple-podded variety when dry are of a pale greenish-buff, it would at first appear that this remarkable change of colour in the peas in the crossed pod could not have been caused by the direct action of the pollen of the purple-pod: but when we bear in mind that this latter variety has purple flowers, purple marks on its stipules, and purple pods; and that the Tall Sugar-pea likewise has purple flowers and stipules, and microscopically minute purple dots on the peas, we can hardly doubt that the tendency to the production of purple in both parents has in combination modified the colour of the peas in the crossed pod. After having examined these specimens, I crossed the same two varieties, and the peas in one pod but not the pods themselves, were clouded and tinted with purplish-red in a much more conspicuous manner than the peas in the uncrossed pods produced at the same time by the same plants. I may notice as a caution that Mr. Laxton sent me various other crossed peas slightly, or even greatly, modified in colour; but the change in these cases was due, as had been suspected by Mr. Laxton, to the altered colour of the cotyledons, seen through the transparent coats of the peas; and as the cotyledons are parts of the embryo, these cases are not in any way remarkable.

Turning now to the genus Matthiola. The pollen of one kind of stock sometimes affects the colour of the seeds of another kind, used as the mother-plant. I give the following case the more readily, as Gartner doubted similar statements previously made with respect to the stock by other observers. A well-known horticulturist, Major Trevor Clarke, informs me (11/131. See also a paper by this observer read before the International Hort. and Bot. Congress of London 1866.) that the seeds of the large red- flowered BIENNIAL stock, Matthiola annua (Cocardeau of the French), are light brown, and those of the purple branching Queen stock (M. incana) are violet-black; and he found that, when flowers of the red stock were fertilised by pollen from the purple stock, they yielded about fifty per cent of BLACK seeds. He sent me four pods from a red flowered plant, two of which had been fertilised by their own pollen, and they included pale brown seed; and two which had been crossed by pollen from the purple kind, and they included seeds all deeply tinged with black. These latter seeds yielded purple-flowered plants like their father; whilst the pale brown seeds yielded normal red-flowered plants; and Major Clarke, by sowing similar seeds, has observed on a greater scale the same result. The evidence in this case of the direct action of the pollen of one species on the colour of the seeds of another species appears to me conclusive.

Gallesio (11/132. 'Traite du Citrus' page 40.) fertilised the flowers of an orange with pollen from the lemon; and one fruit thus produced bore a longitudinal stripe of peel having the colour, flavour, and other characters of the lemon. Mr. Anderson (11/133. 'Transact. Hort. Soc.' volume 3 page 318. See also volume 5 page 65.) fertilised a green-fleshed melon with pollen from a scarlet-fleshed kind; in two of the fruits "a sensible change was perceptible: and four other fruits were somewhat altered both internally and externally." The seeds of the two first- mentioned fruits produced plants partaking of the good properties of both parents. In the United States, where Cucurbitaceae are largely cultivated, it is the popular belief (11/134. Prof. Asa Gray 'Proc. Acad. Sc.' Boston volume 4 1860 page 21. I have received statements to the same effect from other persons in the United States.) that the fruit is thus directly affected hy foreign pollen; and I have received a similar statement with respect to the cucumber in England. It is believed that grapes have been thus affected in colour, size, and shape: in France a pale-coloured grape had its juice tinted by the pollen of the dark-coloured Teinturier; in Germany a variety bore berries which were affected by the pollen of two adjoining kinds; some of the berries being only partially affected or mottled. (11/135. For the French case see 'Journ. Hort. Soc.' volume 1 new series 1866 page 50. For Germany see M. Jack quoted in Henfrey's 'Botanical Gazette' volume 1 page 277. A case in England has recently been alluded to by the Rev. J.M. Berkeley before the Hort. Soc. of London.)

