The Variation of Animals and Plants under Domestication — Volume 2

The same law, namely, that the crossed offspring both of varieties and species are larger than the parent-forms, holds good in the most striking manner with hybrid animals as well as with mongrels. Mr. Bartlett, who has had such large experience says, "Among all hybrids of vertebrated animals there is a marked increase of size." He then enumerates many cases with mammals, including monkeys, and with various families of birds. (17/65. Quoted by Dr. Murie in 'Proc. Zoolog. Soc.' 1870 page 40.)]

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 the foregoing, as self-sterility is not here the result of long-continued 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 (17/66. 'Botanische Zeitung' January 1864 s. 3.): —

[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 and 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. (17/67. 'Monatsbericht Akad. Wissen.' Berlin 1866 s. 372.)]

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 is the fact, 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 (17/68. International Hort. Congress, London 1866.) with pollen from other plants of the same species; and fifty-eight capsules were obtained, including on an average 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. (17/69. 'Proc. Bot. Soc. of Edinburgh' May 1863: these observations are given in abstract, and others are added, in the 'Journal of Proc. of Linn. Soc.' volume 8 Bot. 1864 page 162.) Oncidium sphacelatum has effective pollen, for Mr. Scott fertilised two distinct species with it; the 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. Robertson 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 effective, 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. Riviere (17/70. Prof. Lecoq 'De la Fecondation' 2nd edition 1862 page 76.) 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 been grown under unnatural conditions in hot-houses, I concluded that their self-sterility was due to this cause. But Fritz Muller 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 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 only those flowers 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 Muller 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 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 Muller 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 fertilised 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 seeds produced by the plant's own pollen, were destitute of an embryo. Some self-fertilised capsules of a Maxillaria were in a similar state.

Another observation made by Fritz Muller 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 discoloration and decay are not caused by parasitic cryptograms, which were observed by Fritz Muller 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 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 Muller 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 Muller 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 fertilised 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 fertilised 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 fertilised 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 Muller suggests that, as in all these cases the plant's own pollen is not only impotent (thus effectually preventing self-fertilisation), 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.

The same naturalist found in Brazil three plants of a Bignonia growing near together. He fertilised twenty-nine flowerets on one of them with their own pollen, and they did not set a single capsule. Thirty flowers were then fertilised with pollen from a distinct plant, one of the three, and they yielded only two capsules. Lastly, five flowers were fertilised with pollen from a fourth plant growing at a distance, and all five produced capsules. Fritz Muller thinks that the three plants which grew near one another were probably seedlings from the same parent, and that from being closely related, they acted very feebly on one another. This view is extremely probable, for he has since shown in a remarkable paper (17/71. 'Jenaische Zeitschrift fur Naturwiss.' b. 7 page 22 1872 and page 441 1873. A large part of this paper has been translated in the 'American Naturalist' 1874 page 223.), that in the case of some Brazilian species of Abutilon, which are self-sterile, and between which he raised some complex hybrids, that these, if near relatives, were much less fertile inter se, than when not closely related.]

We now come to cases closely analogous with those just given, but different in so far that only certain individuals of the species are self-sterile. This self-impotence does not depend on the pollen or ovules being in an unfit state for fertilisation, for both have been found effective in union with other plants of the same or of a distinct species. The fact of plants having acquired so peculiar a constitution, that they can be fertilised more readily by the pollen of a distinct species than by their own, is exactly the reverse of what occurs with all ordinary species. For in the latter the two sexual elements of the same individual plant are of course 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.

[Gartner experimented on two plants of Lobelia fulgens, brought from separate places, and found (17/72. 'Bastarderzeugung' s. 64, 357.) 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 Gartner (17/73. Ibid s. 357.) 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. Kolreuter, also (17/74. 'Zweite Fortsetzung' s. 10; 'Dritte Forts.' s. 40. Mr. Scott likewise fertilised fifty-four flowers of Verbascum phoeniceum, including two varieties, with their own pollen, and not a single capsule was produced. Many of the pollen-grains emitted their tubes, but only a few of them penetrated the stigmas; some slight effect however was produced, as many of the ovaries became somewhat developed: 'Journal Asiatic Soc. Bengal' 1867 page 150.), gives the case of three garden plants of Verbascum phoeniceum, which bore during two years many flowers; these he fertilised successfully with 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.

With Reseda odorata I have found certain individuals quite sterile with their own pollen, and so it is with the indigenous Reseda lutea. The self-sterile plants of both species were perfectly fertile when crossed with pollen from any other individual of the same species. These observations will hereafter be published in another work, in which I shall also show that seeds sent to me by Fritz Muller produced by plants of Eschscholtzia californica which were quite self-sterile in Brazil, yielded in this country plants which were only slightly self-sterile.

It appears (17/75. Duvernoy quoted by Gartner 'Bastarderzeugung' s. 334) that certain flowers on certain plants of Lilium candidum can be fertilised more freely by pollen from a distinct individual than by their own. So, again, with the varieties of the potato. Tinzmann (17/76. 'Gardener's Chronicle' 1846 page 183.), 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 (17/77. 'Transact. Hort. Soc.' volume 7 1830 page 95.) found that he could not get fruit from P. alata and racemosa except by reciprocally fertilising them with each other's pollen; and similar facts have been observed in Germany and France. (17/78. Prof. Lecoq 'De la Fecondation' 1845 page 70; Gartner 'Bastarderzeugung' s. 64.) I have received two accounts of P. quadrangularis never producing fruit from its own pollen, but doing so freely when fertilised in one case with the pollen of P. coerulea, and in another case with that of P. edulis. But in three other cases this species fruited freely when fertilised with its own pollen; and the writer in one case attributed the favourable result to the temperature of the house having been raised from 5 deg to 10 deg Fahr. above the former temperature, after the flowers were fertilised. (17/79. 'Gardener's Chronicle' 1868 page 1341.) With respect to P. laurifolia, a cultivator of much experience has recently remarked (17/80. 'Gardener's Chronicle' 1866 page 1068.) 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 Messrs. Scott and Robertson Munro (17/81. 'Journal of Proc. of Linn. Soc.' volume 8 1864 page 1168. Mr. Robertson Munro in 'Trans. Bot. Soc.' of Edinburgh volume 9 page 399.): plants of Passiflora racemosa, coerulea, and alata flowered profusely during many years in the Botanic Gardens of Edinburgh, and, though repeatedly fertilised 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. In the case of P. coerulea three plants, two of which grew in the Botanic Gardens, were all rendered fertile, merely by impregnating each with pollen of one of the others. The same result was attained in the same manner with P. alata, but with only one plant out of three. As so many self-sterile species of Passiflora have been mentioned, it should be stated that the flowers of the annual P. gracilis are nearly as fertile with their own pollen as with that from a distinct plant; thus sixteen flowers 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. Robertson 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 (17/82. 'Amaryllidaceae' 1837 page 371; 'Journal of Hort. Soc.' volume 2 1847 page 19.) 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. 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, reginae, 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 (17/83. Loudon's 'Gardener's Magazine' volume 11 1835 page 260.) 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 (17/84. 'Gardener's Chronicle' 1850 page 470.) asserts that Amaryllis belladonna bears many more seeds when fertilised by the pollen of Brunswigia (Amaryllis of some authors) josephinae 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." (17/85. 'Journal Hort. Soc.' volume 5 page 135. The seedlings thus raised were given to the Hort. Soc.; but I find, on inquiry, that they unfortunately died the following winter.) These latter cases, however, relate to uncrossed species, like those before given with respect to Passiflora, Orchids, etc., and are here referred to only because the plants belong to the same group of Amaryllidaceae.