General character of cleistogamic flowers.
List of the genera producing such flowers, and their distribution in the vegetable series.
Viola, description of the cleistogamic flowers in the several species; their fertility compared with that of the perfect flowers.
O. sensitiva, three forms of cleistogamic flowers.
Miscellaneous observations on various other cleistogamic plants.
Anemophilous species producing cleistogamic flowers.
Leersia, perfect flowers rarely developed.
Summary and concluding remarks on the origin of cleistogamic flowers.
The chief conclusions which may be drawn from the observations in this volume.
It was known even before the time of Linnaeus that certain plants produced two kinds of flowers, ordinary open, and minute closed ones; and this fact formerly gave rise to warm controversies about the sexuality of plants. These closed flowers have been appropriately named cleistogamic by Dr. Kuhn. (8/1. 'Botanische Zeitung' 1867 page 65.) They are remarkable from their small size and from never opening, so that they resemble buds; their petals are rudimentary or quite aborted; their stamens are often reduced in number, with the anthers of very small size, containing few pollen-grains, which have remarkably thin transparent coats, and generally emit their tubes whilst still enclosed within the anther-cells; and, lastly, the pistil is much reduced in size, with the stigma in some cases hardly at all developed. These flowers do not secrete nectar or emit any odour; from their small size, as well as from the corolla being rudimentary, they are singularly inconspicuous. Consequently insects do not visit them; nor if they did, could they find an entrance. Such flowers are therefore invariably self-fertilised; yet they produce an abundance of seed. In several cases the young capsules bury themselves beneath the ground, and the seeds are there matured. These flowers are developed before, or after, or simultaneously with the perfect ones. Their development seems to be largely governed by the conditions to which the plants are exposed, for during certain seasons or in certain localities only cleistogamic or only perfect flowers are produced.
Dr. Kuhn, in the article above referred to, gives a list of 44 genera including species which bear flowers of this kind. To this list I have added some genera, and the authorities are appended in a footnote. I have omitted three names, from reasons likewise given in the footnote. But it is by no means easy to decide in all cases whether certain flowers ought to be ranked as cleistogamic. For instance, Mr. Bentham informs me that in the South of France some of the flowers on the vine do not fully open and yet set fruit; and I hear from two experienced gardeners that this is the case with the vine in our hothouses; but as the flowers do not appear to be completely closed it would be imprudent to consider them as cleistogamic. The flowers of some aquatic and marsh plants, for instance of Ranunculus aquatalis, Alisma natans, Subularia, Illecebrum, Menyanthes, and Euryale, remain closely shut as long as they are submerged, and in this condition fertilise themselves. (8/2. Delpino 'Sull' Opera, la Distribuzione dei Sessi nelle Piante' etc. 1867 page 30. Subularia, however, sometimes has its flowers fully expanded beneath the water, see Sir J.E. Smith 'English Flora' volume 3 1825 page 157. For the behaviour of Menyanthes in Russia see Gillibert in 'Act. Acad. St. Petersb.' 1777 part 2 page 45.--On Euryale 'Gardener's Chronicle' 1877 page 280.) They behave in this manner, apparently as a protection to their pollen, and produce open flowers when exposed to the air; so that these cases seem rather different from those of true cleistogamic flowers, and have not been included in the list. Again, the flowers of some plants which are produced very early or very late in the season do not properly expand; and these might perhaps be considered as incipiently cleistogamic; but as they do not present any of the remarkable peculiarities proper to the class, and as I have not found any full record of such cases, they are not entered in the list. When, however, it is believed on fairly good evidence that the flowers on a plant in its native country do not open at any hour of the day or night, and yet set seeds capable of germination, these may fairly be considered as cleistogamic, notwithstanding that they present no peculiarities of structure. I will now give as complete a list of the genera containing cleistogamic species as I have been able to collect.
TABLE 8.38. List of genera including cleistogamic species (chiefly after Kuhn). (8/3. I have omitted Trifolium and Arachis from the list, because Von Mohl says 'Botanische Zeitung' 1863 page 312, that the flower-stems merely draw the flowers beneath the ground, and that these do not appear to be properly cleistogamic. Correa de Mello 'Journal of the Linnean Society Botany' volume 11 1870 page 254, observed plants of Arachis in Brazil, and could never find such flowers. Plantago has been omitted because as far as I can discover it produces hermaphrodite and female flower-heads, but not cleistogamic flowers. Krascheninikowia (vel Stellaria) has been omitted because it seems very doubtful from Maximowicz' description whether the lower flowers which have no petals or very small ones, and barren stamens or none, are cleistogamic; the upper hermaphrodite flowers are said never to produce fruit, and therefore probably act as males. Moreover in Stellaria graminea, as Babington remarks 'British Botany' 1851 page 51, "shorter and longer petals accompany an imperfection of the stamens or germen."
