The Different Forms of Flowers on Plants of the Same Species by Charles Robert Darwin (rainbow fish read aloud txt) 📕

completely closed sepals. The pollen, which was scanty in amount, but not so scanty as in Viola or Oxalis, remained enclosed within the anthers, whence the tubes proceeded and penetrated the stigma. As the ovarium swelled the little withered corolla was carried upwards in the form of a cap. These cleistogamic flowers produced an abundance of seed. Later in the season perfect flowers appeared. With plants in a state of nature the flowers open only in the early morning, as I have been informed by Mr. Wallis, who particularly attended to the time of their flowering. In the case of D. Anglica, the still folded petals on some plants in my greenhouse opened just sufficiently to leave a minute aperture; the anthers dehisced properly, but the pollen-grains adhered in a mass to them, and thence emitted their tubes, which penetrated the stigmas. These flowers, therefore, were in an intermediate condition, and could not be called either perfect or cleistogamic.
A few miscellaneous observations may be added with respect to some other species, as throwing light on our subject. Mr. Scott states that Eranthemum ambiguum bears three kinds of flowers,--large, conspicuous, open ones, which are quite sterile,--others of intermediate size, which are open and moderately fertile--and lastly small closed or cleistogamic ones, which are perfectly fertile. (8/16. 'Journal of Botany' London new series volume 1 1872 pages 161- 4.) Ruellia tuberosa, likewise one of the Acanthaceae, produces both open and cleistogamic flowers; the latter yield from 18 to 24, whilst the former only from 8 to 10 seeds; these two kinds of flowers are produced simultaneously, whereas in several other members of the family the cleistogamic ones appear only during the hot season. According to Torrey and Gray, the North American species of Helianthemum, when growing in poor soil, produce only cleistogamic flowers. The cleistogamic flowers of Specularia perfoliata are highly remarkable, as they are closed by a tympanum formed by the rudimentary corolla, and without any trace of an opening. The stamens vary from 3 to 5 in number, as do the sepals. (8/17. Von Mohl 'Botanische Zeitung' 1863 pages 314 and 323. Dr. Bromfield 'Phytologist' volume 3 page 530, also remarks that the calyx of the cleistogamic flowers is usually only 3-cleft, while that of the perfect flower is mostly 5- cleft.) The collecting hairs on the pistil, which play so important a part in the fertilisation of the perfect flowers, are here quite absent. Drs. Hooker and Thomson state that some of the Indian species of Campanula produce two kinds of flowers; the smaller ones being borne on longer peduncles with differently formed sepals, and producing a more globose ovary. (8/18. 'Journal of the Linnean Society' volume 2 1857 page 7. See also Professor Oliver in 'Natural History Review' 1862 page 240.) The flowers are closed by a tympanum like that in Specularia. Some of the plants produce both kinds of flowers, others only one kind; both yield an abundance of seeds. Professor Oliver adds that he has seen flowers on Campanula colorata in an intermediate condition between cleistogamic and perfect ones.
The solitary almost sessile cleistogamic flowers produced by Monochoria vaginalis are differently protected from those in any of the previous cases, namely, within "a short sack formed of the membranous spathe, without any opening or fissure." There is only a single fertile stamen; the style is almost obsolete, with the three stigmatic surfaces directed to one side. Both the perfect and cleistogamic flowers produce seeds. (8/19. Dr. Kirk 'Journal of the Linnean Society' volume 8 1864 page 147.)
The cleistogamic flowers on some of the Malpighiaceae seem to be more profoundly modified than those in any of the foregoing genera. According to A. de Jussieu they are differently situated from the perfect flowers; they contain only a single stamen, instead of 5 or 6; and it is a strange fact that this particular stamen is not developed in the perfect flowers of the same species. (8/20. 'Archives du Museum' tome 3 1843 pages 35-38, 82-86, 589, 598.) The style is absent or rudimentary; and there are only two ovaries instead of three. Thus these degraded flowers, as Jussieu remarks, "laugh at our classifications, for the greater number of the characters proper to the species, to the genus, to the family, to the class disappear." I may add that their calyces are not glandular, and as, according to Kerner, the fluid secreted by such glands generally serves to protect the flowers from crawling insects, which steal the nectar without aiding in their cross-fertilisation (8/21. 'Die Schutzmittel der Bluthen gegen unberufene Gaste' 1876 page 25.), the deficiency of the glands in the cleistogamic flowers of these plants may perhaps be accounted for by their not requiring any such protection.
As the Asclepiadous genus Stapelia is said to produce cleistogamic flowers, the following case may be worth giving. I have never heard of the perfect flowers of Hoya carnosa setting seeds in this country, but some capsules were produced in Mr. Farrer's hothouse; and the gardener detected that they were the product of minute bud-like bodies, three or four of which could sometimes be found on the same umbel with the perfect flowers. They were quite closed and hardly thicker than their peduncles. The sepals presented nothing particular, but internally and alternating with them, there were five small flattened heart-shaped papillae, like rudiments of petals; but the homological nature of which appeared doubtful to Mr. Bentham and Dr. Hooker. No trace of anthers or of stamens could be detected; and I knew from having examined many cleistogamic flowers what to look for. There were two ovaries, full of ovules, quite open at their upper ends, with their edges festooned, but with no trace of a proper stigma. In all these flowers one of the two ovaries withered and blackened long before the other. The one perfect capsule, 3 1/2 inches in length, which was sent me, had likewise been developed from a single carpel. This capsule contained an abundance of plumose seeds, many of which appeared quite sound, but they did not germinate when sown at Kew. Therefore the little bud-like flower which produced this capsule probably was as destitute of pollen as were those which I examined.
