The Different Forms of Flowers on Plants of the Same Species, Charles Robert Darwin [tohfa e dulha read online .TXT] 📗
- Author: Charles Robert Darwin
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some dimorphic species, namely, Primula vulgaris and Pulmonaria angustifolia; but it entirely fails in other cases, as with Hottonia palustris and Linum grandiflorum. We shall, however, best perceive the difficulty of understanding the nature and origin of the co-adaptation between the reproductive organs of the two forms of heterostyled plants, by considering the case of Linum grandiflorum: the two forms of this plant differ exclusively, as far as we can see, in the length of their pistils; in the long-styled form, the stamens equal the pistil in length, but their pollen has no more effect on it than so much inorganic dust; whilst this pollen fully fertilises the short pistil of the other form. Now, it is scarcely credible that a mere difference in the length of the pistil can make a wide difference in its capacity for being fertilised. We can believe this the less because with some plants, for instance, Amsinckia spectabilis, the pistil varies greatly in length without affecting the fertility of the individuals which are intercrossed. So again I observed that the same plants of Primula veris and vulgaris differed to an extraordinary degree in the length of their pistils during successive seasons; nevertheless they yielded during these seasons exactly the same average number of seeds when left to fertilise themselves spontaneously under a net.
We must therefore look to the appearance of inner or hidden constitutional differences between the individuals of a varying species, of such a nature that the male element of one set is enabled to act efficiently only on the female element of another set. We need not doubt about the possibility of variations in the constitution of the reproductive system of a plant, for we know that some species vary so as to be completely self-sterile or completely self-fertile, either in an apparently spontaneous manner or from slightly changed conditions of life. Gartner also has shown that the individual plants of the same species vary in their sexual powers in such a manner that one will unite with a distinct species much more readily than another. (6/6. Gartner 'Bastarderzeugung im Pflanzenreich' 1849 page 165.) But what the nature of the inner constitutional differences may be between the sets or forms of the same varying species, or between distinct species, is quite unknown. It seems therefore probable that the species which have become heterostyled at first varied so that two or three sets of individuals were formed differing in the length of their pistils and stamens and in other co-adapted characters, and that almost simultaneously their reproductive powers became modified in such a manner that the sexual elements in one set were adapted to act on the sexual elements of another set; and consequently that these elements in the same set or form incidentally became ill-adapted for mutual interaction, as in the case of distinct species. I have elsewhere shown that the sterility of species when first crossed and of their hybrid offspring must also be looked at as merely an incidental result, following from the special co-adaptation of the sexual elements of the same species. (6/7. 'Origin of Species' 6th edition page 247; 'Variation of Animals and Plants under Domestication' 2nd edition volume 2 page 169; 'The Effects of Cross and Self-fertilisation' page 463. It may be well here to remark that, judging from the remarkable power with which abruptly changed conditions of life act on the reproductive system of most organisms, it is probable that the close adaptation of the male to the female elements in the two forms of the same heterostyled species, or in all the individuals of the same ordinary species, could be acquired only under long-continued nearly uniform conditions of life.) We can thus understand the striking parallelism, which has been shown to exist between the effects of illegitimately uniting heterostyled plants and of crossing distinct species. The great difference in the degree of sterility between the various heterostyled species when illegitimately fertilised, and between the two forms of the same species when similarly fertilised, harmonises well with the view that the result is an incidental one which follows from changes gradually effected in their reproductive systems, in order that the sexual elements of the distinct forms should act perfectly on one another.
TRANSMISSION OF THE TWO FORMS BY HETEROSTYLED PLANTS.
The transmission of the two forms by heterostyled plants, with respect to which many facts were given in the last chapter, may perhaps be found hereafter to throw some light on their manner of development. Hildebrand observed that seedlings from the long-styled form of Primula Sinensis when fertilised with pollen from the same form were mostly long-styled, and many analogous cases have since been observed by me. All the known cases are given in Tables 6.36 and 6.37.