I have added to the list the following cases: Several Acanthaceae, for which see J. Scott in 'Journal of Botany' London new series volume 1 1872 page 161. With respect to salvia see Dr. Ascherson in 'Botanische Zeitung' 1871 page 555. For Oxybaphus and Nyctaginia see Asa Gray in 'American Naturalist' November 1873 page 692. From Dr. Torrey's account of Hottonia inflata 'Bulletin of the Torrey Botanical Club' volume 2 June 1871, it is manifest that this plant produces true cleistogamic flowers. For Pavonia see Bouche in 'Sitzungsberichte d. Gesellsch. Natur. Freunde' October 20, 1874 page 90. I have added Thelymitra, as from the account given by Mr. Fitzgerald in his magnificent work on 'Australian Orchids' it appears that the flowers of this plant in its native home never open, but they do not appear to be reduced in size. Nor is this the case with the flowers of certain species of Epidendron, Cattleya, etc. see second edition of my 'Fertilisation of Orchids' page 147, which without expanding produce capsules. It is therefore doubtful whether these Orchideae ought to have been included in the list. From what Duval-Jouve says about Cryptostachys in 'Bulletin of the Soc. Bot. de France' tome 10 1863 page 195, this plant appears to produce cleistogamic flowers. the other additions to the list are noticed in my text.)
SCROPHULARINEAE: Scrophularia. Linaria. Vandellia.
ACANTHACEAE: Cryphiacanthus. Eranthemum. Daedalacanthus. Dipteracanthus. Aechmanthera. Ruellia.
LABIATAE: Lamium. Salvia.
NYCTAGINEAE: Oxybaphus. Nyctaginia.
CAMPANULACEAE: Specularia. Campanula.
CISTINEAE: Helianthemum. Lechea.
MALPIGHIACEAE: Gaudichaudia. Aspicarpa. Camarea. Janusia.
LEGUMINOSAE: Ononis. Parochaetus. Chapmannia. Stylosanthus. Lespedeza. Vicia. Lathyrus. Martinsia vel Neurocarpum. Amphicarpaea. Glycine. Galactia. Voandzeia.
GRAMINEAE: Leersia. Hordeum. Cryptostachys.
ORCHIDEAE: Schomburgkia. Cattleya. Epidendron. Thelymitra.
The first point that strikes us in considering this list of 55 genera, is that they are very widely distributed in the vegetable series. They are more common in the family of the Leguminosae than in any other, and next in order in that of the Acanthaceae and Malpighiaceae. A large number, but not all the species, of certain genera, as of Oxalis and Viola, bear cleistogamic as well as ordinary flowers. A second point which deserves notice is that a considerable proportion of the genera produce more or less irregular flowers; this is the case with about 32 out of the 55 genera, but to this subject I shall recur.
I formerly made many observations on cleistogamic flowers, but only a few of them are worth giving, since the appearance of an admirable paper by Hugo Von Mohl, whose examination was in some respects much more complete than mine. (8/4. 'Botanische Zeitung' 1863 page 309-28.) His paper includes also an interesting history of our knowledge on the subject.
The calyx of the cleistogamic flowers differs in no respect from that of the perfect ones. The petals are reduced to five minute scales; the lower one, which represents the lower lip, is considerably larger than the others, but with no trace of the spur-like nectary; its margins are smooth, whilst those of the other four scale-like petals are papillose. D. Muller of Upsala says that in the specimens which he observed the petals were completely aborted. (8/5. Ibid. 1857 page 730. This paper contains the first full and satisfactory account of any cleistogamic flower.) The stamens are very small, and only the two lower ones are provided with anthers, which do not cohere together as in the perfect flowers. The anthers are minute, with the two cells or loculi remarkably distinct; they contain very little pollen in comparison with those of the perfect flowers. The connective expands into a membranous hood-like shield which projects above the anther-cells. These two lower stamens have no vestige of the curious appendages which secrete nectar in the perfect flowers. The three other stamens are destitute of anthers and have broader filaments, with their terminal membranous expansions flatter or not so hood-like as those of the two antheriferous stamens. The pollen-grains have remarkably thin transparent coats; when exposed to the air they shrivel up quickly; when placed in water they swell, and are then 8-10/7000 of an inch in diameter, and therefore of smaller size than the ordinary pollen-grains similarly treated, which have a diameter of 13-14/7000 of an inch. In the cleistogamic flowers, the pollen-grains, as far as I could see, never naturally fall out of the anther-cells, but emit their tubes through a pore at the upper end. I was able to trace the tubes from the grains some way down the stigma. The pistil is very short, with the style hooked, so that its extremity, which is a little enlarged or funnel-shaped and represents the stigma, is directed downwards, being covered by the two membranous expansions of the antheriferous stamens. It is remarkable that there is an open passage from the enlarged funnel-shaped extremity to within the ovarium; this was evident, as slight pressure caused a bubble of air, which had been drawn in by some accident, to travel freely from one end to the other: a similar passage was observed by Michalet in V. alba. The pistil therefore differs considerably from that of the perfect flower; for in the latter it is much longer, and straight with the exception of the rectangularly bent stigma; nor is it perforated by an open passage.
The ordinary or perfect flowers have been said by some authors never to produce capsules; but this is an error, though only a small proportion of them do so. This appears to depend in some cases on their anthers not containing even a trace of pollen, but more generally on bees not visiting the flowers. I twice covered with a net a group of flowers, and marked with threads twelve of them which had not as yet expanded. This precaution is necessary, for though as a general rule the perfect flowers appear considerably before the cleistogamic ones, yet occasionally some of the latter are produced early in the season, and their capsules might readily be mistaken for those produced by the perfect flowers. Not one of the twelve marked perfect flowers yielded a capsule, whilst others under the net which had been artificially fertilised produced five capsules; and these contained exactly the same average number of seeds as some capsules from flowers outside the net which had been fertilised by bees. I have repeatedly seen Bombus hortorum, lapidarius, and a third species, as well as hive-bees, sucking the flowers of this violet: I marked six which were thus visited, and four of them produced fine capsules; the two others were gnawed off by some animal. I watched Bombus hortorum for some time, and whenever it came to a flower which did not stand in a convenient position to be sucked, it bit a hole through the spur-like nectary. Such ill-placed flowers would not yield any seed or leave descendants; and the plants bearing them would thus tend to be eliminated through natural selection.