As long ago as 1751 (11/136. 'Philosophical Transactions' volume 47 1751-52 page 206.) it was observed that, when differently-coloured varieties of maize grew near each other, they mutually affected each other's seeds, and this is now a popular belief in the United States. Dr. Savi (11/137. Gallesio 'Teoria della Riproduzione' 1816 page 95.) tried the experiment with care: he sowed yellow and black-seeded maize together, and on the same ear some of the seeds were yellow, some black, and some mottled, the differently coloured seeds being arranged irregularly or in rows. Prof. Hildebrand has repeated the experiment (11/138. 'Bot. Zeitung' May 1868 page 326.) with the precaution of ascertaining that the mother-plant was true. A kind bearing yellow grains was fertilised with pollen of a kind having brown grains, and two ears produced yellow grains mingled with others of a dirty violet tint. A third ear had only yellow grains, but one side of the spindle was tinted of a reddish-brown; so that here we have the important fact of the influence of the foreign pollen extending to the axis. Mr. Arnold, in Canada, varied the experiment in an interesting manner: "a female flower was subjected first to the action of pollen from a yellow variety, and then to that from a white variety; the result was an ear, each grain of which was yellow below and white above." (11/139. See Dr. J. Stockton-Hough 'American Naturalist' January 1874 page 29.) With other plants it has occasionally been observed that the crossed offspring showed the influence of two kinds of pollen, but in this case the two kinds affected the mother-plant.

Mr. Sabine states (11/140. 'Transact. Hort. Soc.' volume 5 page 69.) that he has seen the form of the nearly globular seed-capsule of Amaryllis vittata altered by the application of the pollen of another species, of which the capsule has gibbous angles. With an allied genus, a well-known botanist, Maximowicz, has described in detail the striking results of reciprocally fertilising Lilium bulbiferum and davuricum with each other's pollen. Each species produced fruit not like its own, but almost identical with that of the pollen-bearing species; but from an accident only the fruit of the latter species was carefully examined; the seeds were intermediate in the development of their wings. (11/141. 'Bull. de l'Acad. Imp. de St. Petersburg' tome 17 page 275, 1872. The author gives references to those cases in the So1anaceae of fruit affected by foreign pollen, but as it does not appear that the mother-plant was artificially fertilised, I have not entered into details.)

Fritz Muller fertilised Cattleya leopoldi with pollen of Epidendron cinnabarinum; and the capsules contained very few seeds; but these presented a most wonderful appearance, which, from the description given, two botanists, Hildebrand and Maximowicz, attribute to the direct action of the pollen of the Epidendron. (11/142. 'Bot. Zeitung' September 1868 page 631. For Maximowicz's judgment see the paper last referred to.)

Mr. J. Anderson Henry (11/143. 'Journal of Horticulture' January 20, 1863 page 46.) crossed Rhododendron dalhousiae with the pollen of R. nuttallii, which is one of the largest-flowered and noblest species of the genus. The largest pod produced by the former species, when fertilised with its own pollen, measured 1 2/8 inch in length and 1 1/2 in girth; whilst three of the pods which had been fertilised by pollen of R. nuttallii measured 1 5/8 inch in length and no less than 2 inches in girth. Here the effect of the foreign pollen was apparently confined to increasing the size of the ovarium; but we must be cautious in assuming, as the following case shows, that size had been transferred from the male parent to the capsule of the female plant. Mr. Henry fertilised Arabis blepharophylla with pollen of A. soyeri, and the pods thus produced, of which he was so kind as to send me detailed measurements and sketches, were much larger in all their dimensions than those naturally produced by either the male or female parent-species. In a future chapter we shall see that the organs of vegetation in hybrid plants, independently of the character of either parent, are sometimes developed to a monstrous size; and the increased size of the pods in the foregoing cases may be an analogous fact. On the other hand, M. de Saporta informs me that an isolated female plant of Pistacia vera is very apt to be fertilised by the pollen of neighbouring plants of P. terebinthus, and in this case the fruits are only half their proper size, which he attributes to the influence of the pollen of P. terebinthus.