Juncus bufonius and Hordeum.
All the species hitherto mentioned which produce cleistogamic flowers are entomophilous; but four genera, Juncus, Hordeum, Cryptostachys, and Leersia are anemophilous. Juncus bufonius is remarkable by bearing in parts of Russia only cleistogamic flowers, which contain three instead of the six anthers found in the perfect flowers. (8/22. See Dr. Ascherson's interesting paper in 'Botanische Zeitung' 1871 page 551.) In the genus Hordeum it has been shown by Delpino that the majority of the flowers are cleistogamic, some of the others expanding and apparently allowing of cross-fertilisation. (8/23. 'Bollettini del Comizio agrario Parmense.' Marzo e Aprile 1871. An abstract of this valuable paper is given in 'Botanische Zeitung' 1871 page 537. See also Hildebrand on Hordeum in 'Monatsbericht d. K. Akad Berlin' October 1872 page 760.) I hear from Fritz Muller that there is a grass in Southern Brazil, in which the sheath of the uppermost leaf, half a metre in length, envelopes the whole panicle; and this sheath never opens until the self-fertilised seeds are ripe. On the roadside some plants had been cut down, whilst the cleistogamic panicles were developing, and these plants afterwards produced free or unenclosed panicles of small size, bearing perfect flowers.
Leersia oryzoides.
It has long been known that this plant produces cleistogamic flowers, but these were first described with care by M. Duval-Jouve. (8/24. 'Bulletin Bot. Soc. de France' tome 10 1863 page 194.) I procured plants from a stream near Reigate, and cultivated them for several years in my greenhouse. The cleistogamic flowers are very small, and usually mature their seeds within the sheaths of the leaves. These flowers are said by Duval-Jouve to be filled by slightly viscid fluid; but this was not the case with several that I opened; but there was a thin film of fluid between the coats of the glumes, and when these were pressed the fluid moved about, giving a similarly deceptive appearance of the whole inside of the flower being thus filled. The stigma is very small and the filaments extremely short; the anthers are less than 1/50 of an inch in length or about one-third of the length of those in the perfect flowers. One of the three anthers dehisces before the two others. Can this have any relation with the fact that in some other species of Leersia only two stamens are fully developed? (8/25. Asa Gray 'Manual of Botany of the United States' 1856 page 540.) The anthers shed their pollen on the stigma; at least in one instance this was clearly the case, and by tearing open the anthers under water the grains were easily detached. Towards the apex of the anther the grains are arranged in a single row and lower down in two or three rows, so that they could be counted; and there were about 35 in each cell, or 70 in the whole anther; and this is an astonishingly small number for an anemophilous plant. The grains have very delicate coats, are spherical and about 5/7000 of an inch (.0181 millimetres), whilst those of the perfect flowers are about 7/7000 of an inch (.0254 millimetres) in diameter.
M. Duval-Jouve states that the panicles very rarely protrude from their sheaths, but that when this does happen the flowers expand and exhibit well-developed ovaries and stigmas, together with full-sized anthers containing apparently sound pollen; nevertheless such flowers are invariably quite sterile. Schreiber had previously observed that if a panicle is only half protruded, this half is sterile, whilst the still included half is fertile. Some plants which grew in a large tub of water in my greenhouse behaved on one occasion in a very different manner. They protruded two very large much-branched panicles; but the florets never opened, though these included fully developed stigmas, and stamens supported on long filaments with large anthers that dehisced properly. If these florets had opened for a short time unperceived by me and had then closed again, the empty anthers would have been left dangling outside. Nevertheless they yielded on August 17th an abundance of fine ripe seeds. Here then we have a near approach to the single case as yet known of this grass producing in a state of nature (in Germany) perfect flowers which yielded a copious supply of fruit. (8/26. Dr. Ascherson 'Botanische Zeitung' 1864 page 350.) Seeds from the cleistogamic flowers were sent by me to Mr. Scott in Calcutta, who there cultivated the plants in various ways, but they never produced perfect flowers.
In Europe Leersia oryzoides is the sole representative of its genus, and Duval- Jouve, after examining several exotic species, found that it apparently is the sole one which bears cleistogamic flowers. It ranges from Persia to North America, and specimens from Pennsylvania resembled the European ones in their concealed manner of fructification. There can therefore be little doubt that this plant generally propagates itself throughout an immense area by cleistogamic seeds, and that it can hardly ever be invigorated by cross- fertilisation. It resembles in this respect those plants which are now widely spread, though they increase solely by asexual generation. (8/27. I have collected several such cases in my 'Variation under Domestication' chapter 18 2nd edition volume 2 page 153.)
CONCLUDING REMARKS ON CLEISTOGAMIC FLOWERS.
That these flowers owe their structure primarily to the arrested development of perfect ones, we may infer from such cases as that of the lower rudimentary petal in Viola being larger than the others, like the lower lip of the perfect flower,--from a vestige of a spur in the cleistogamic flowers of Impatiens,-- from the ten stamens of Ononis being united into a tube,--and other such structures. The same inference may be drawn from the occurrence, in some instances, on the same plant of a series