The seeds produced by the cleistogamic and perfect flowers do not differ in appearance or number. On two occasions I fertilised several perfect flowers with pollen from other individuals, and afterwards marked some cleistogamic flowers on the same plants; and the result was that 14 capsules produced by the perfect flowers contained on an average 9.85 seeds; and 17 capsules from the cleistogamic ones contained 9.64 seeds,--an amount of difference of no significance. It is remarkable how much more quickly the capsules from the cleistogamic flowers are developed than those from the perfect ones; for instance, several perfect flowers were cross-fertilised on April 14th, 1863, and a month afterwards (May 15th) eight young cleistogamic flowers were marked with threads; and when the two sets of capsules thus produced were compared on June 3rd, there was scarcely any difference between them in size.
Viola odorata (WHITE-FLOWERED, SINGLE, CULTIVATED VARIETY).
The petals are represented by mere scales as in the last species; but differently from in the last, all five stamens are provided with diminutive anthers. Small bundles of pollen-tubes were traced from the five anthers into the somewhat distant stigma. The capsules produced by these flowers bury themselves in the soil, if it be loose enough, and there mature themselves. (8/6. Vaucher says 'Hist. Phys. des Plantes d'Europe' tome 3 1844 page 309, that V. hirta and collina likewise bury their capsules. See also Lecoq 'Geograph. Bot.' tome 5 1856 page 180.) Lecoq says that it is only these latter capsules which possess elastic valves; but I think this must be a misprint, as such valves would obviously be of no use to the buried capsules, but would serve to scatter the seeds of the sub-aerial ones, as in the other species of Viola. It is remarkable that this plant, according to Delpino, does not produce cleistogamic flowers in one part of Liguria, whilst the perfect flowers are there abundantly fertile (8/7. 'Sull' Opera, la Distribuzione dei Sessi nelle Piante' etc. 1867 page 30.); on the other hand, cleistogamic flowers are produced by it near Turin. Another fact is worth giving as an instance of correlated development: I found on a purple variety, after it had produced its perfect double flowers, and whilst the white single variety was bearing its cleistogamic flowers, many bud-like bodies which from their position on the plant were certainly of a cleistogamic nature. They consisted, as could be seen on bisecting them, of a dense mass of minute scales closely folded over one another, exactly like a cabbage-head in miniature. I could not detect any stamens, and in the place of the ovarium there was a little central column. The doubleness of the perfect flowers had thus spread to the cleistogamic ones, which therefore were rendered quite sterile.
The five stamens of the cleistogamic flowers are provided, as in the last case, with small anthers, from all of which pollen-tubes proceed to the stigma. The petals are not quite so much reduced as in V. canina, and the short pistil instead of being hooked is merely bent into a rectangle. Of several perfect flowers which I saw visited by hive-and humble-bees, six were marked, but they produced only two capsules, some of the others having been accidentally injured. M. Monnier was therefore mistaken in this case as in that of V. odorata, in supposing that the perfect flowers always withered away and aborted. He states that the peduncles of the cleistogamic flowers curve downwards and bury the ovaries beneath the soil. (8/8. These statements are taken from Professor Oliver's excellent article in the 'Natural History Review' July 1862 page 238. With respect to the supposed sterility of the perfect flowers in this genus see also Timbal-Lagrave 'Botanische Zeitung' 1854 page 772.) I may here add that Fritz Muller, as I hear from his brother, has found in the highlands of Southern Brazil a white-flowered species of violet which bears subterranean cleistogamic flowers.
Mr. Scott sent me seeds of this Indian species from the Sikkim Terai, from which I raised many plants, and from these other seedlings during several successive generations. They produced an abundance of cleistogamic flowers during the whole of each summer, but never a perfect one. When Mr. Scott wrote to me his plants in Calcutta were behaving similarly, though his collector saw the species in flower in its native site. This case is valuable as showing that we ought not to infer, as has sometimes been done, that a species does not bear perfect flowers when growing naturally, because it produces only cleistogamic flowers under culture. The calyx of these flowers is sometimes formed of only three sepals; two being actually suppressed and not merely coherent with the others; this occurred with five out of thirty flowers which were examined for this purpose. The petals are represented by extremely minute scales. Of the stamens, two bear anthers which are in the same state as in the previous species, but, as far as I could judge, each of the two cells contained only from 20 to 25 delicate transparent pollen-grains. These emitted their tubes in the usual manner. The three other stamens bore very minute rudimentary anthers, one of which was generally larger than the other two, but none of them contained any pollen. In one instance, however, a single cell of the larger rudimentary anther included a little pollen. The style consists of a short flattened tube, somewhat expanded at its upper end, and this forms an open channel leading into the ovarium, as described under V. canina. It is slightly bent towards the two fertile anthers.
