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 of gradations between the cleistogamic and perfect flowers. But that the former owe their origin wholly to arrested development is by no means the case; for various parts have been specially modified, so as to aid in the self-fertilisation of the flowers, and as a protection to the pollen; for instance, the hook-shaped pistil in Viola and in some other genera, by which the stigma is brought close to the fertile anthers,- -the rudimentary corolla of Specularia modified into a perfectly closed tympanum, and the sheath of Monochoria modified into a closed sack,—the excessively thin coats of the pollen-grains,—the anthers not being all equally aborted, and other such cases. Moreover Mr. Bennett has shown that the buds of the cleistogamic and perfect flowers of Impatiens differ at a very early period of growth.
The degree to which many of the most important organs in these degraded flowers have been reduced or even wholly obliterated, is one of their most remarkable peculiarities, reminding us of many parasitic animals. In some cases only a single anther is left, and this contains but few pollen-grains of diminished size; in other cases the stigma has disappeared, leaving a simple open passage into the ovarium. It is also interesting to note the complete loss of trifling points in the structure or functions of certain parts, which though of service to the perfect flowers, are of none to the cleistogamic; for instance the collecting hairs on the pistil of Specularia, the glands on the calyx of the Malpighiaceae, the nectar-secreting appendages to the lower stamens of Viola, the secretion of nectar by other parts, the emission of a sweet odour, and apparently the elasticity of the valves in the buried capsules of Viola odorata. We here see, as throughout nature, that as soon as any part or character becomes superfluous it tends sooner or later to disappear.