TABLE 6.36. Nature of the offspring from illegitimately fertilised dimorphic plants.
Column 1: Species and form. Column 2: Number of long-styled offspring. Column 3: Number of short-styled offspring.
Primula veris. Long-styled form, fertilised by own-form pollen during five successive generations : 156 : 6.
Primula veris. Short-styled form, fertilised by own-form pollen : 5 : 9.
Primula vulgaris. Long-styled form, fertilised by own-form pollen during two successive generations : 69 : 0.
Primula auricula. Short-styled form, fertilised by own-form pollen, is said to produce during successive generations offspring in about the following proportions : 25 : 75.
Primula Sinensis. Long-styled form, fertilised by own-form pollen during two successive generations : 52 : 0.
Primula Sinensis. Long-styled form, fertilised by own-form pollen (Hildebrand) : 14 : 3.
Primula Sinensis. Short-styled form, fertilised by own-form pollen: 1 : 24.
Pulmonaria officinalis. Long-styled form, fertilised by own-form pollen : 11 : 0.
Polygonum fagopyrum. Long-styled form, fertilised by own-form pollen : 45 : 4.
Polygonum fagopyrum. Short-styled form, fertilised by own-form pollen : 13 : 20.
TABLE 6.37. Nature of the offspring from illegitimately fertilised trimorphic plants.
Column 1: Species and form. Column 2: Number of long-styled offspring. Column 3: Number of mid-styled offspring. Column 4: Number of short-styled offspring.
Lythrum salicaria. Long-styled form, fertilised by own-form pollen : 56 : 0 : 0.
Lythrum salicaria. Short-styled form, fertilised by own-form pollen : 1 : 0 : 8.
Lythrum salicaria. Short-styled form, fertilised by pollen from mid-length stamens of long-styled form : 4 : 0 : 8.
Lythrum salicaria. Mid-styled form, fertilised by own-form pollen : 1 : 3 : 0.
Lythrum salicaria. Mid-styled form, fertilised by pollen from shortest stamens of long-styled form : 17 : 8 : 0.
Lythrum salicaria. Mid-styled form, fertilised by pollen from longest stamens of short-styled form : 14 : 8 : 18.
Oxalis rosea. Long-styled form, fertilised during several generations by own- form pollen, produced offspring in the ratio of : 100 : 0 : 0.
Oxalis hedysaroides. Mid-styled form, fertilised by own-form pollen : 0 : 17 : 0.
We see in these two tables that the offspring from a form illegitimately fertilised with pollen from another plant of the same form belong, with a few exceptions, to the same form as their parents. For instance, out of 162 seedlings from long-styled plants of Primula veris fertilised during five generations in this manner, 156 were long-styled and only 6 short-styled. Of 69 seedlings from P. vulgaris similarly raised all were long-styled. So it was with 56 seedlings from the long-styled form of the trimorphic Lythrum salicaria, and with numerous seedlings from the long-styled form of Oxalis rosea. The offspring from the short-styled forms of dimorphic plants, and from both the mid-styled and short-styled forms of trimorphic plants, fertilised with their own-form pollen, likewise tend to belong to the same form as their parents, but not in so marked a manner as in the case of the long-styled form. There are three cases in Table 6.37, in which a form of Lythrum was fertilised illegitimately with pollen from another form; and in two of these cases all the offspring belonged to the same two forms as their parents, whilst in the third case they belonged to all three forms.