This species bore in my hothouse during two years a multitude of cleistogamic flowers, which resembled in all respects those of the last species; but no perfect ones were produced. Mr. Scott informs me that in India it bears perfect flowers only during the cold season, and that these are quite fertile. During the hot, and more especially during the rainy season, it bears an abundance of cleistogamic flowers.
Many other species, besides the five now described, produce cleistogamic flowers; this is the case, according to D. Muller, Michalet, Von Mohl, and Hermann Muller, with V. elatior, lancifolia, sylvatica, palustris, mirabilis, bicolor, ionodium, and biflora. But V. tricolor does not produce them.
Michalet asserts that V. palustris produces near Paris only perfect flowers, which are quite fertile; but that when the plant grows on mountains cleistogamic flowers are produced; and so it is with V. biflora. The same author states that he has seen in the case of V. alba flowers intermediate in structure between the perfect and cleistogamic ones. According to M. Boisduval, an Italian species, V. Ruppii, never bears in France "des fleurs bien apparentes, ce qui ne l'empeche pas de fructifier."
It is interesting to observe the gradation in the abortion of the parts in the cleistogamic flowers of the several foregoing species. It appears from the statements by D. Muller and Von Mohl that in V. mirabilis the calyx does not remain quite closed; all five stamens are provided with anthers, and some pollen-grains probably fall out of the cells on the stigma, instead of protruding their tubes whilst still enclosed, as in the other species. In V. hirta all five stamens are likewise antheriferous; the petals are not so much reduced and the pistil not so much modified as in the following species. In V. nana and elatior only two of the stamens properly bear anthers, but sometimes one or even two of the others are thus provided. Lastly, in V. canina never more than two of the stamens, as far as I have seen, bear anthers; the petals are much more reduced than in V. hirta, and according to D. Muller are sometimes quite absent.
The existence of cleistogamic flowers on this plant was discovered by Michalet. (8/9. 'Bulletin Soc. Bot. de France' tome 7 1860 page 465.) They have been fully described by Von Mohl, and I can add hardly anything to his description. In my specimens the anthers of the five longer stamens were nearly on a level with the stigmas; whilst the smaller and less plainly bilobed anthers of the five shorter stamens stood considerably below the stigmas, so that their tubes had to travel some way upwards. According to Michalet these latter anthers are sometimes quite aborted. In one case the tubes, which ended in excessively fine points, were seen by me stretching upwards from the lower anthers towards the stigmas, which they had not as yet reached. My plants grew in pots, and long after the perfect flowers had withered they produced not only cleistogamic but a few minute open flowers, which were in an intermediate condition between the two kinds. In one of these the pollen-tubes from the lower anthers had reached the stigmas, though the flower was open. The footstalks of the cleistogamic flowers are much shorter than those of the perfect flowers, and are so much bowed downwards that they tend, according to Von Mohl, to bury themselves in the moss and dead leaves on the ground. Michalet also says that they are often hypogean. In order to ascertain the number of seeds produced by these flowers, I marked eight of them; two failed, one cast its seed abroad, and the remaining five contained on an average 10.0 seeds per capsule. This is rather above the average 9.2, which eleven capsules from perfect flowers fertilised with their own pollen yielded, and considerably above the average 7.9, from the capsules of perfect flowers fertilised with pollen from another plant; but this latter result must, I think, have been accidental.
Hildebrand, whilst searching various Herbaria, observed that many other species of Oxalis besides O. acetosella produce cleistogamic flowers (8/10. 'Monatsbericht der Akad. der Wiss. zu Berlin' 1866 page 369.); and I hear from him that this is the case with the heterostyled trimorphic O. incarnata from the Cape of Good Hope.
Oxalis (Biophytum) sensitiva.
This plant is ranked by many botanists as a distinct genus, but as a sub-genus by Bentham and Hooker. Many of the early flowers on a mid-styled plant in my hothouse did not open properly, and were in an intermediate condition between cleistogamic and perfect. Their petals varied from a rudiment to about half their proper size; nevertheless they produced capsules. I attributed their state to unfavourable conditions, for later in the season fully expanded flowers of the proper size appeared. But Mr. Thwaites afterwards sent me from Ceylon a number of long-styled, mid-styled, and short-styled flower-stalks preserved in spirits; and on the same stalks with the perfect flowers, some of which were fully expanded and others still in bud, there were small bud-like bodies containing mature pollen, but with their calyces closed. These cleistogamic flowers do not differ much in structure from the perfect ones of the corresponding form, with the exception that their petals are reduced to extremely minute, barely visible scales, which adhere firmly to the rounded bases of the shorter stamens. Their stigmas are much less papillose, and smaller in about the ratio of 13 to 20 divisions of the micrometer, as measured transversely from apex to apex, than the stigmas of the perfect flowers. The styles are furrowed longitudinally, and are clothed with simple as well as glandular hairs, but only in the cleistogamic flowers produced by the long- styled and mid-styled forms. The anthers of the longer stamens are a little smaller than the corresponding ones of the perfect flowers, in about the ratio of 11 to 14. They dehisce properly, but do not appear to contain much pollen. Many pollen-grains were attached by short tubes to the stigmas; but many others, still adhering to the anthers, had emitted their tubes to a considerable length, without having come in contact with the stigmas. Living plants ought to be examined, as the stigmas, at least of the long-styled form, project beyond the calyx, and if visited by insects (which, however, is very improbable) might be fertilised with pollen from a perfect flower. The most singular fact about the present species is that long-styled cleistogamic flowers are produced by the long-styled plants, and mid-styled as well as short-styled cleistogamic flowers by the other two forms; so that there are three kinds of cleistogamic and three kinds of perfect flowers produced by this one species! Most of the heterostyled species of Oxalis are more or less sterile, many absolutely so, if illegitimately fertilised with their own-form pollen. It is therefore probable that the pollen of the cleistogamic flowers has been modified in power, so as to act on their own stigmas, for they yield an abundance of seeds. We may perhaps account for the cleistogamic flowers consisting of the three forms, through the principle of correlated growth, by which the cleistogamic flowers of the double violet have been rendered double.
