The Different Forms of Flowers on Plants of the Same Species by Charles Robert Darwin (rainbow fish read aloud txt) π
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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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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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Publication Date: 12-15-2009
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