The cases hitherto given relate to illegitimate unions, but Hildebrand, Fritz Muller, and myself found that a very large proportion, or all of the offspring, from a legitimate union between any two forms of the trimorphic species of Oxalis belonged to the same two forms. A similar rule therefore holds good with unions which are fully fertile, as with those of an illegitimate nature which are more or less sterile. When some of the seedlings from a heterostyled plant belong to a different form from that of its parents, Hildebrand accounts for the fact by reversion. For instance, the long-styled parent-plant of Primula veris, from which the 162 illegitimate seedlings in Table 6.36 were derived in the course of five generations, was itself no doubt derived from the union of a long-styled and a short-styled parent; and the 6 short-styled seedlings may be attributed to reversion to their short-styled progenitor. But it is a surprising fact in this case, and in other similar ones, that the number of the offspring which thus reverted was not larger. The fact is rendered still more strange in the particular instance of P. veris, for there was no reversion until four or five generations of long-styled plants had been raised. It may be seen in both tables that the long-styled form transmits its form much more faithfully than does the short-styled, when both are fertilised with their own-form pollen; and why this should be so it is difficult to conjecture, unless it be that the aboriginal parent-form of most heterostyled species possessed a pistil which exceeded its own stamens considerably in length. (6/8. It may be suspected that this was the case with Primula, judging from the length of the pistil in several allied genera (see Mr. J. Scott 'Journal of the Linnean Society Botany' volume 8 1864 page 85). Herr Breitenbach found many specimens of Primula elatior growing in a state of nature with some flowers on the same plant long-styled, others short-styled and others equal-styled; and the long-styled form greatly preponderated in number; there being 61 of this form to 9 of the short-styled and 15 of the equal-styled.) I will only add that in a state of nature any single plant of a trimorphic species no doubt produces all three forms; and this may be accounted for either by its several flowers being separately fertilised by both the other forms, as Hildebrand supposes; or by pollen from both the other forms being deposited by insects on the stigma of the same flower.
EQUAL-STYLED VARIETIES.
The tendency of the dimorphic species of Primula to produce equal-styled varieties deserves notice. Cases of this kind have been observed, as shown in the last chapter, in no less than six species, namely, P. veris, vulgaris, Sinensis, auricula, farinosa, and elatior. In the case of P. veris, the stamens resemble in length, position and size of their pollen-grains the stamens of the short-styled form; whilst the pistil closely resembles that of the long-styled, but as it varies much in length, one proper to the short-styled form appears to have been elongated and to have assumed at the same time the functions of a long-styled pistil. Consequently the flowers are capable of spontaneous self- fertilisation of a legitimate nature and yield a full complement of seed, or even more than the number produced by ordinary flowers legitimately fertilised. With P. Sinensis, on the other hand, the stamens resemble in all respects the shorter ones proper to the long-styled form, whilst the pistil makes a near approach to that of the short-styled, but as it varies in length, it would appear as if a long-styled pistil had been reduced in length and modified in function. The flowers in this case as in the last are capable of spontaneous legitimate fertilisation, and are rather more productive than ordinary flowers legitimately fertilised. With P. auricula and farinosa the stamens resemble those of the short-styled form in length, but those of
We must therefore look to the appearance of inner or hidden constitutional differences between the individuals of a varying species, of such a nature that the male element of one set is enabled to act efficiently only on the female element of another set. We need not doubt about the possibility of variations in the constitution of the reproductive system of a plant, for we know that some species vary so as to be completely self-sterile or completely self-fertile, either in an apparently spontaneous manner or from slightly changed conditions of life. Gartner also has shown that the individual plants of the same species vary in their sexual powers in such a manner that one will unite with a distinct species much more readily than another. (6/6. Gartner 'Bastarderzeugung im Pflanzenreich' 1849 page 165.) But what the nature of the inner constitutional differences may be between the sets or forms of the same varying species, or between distinct species, is quite unknown. It seems therefore probable that the species which have become heterostyled at first varied so that two or three sets of individuals were formed differing in the length of their pistils and stamens and in other co-adapted characters, and that almost simultaneously their reproductive powers became modified in such a manner that the sexual elements in one set were adapted to act on the sexual elements of another set; and consequently that these elements in the same set or form incidentally became ill-adapted for mutual interaction, as in the case of distinct species. I have elsewhere shown that the sterility of species when first crossed and of their hybrid offspring must also be looked at as merely an incidental result, following from the special co-adaptation of the sexual elements of the same species. (6/7. 'Origin of Species' 6th edition page 247; 'Variation of Animals and Plants under Domestication' 2nd edition volume 2 page 169; 'The Effects of Cross and Self-fertilisation' page 463. It may be well here to remark that, judging from the remarkable power with which abruptly changed conditions of life act on the reproductive system of most organisms, it is probable that the close adaptation of the male to the female elements in the two forms of the same heterostyled species, or in all the individuals of the same ordinary species, could be acquired only under long-continued nearly uniform conditions of life.) We can thus understand the striking parallelism, which has been shown to exist between the effects of illegitimately uniting heterostyled plants and of crossing distinct species. The great difference in the degree of sterility between the various heterostyled species when illegitimately fertilised, and between the two forms of the same species when similarly fertilised, harmonises well with the view that the result is an incidental one which follows from changes gradually effected in their reproductive systems, in order that the sexual elements of the distinct forms should act perfectly on one another.