Dr. Kuhn has collected all the notices with respect to cleistogamic flowers in this genus, and has described from dried specimens those produced by an Abyssinian species. (8/11. 'Botanische Zeitung' 1867 page 65.) Mr. Scott sent me from Calcutta seeds of the above common Indian weed, from which many plants were successively raised during several years. The cleistogamic flowers are very small, being when fully mature under 1/20 of an inch (1.27 millimetres) in length. The calyx does not open, and within it the delicate transparent corolla remains closely folded over the ovarium. There are only two anthers instead of the normal number of four, and their filaments adhere to the corolla. The cells of the anthers diverge much at their lower ends and are only 5/700 of an inch (.181 millimetres) in their longer diameter. They contain but few pollen-grains, and these emit their tubes whilst still within the anther. The pistil is very short, and is surmounted by a bilobed stigma. As the ovary grows the two anthers together with the shrivelled corolla, all attached by the dried pollen-tubes to the stigma, are torn off and carried upwards in the shape of a little cap. The perfect flowers generally appear before the cleistogamic, but sometimes simultaneously with them. During one season a large number of plants produced no perfect flowers. It has been asserted that the latter never yield capsules; but this is a mistake, as they do so even when insects are excluded. Fifteen capsules from cleistogamic flowers on plants growing under favourable conditions contained on an average 64.2 seeds, with a maximum of 87; whilst 20 capsules from plants growing much crowded yielded an average of only 48. Sixteen capsules from perfect flowers artificially crossed with pollen from another plant contained on an average 93 seeds, with a maximum of 137. Thirteen capsules from self-fertilised perfect flowers gave an average of 62 seeds, with a maximum of 135. Therefore the capsules from the cleistogamic flowers contained fewer seeds than those from perfect flowers when cross-fertilised, and slightly more than those from perfect flowers self-fertilised.
Dr. Kuhn believes that the Abyssinian V. sessiflora does not differ specifically from the foregoing species. But its cleistogamic flowers apparently include four anthers instead of two as above described. The plants, moreover, of V. sessiflora produce subterranean runners which yield capsules; and I never saw a trace of such runners in V. nummularifolia, although many plants were cultivated.
Michalet says that short, thin, twisted branches are developed from the buds in the axils of the lower leaves, and that these bury themselves in the ground. (8/12. 'Bulletin Soc. Bot. de France' tome 7 1860 page 468.) They there produce flowers not offering any peculiarity in structure, excepting that their corollas, though properly coloured, are deformed. These flowers may be ranked as cleistogamic, as they are developed, and not merely drawn, beneath the ground.
Plants were raised from seeds sent me from Northern Italy. The sepals of the cleistogamic flowers are elongated and closely pressed together; the petals are much reduced in size, colourless, and folded over the interior organs. The filaments of the ten stamens are united into a tube, and this is not the case, according to Von Mohl, with the cleistogamic flowers of other Leguminosae. Five of the stamens are destitute of anthers, and alternate with the five thus provided. The two cells of the anthers are minute, rounded and separated from one another by connective tissue; they contain but few pollen-grains, and these have extremely delicate coats. The pistil is hook-shaped, with a plainly enlarged stigma, which is curled down, towards the anthers; it therefore differs much from that of the perfect flower. During the year 1867 no perfect flowers were produced, but in the following year there were both perfect and cleistogamic ones.
My plants produced both perfect and cleistogamic flowers; but I did not examine the latter. Some of the former were crossed with pollen from a distinct plant, and six capsules thus obtained yielded on an average 3.66 seeds, with a maximum of 5 in one. Twelve perfect flowers were marked and allowed to fertilise themselves spontaneously under a net, and they yielded eight capsules, containing on an average 2.38 seeds, with a maximum of 3 in one. Fifty-three capsules produced by the cleistogamic flowers contained on an average 4.1 seeds, so that these were the most productive of all; and the seeds themselves looked finer even than those from the crossed perfect flowers. According to Mr. Bentham O. parviflora likewise bears cleistogamic flowers; and he informs me that these flowers are produced by all three species early in the spring; whilst the perfect ones appear afterwards, and therefore in a reversed order compared with those of Viola and Oxalis. Some of the species, for instance Ononis columnae, bear a fresh crop of cleistogamic flowers in the autumn.
This plant apparently offers a case of the first stage in the production of cleistogamic flowers, for on plants growing in a state of nature, many of the flowers never expand and yet produce fine pods. Some of the buds are so large that they seem on the point of expansion; others are much smaller, but none so small as the true cleistogamic flowers of the foregoing species. As I marked these buds with thread and examined them daily, there could be no mistake about their producing fruit without having expanded.
Several other Leguminous genera produce cleistogamic flowers, as may be seen in Table 8.38; but much does not appear to be known about them. Von Mohl says that their petals are commonly rudimentary, that only a few of their anthers are developed, their filaments are not united into a tube and their pistils are hook-shaped. In three of the genera, namely Vicia, Amphicarpaea, and Voandzeia, the cleistogamic flowers are produced on subterranean stems. The perfect flowers of Voandzeia, which is a cultivated plant, are said never to produce fruit (8/13. Correa de Mello 'Journal of the Linnean Society Botany' volume 11 1870 page 254, particularly attended to the flowering and fruiting of this African plant, which is sometimes cultivated in Brazil.); but we should remember how often fertility is affected by cultivation.
Mr. A.W. Bennett has published an excellent description, with figures, of this plant. (8/14. 'Journal of the Linnean Society Botany' volume 13 1872 page 147.) He shows that the cleistogamic and perfect flowers differ in structure at a very early period of growth, so that the existence of the former cannot be due merely to the arrested development of the latter,--a conclusion which indeed follows from most of the previous descriptions. Mr. Bennett found on the banks of the Wey that the plants which bore cleistogamic flowers alone were to those bearing perfect flowers as 20 to 1; but we should remember that this is a naturalised species. The perfect flowers are usually barren in England; but Professor Asa Gray writes to me that after midsummer in the United States some or many of them produce capsules.
I can add nothing of importance to Von Mohl's description, excepting that one of the rudimentary petals shows a vestige of a nectary, as Mr. Bennett likewise found to be the case with I. fulva. As in this latter species all five stamens produce some pollen, though small in amount; a single anther contains, according to Von Mohl, not more than 50 grains, and these emit their tubes while still enclosed within it. The pollen-grains of the perfect flowers are tied together by threads, but not, so far as I could see, those of the cleistogamic flowers; and a provision of this kind would here have been useless, as the grains can never be transported by insects. The flowers of I. balsamina are visited by humble-bees (8/15. H. Muller 'Die Befruchtung' etc. page 170.), and I am almost sure that this is the case with the perfect flowers of I. noli-me-tangere. From the perfect flowers of this latter species covered with a net eleven spontaneously self-fertilised capsules were produced, and these yielded on an average 3.45 seeds. Some perfect flowers with their anthers still containing an abundance of pollen were fertilised with pollen from a distinct plant; and the three capsules thus produced contained, to my surprise, only 2, 2, and 1 seed. As I. balsamina is proterandrous, so probably is the present species; and if so, cross-fertilisation was effected by me at too early a period, and this may account for the capsules yielding so few seeds.
The first flower-stems which were thrown up by some plants in my greenhouse bore only cleistogamic flowers. The petals of small size remained permanently closed over the reproductive organs, but their white tips could just be seen between the almost 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.
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.
Another peculiarity in these flowers is that the pollen-grains generally emit their tubes whilst still enclosed within the anthers; but this is not so remarkable a fact as was formerly thought, when the case of Asclepias was alone known. (8/28. The case of Asclepias was described by R. Brown. Baillon asserts 'Adansonia' tome 2 1862 page 58, that with many plants the tubes are emitted from pollen-grains which have not come into contact with the stigma; and that they may be seen advancing horizontally through the air towards the stigma. I have observed the emission of the tubes from the pollen-masses whilst still within the anthers, in three widely distinct Orchidean genera namely Aceras, Malaxis, and Neottia: see 'The Various Contrivances by which Orchids are Fertilised' 2nd edition page 258.) It is, however, a wonderful sight to behold the tubes directing themselves in a straight line to the stigma, when this is at some little distance from the anthers. As soon as they reach the stigma or the open passage leading into the ovarium, no doubt they penetrate it, guided by the same means, whatever these may be, as in the case of ordinary flowers. I thought that they might be guided by the avoidance of light: some pollen-grains of a willow were therefore immersed in an extremely weak solution of honey, and the vessel was placed so that the light entered only in one direction, laterally or from below or from above, but the long tubes were in each case protruded in every possible direction.
As cleistogamic flowers are completely closed they are necessarily self- fertilised, not to mention the absence of any attraction to insects; and they thus differ widely from the great majority of ordinary flowers. Delpino believes that cleistogamic flowers have been developed in order to ensure the production of seeds under climatic or other conditions which tend to prevent the fertilisation of the perfect flowers. (8/29. 'Sull' Opera la Distribuzione dei Sessi nelle Piante' 1867 page 30.) I do not doubt that this holds good to a certain limited extent, but the production of a large supply of seeds with little consumption of nutrient matter or expenditure of vital force is probably a far more efficient motive power. The whole flower is much reduced in size; but what is much more important, an extremely small quantity of pollen has to be formed, as none is lost through the action of insects or the weather; and pollen contains much nitrogen and phosphorus. Von Mohl estimated that a single cleistogamic anther-cell of Oxalis acetosella contained from one to two dozen pollen-grains; we will say 20, and if so the whole flower can have produced at most 400 grains; with Impatiens the whole number may be estimated in the same manner at 250; with Leersia at 210; and with Viola nana at only 100. These figures are wonderfully low compared with the 243,600 pollen-grains produced by a flower of Leontodon, the 4,863 by an Hibiscus, or the 3,654,000 by a Paeony. (8/30. The authorities for these statements are given in my 'Effects of Cross and Self-Fertilisation' page 376.) We thus see that cleistogamic flowers produce seeds with a wonderfully small expenditure of pollen; and they produce as a general rule quite as many seeds as the perfect flowers.