TRANSMISSION OF THE TWO FORMS BY HETEROSTYLED PLANTS.
The transmission of the two forms by heterostyled plants, with respect to which many facts were given in the last chapter, may perhaps be found hereafter to throw some light on their manner of development. Hildebrand observed that seedlings from the long-styled form of Primula Sinensis when fertilised with pollen from the same form were mostly long-styled, and many analogous cases have since been observed by me. All the known cases are given in Tables 6.36 and 6.37.
TABLE 6.36. Nature of the offspring from illegitimately fertilised dimorphic plants.
Column 1: Species and form. Column 2: Number of long-styled offspring. Column 3: Number of short-styled offspring.
Primula veris. Long-styled form, fertilised by own-form pollen during five successive generations : 156 : 6.
Primula veris. Short-styled form, fertilised by own-form pollen : 5 : 9.
Primula vulgaris. Long-styled form, fertilised by own-form pollen during two successive generations : 69 : 0.
Primula auricula. Short-styled form, fertilised by own-form pollen, is said to produce during successive generations offspring in about the following proportions : 25 : 75.
Primula Sinensis. Long-styled form, fertilised by own-form pollen during two successive generations : 52 : 0.
Primula Sinensis. Long-styled form, fertilised by own-form pollen (Hildebrand) : 14 : 3.
Primula Sinensis. Short-styled form, fertilised by own-form pollen: 1 : 24.
Pulmonaria officinalis. Long-styled form, fertilised by own-form pollen : 11 : 0.
Polygonum fagopyrum. Long-styled form, fertilised by own-form pollen : 45 : 4.
Polygonum fagopyrum. Short-styled form, fertilised by own-form pollen : 13 : 20.
TABLE 6.37. Nature of the offspring from illegitimately fertilised trimorphic plants.
Column 1: Species and form. Column 2: Number of long-styled offspring. Column 3: Number of mid-styled offspring. Column 4: Number of short-styled offspring.
Lythrum salicaria. Long-styled form, fertilised by own-form pollen : 56 : 0 : 0.
Lythrum salicaria. Short-styled form, fertilised by own-form pollen : 1 : 0 : 8.
Lythrum salicaria. Short-styled form, fertilised by pollen from mid-length stamens of long-styled form : 4 : 0 : 8.
Lythrum salicaria. Mid-styled form, fertilised by own-form pollen : 1 : 3 : 0.
Lythrum salicaria. Mid-styled form, fertilised by pollen from shortest stamens of long-styled form : 17 : 8 : 0.
Lythrum salicaria. Mid-styled form, fertilised by pollen from longest stamens of short-styled form : 14 : 8 : 18.
Oxalis rosea. Long-styled form, fertilised during several generations by own- form pollen, produced offspring in the ratio of : 100 : 0 : 0.
Oxalis hedysaroides. Mid-styled form, fertilised by own-form pollen : 0 : 17 : 0.