That the production of a large number of seeds is necessary or beneficial to many plants needs no evidence. So of course is their preservation before they are ready for germination; and it is one of the many remarkable peculiarities of the plants which bear cleistogamic flowers, that an incomparably larger proportion of them than of ordinary plants bury their young ovaries in the ground;--an action which it may be presumed serves to protect them from being devoured by birds or other enemies. But this advantage is accompanied by the loss of the power of wide dissemination. No less than eight of the genera in the list at the beginning of this chapter include species which act in this manner, namely, several kinds of Viola, Oxalis, Vandellia, Linaria, Commelina, and at least three genera of Leguminosae. The seeds also of Leersia, though not buried, are concealed in the most perfect manner within the sheaths of the leaves. Cleistogamic flowers possess great facilities for burying their young ovaries or capsules, owing to their small size, pointed shape, closed condition and the absence of a corolla; and we can thus understand how it is that so many of them have acquired this curious habit.
It has already been shown that in about 32 out of the 55 genera in the list just referred to, the perfect flowers are irregular; and this implies that they have been specially adapted for fertilisation by insects. Moreover three of the genera with regular flowers are adapted by other means for the same end. Flowers thus constructed are liable during certain seasons to be imperfectly fertilised, namely, when the proper insects are scarce; and it is difficult to avoid the belief that the production of cleistogamic flowers, which ensures under all circumstances a full supply of seed, has been in part determined by the perfect flowers being liable to fail in their fertilisation. But if this determining cause be a real one, it must be of subordinate importance, as four of the genera in the list are fertilised by the wind; and there seems no reason why their perfect flowers should fail to be fertilised more frequently than those in any other anemophilous genus. In contrast with what we here see with respect to the large proportion of the perfect flowers being irregular, one genus alone out of the 38 heterostyled genera described in the previous chapters bears such flowers; yet all these genera are absolutely dependent on insects for their legitimate fertilisation. I know not how to account for this difference in the proportion of the plants bearing regular and irregular flowers in the two classes, unless it be that the heterostyled flowers are already so well adapted for cross-fertilisation, through the position of their stamens and pistils and the difference in power of their two or three kinds of pollen, that any additional adaptation, namely, through the flowers being made irregular, has been rendered superfluous.
Although cleistogamic flowers never fail to yield a large number of seeds, yet the plants bearing them usually produce perfect flowers, either simultaneously or more commonly at a different period; and these are adapted for or admit of cross-fertilisation. From the cases given of the two Indian species of Viola, which produced in this country during several years only cleistogamic flowers, and of the numerous plants of Vandellia and of some plants of Ononis which behaved during one whole season in the same manner, it appears rash to infer from such cases as that of Salvia cleistogama not having produced perfect flowers during five years in Germany (8/31. Dr. Ascherson 'Botanische Zeitung' 1871 page 555.), and of an Aspicarpa not having done so during several years in Paris, that these plants would not bear perfect flowers in their native homes. Von Mohl and several other botanists have repeatedly insisted that as a general rule the perfect flowers produced by cleistogamic plants are sterile; but it has been shown under the head of the several species that this is not the case. The perfect flowers Viola are indeed sterile unless they are visited by bees; but when thus visited they yield the full number of seeds. As far as I have been able to discover there is only one absolute exception to the rule that the perfect flowers are fertile, namely, that of Voandzeia; and in this case we should remember that cultivation often affects injuriously the reproductive organs. Although the perfect flowers of Leersia sometimes yield seeds, yet this occurs so rarely, as far as hitherto observed, that it practically forms a second exception to the rule.
As cleistogamic flowers are invariably fertilised, and as they are produced in large numbers, they yield altogether a much larger supply of seeds than do the perfect flowers on the same plant. But the latter flowers will occasionally be cross-fertilised, and their offspring will thus be invigorated, as we may infer from a wide-spread analogy. But of such invigoration I have only a small amount of direct evidence: two crossed seedlings of Ononis minutissima were put into competition with two seedlings raised from cleistogamic flowers; they were at first all of equal height; the crossed were then slightly beaten; but on the following year they showed the usual superiority of their class, and were to the self-fertilised plants of cleistogamic origin as 100 to 88 in mean height. With Vandellia twenty crossed plants exceeded in height twenty plants raised from cleistogamic seeds only by a little, namely, in the ratio of 100 to 94.
It is a natural inquiry how so many plants belonging to various very distinct families first came to have the development of their flowers arrested, so as ultimately to become cleistogamic. That a passage from the one state to the other is far from difficult is shown by the many recorded cases of gradations between the two states on the same plant, in Viola, Oxalis, Biophytum, Campanula, etc. In the several species of Viola the various parts of the flowers have also been modified in very different degrees. Those plants which in their own country produce flowers of full or nearly full size, but never expand (as with Thelymitra), and yet set fruit, might easily be rendered cleistogamic. Lathyrus nissolia seems to be in an incipient transitional state, as does Drosera Anglica, the flowers of which are not perfectly closed. There is good evidence that flowers sometimes fail to expand and are somewhat reduced in size, owing to exposure to unfavourable conditions, but still retain their fertility unimpaired. Linnaeus observed in 1753 that the flowers on several plants brought from Spain and grown at Upsala did not show any corolla and yet produced seeds. Asa Gray has seen flowers on exotic plants in the Northern United States which never expanded and yet fruited. With certain English plants, which bear flowers during nearly the whole year, Mr. Bennett found that those produced during the winter season were fertilised in the bud; whilst with other species having fixed times for flowering, but "which had been tempted by a mild January to put forth a few wretched flowers," no pollen was discharged from the anthers, and no seed was formed. The flowers of Lysimachia vulgaris if fully exposed to the sun expand properly, while those growing in shady ditches have smaller corollas which open only slightly; and these two forms graduate into one another in intermediate stations. Herr Bouche's observations are of especial interest, for he shows that both temperature and the amount of light affect the size of the corolla; and he gives measurements proving that with some plants the corolla is diminished by the increasing cold and darkness of the changing season, whilst with others it is diminished by the increasing heat and light. (8/32. For the statement by Linnaeus see Mohl in 'Botanische Zeitung' 1863 page 327. Asa Gray 'American Journal of Science' 2nd series volume 39 1865 page 105. Bennett in 'Nature' November 1869 page 11. The Reverend G. Henslow also says 'Gardener's Chronicle' 1877 page 271, also 'Nature' October 19, 1876 page 543, "that when the autumn draws on, and habitually in winter for such of our wild flowers as blossom at that season" the flowers are self-fertilised. On Lysimachia H. Muller 'Nature' September 1873 page 433. Bouche 'Sitzungsbericht der Gesell. Naturforsch. Freunde' October 1874 page 90.)
The belief that the first step towards flowers being rendered cleistogamic was due to the conditions to which they were exposed, is supported by the fact of various plants belonging to this class either not producing their cleistogamic flowers under certain conditions, or, on the other hand, producing them to the complete exclusion of the perfect ones. Thus some species of Viola do not bear cleistogamic flowers when growing on the lowlands or in certain districts. Other plants when cultivated have failed to produce perfect flowers during several successive years; and this is the case with Juncus bufonius in its native land of Russia. Cleistogamic flowers are produced by some species late and by others early in the season; and this agrees with the view that the first step towards their development was due to climate; though the periods at which the two sorts of flowers now appear must since have become much more distinctly defined. We do not know whether too low are too high a temperature or the amount of light acts in a direct manner on the size of the corolla, or indirectly through the male organs being first affected. However this may be, if a plant were prevented either early or late in the season from fully expanding its corolla, with some reduction in its size, but with no loss of the power of self-fertilisation, then natural selection might well complete the work and render it strictly cleistogamic. The various organs would also, it is probable, be modified by the peculiar conditions to which they are subjected within a completely closed flower; also by the principle of correlated growth, and by the tendency in all reduced organs finally to disappear. The result would be the production of cleistogamic flowers such as we now see them; and these are admirably fitted to yield a copious supply of seed at a wonderfully small cost to the plant.
I will now sum up very briefly the chief conclusions which seem to follow from the observations given in this volume. Cleistogamic flowers afford, as just stated, an abundant supply of seeds with little expenditure; and we can hardly doubt that they have had their structure modified and degraded for this special purpose; perfect flowers being still almost always produced so as to allow of occasional cross-fertilisation. Hermaphrodite plants have often been rendered monoecious, dioecious or polygamous; but as the separation of the sexes would have been injurious, had not pollen been already transported habitually by insects or by the wind from flower to flower, we may assume that the process of separation did not commence and was not completed for the sake of the advantages to be gained from cross-fertilisation. The sole motive for the separation of the sexes which occurs to me, is that the production of a great number of seeds might become superfluous to a plant under changed conditions of life; and it might then be highly beneficial to it that the same flower or the same individual should not have its vital powers taxed, under the struggle for life to which all organisms are subjected, by producing both pollen and seeds. With respect to the plants belonging to the gyno-dioecious sub-class, or those which co-exist as hermaphrodites and females, it has been proved that they yield a much larger supply of seed than they would have done if they had all remained hermaphrodites; and we may feel sure from the large number of seeds produced by many plants that such production is often necessary or advantageous. It is therefore probable that the two forms in this sub-class have been separated or developed for this special end.
Various hermaphrodite plants have become heterostyled, and now exist under two or three forms; and we may confidently believe that this has been effected in order that cross-fertilisation should be assured. For the full and legitimate fertilisation of these plants pollen from the one form must be applied to the stigma of another. If the sexual elements belonging to the same form are united the union is an illegitimate one and more or less sterile. With dimorphic species two illegitimate unions, and with trimorphic species twelve are possible. There is reason to believe that the sterility of these unions has not been specially acquired, but follows as an incidental result from the sexual elements of the two or three forms having been adapted to act on one another in a particular manner, so that any other kind of union is inefficient, like that between distinct species. Another and still more remarkable incidental result is that the seedlings from an illegitimate union are often dwarfed and more or less or completely barren, like hybrids from the union of two widely distinct species.
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