We see in these two tables that the offspring from a form illegitimately fertilised with pollen from another plant of the same form belong, with a few exceptions, to the same form as their parents. For instance, out of 162 seedlings from long-styled plants of Primula veris fertilised during five generations in this manner, 156 were long-styled and only 6 short-styled. Of 69 seedlings from P. vulgaris similarly raised all were long-styled. So it was with 56 seedlings from the long-styled form of the trimorphic Lythrum salicaria, and with numerous seedlings from the long-styled form of Oxalis rosea. The offspring from the short-styled forms of dimorphic plants, and from both the mid-styled and short-styled forms of trimorphic plants, fertilised with their own-form pollen, likewise tend to belong to the same form as their parents, but not in so marked a manner as in the case of the long-styled form. There are three cases in Table 6.37, in which a form of Lythrum was fertilised illegitimately with pollen from another form; and in two of these cases all the offspring belonged to the same two forms as their parents, whilst in the third case they belonged to all three forms.
The cases hitherto given relate to illegitimate unions, but Hildebrand, Fritz Muller, and myself found that a very large proportion, or all of the offspring, from a legitimate union between any two forms of the trimorphic species of Oxalis belonged to the same two forms. A similar rule therefore holds good with unions which are fully fertile, as with those of an illegitimate nature which are more or less sterile. When some of the seedlings from a heterostyled plant belong to a different form from that of its parents, Hildebrand accounts for the fact by reversion. For instance, the long-styled parent-plant of Primula veris, from which the 162 illegitimate seedlings in Table 6.36 were derived in the course of five generations, was itself no doubt derived from the union of a long-styled and a short-styled parent; and the 6 short-styled seedlings may be attributed to reversion to their short-styled progenitor. But it is a surprising fact in this case, and in other similar ones, that the number of the offspring which thus reverted was not larger. The fact is rendered still more strange in the particular instance of P. veris, for there was no reversion until four or five generations of long-styled plants had been raised. It may be seen in both tables that the long-styled form transmits its form much more faithfully than does the short-styled, when both are fertilised with their own-form pollen; and why this should be so it is difficult to conjecture, unless it be that the aboriginal parent-form of most heterostyled species possessed a pistil which exceeded its own stamens considerably in length. (6/8. It may be suspected that this was the case with Primula, judging from the length of the pistil in several allied genera (see Mr. J. Scott 'Journal of the Linnean Society Botany' volume 8 1864 page 85). Herr Breitenbach found many specimens of Primula elatior growing in a state of nature with some flowers on the same plant long-styled, others short-styled and others equal-styled; and the long-styled form greatly preponderated in number; there being 61 of this form to 9 of the short-styled and 15 of the equal-styled.) I will only add that in a state of nature any single plant of a trimorphic species no doubt produces all three forms; and this may be accounted for either by its several flowers being separately fertilised by both the other forms, as Hildebrand supposes; or by pollen from both the other forms being deposited by insects on the stigma of the same flower.
EQUAL-STYLED VARIETIES.
The tendency of the dimorphic species of Primula to produce equal-styled varieties deserves notice. Cases of this kind have been observed, as shown in the last chapter, in no less than six species, namely, P. veris, vulgaris, Sinensis, auricula, farinosa, and elatior. In the case of P. veris, the stamens resemble in length, position and size of their pollen-grains the stamens of the short-styled form; whilst the pistil closely resembles that of the long-styled, but as it varies much in length, one proper to the short-styled form appears to have been elongated and to have assumed at the same time the functions of a long-styled pistil. Consequently the flowers are capable of spontaneous self- fertilisation of a legitimate nature and yield a full complement of seed, or even more than the number produced by ordinary flowers legitimately fertilised. With P. Sinensis, on the other hand, the stamens resemble in all respects the shorter ones proper to the long-styled form, whilst the pistil makes a near approach to that of the short-styled, but as it varies in length, it would appear as if a long-styled pistil had been reduced in length and modified in function. The flowers in this case as in the last are capable of spontaneous legitimate fertilisation, and are rather more productive than ordinary flowers legitimately fertilised. With P. auricula and farinosa the stamens resemble those of the short-styled form in length, but those of
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