Part 25 (1/2)
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. .h.i.therto 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 a.s.sumed 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 the long-styled in the size of their pollen-grains; the pistil also resembles that of the long-styled, so that although the stamens and pistil are of nearly equal length, and consequently pollen is spontaneously deposited on the stigma, yet the flowers are not legitimately fertilised and yield only a very moderate supply of seed.
We thus see, firstly, that equal-styled varieties have originated in various ways, and, secondly, that the combination of the two forms in the same flower differs in completeness. With P. elatior some of the flowers on the same plant have become equal-styled, instead of all of them as in the other species.
Mr. Scott has suggested that the equal-styled varieties arise through reversion to the former h.o.m.ostyled condition of the genus. This view is supported by the remarkable fidelity with which the equal-styled variation is transmitted after it has once appeared. I have shown in Chapter 13 of my 'Variation of Animals and Plants under Domestication,' that any cause which disturbs the const.i.tution tends to induce reversion, and it is chiefly the cultivated species of Primula which become equal-styled. Illegitimate fertilisation, which is an abnormal process, is likewise an exciting cause; and with illegitimately descended long- styled plants of P. Sinensis, I have observed the first appearance and subsequent stages of this variation. With some other plants of P. Sinensis of similar parentage the flowers appeared to have reverted to their original wild condition. Again, some hybrids between P. veris and vulgaris were strictly equal-styled, and others made a near approach to this structure. All these facts support the view that this variation results, at least in part, from reversion to the original state of the genus, before the species had become heterostyled.
On the other hand, some considerations indicate, as previously remarked, that the aboriginal parent-form of Primula had a pistil which exceeded the stamens in length. The fertility of the equal-styled varieties has been somewhat modified, being sometimes greater and sometimes less than that of a legitimate union.
Another view, however, may be taken with respect to the origin of the equal- styled varieties, and their appearance may be compared with that of hermaphrodites amongst animals which properly have their s.e.xes separated; for the two s.e.xes are combined in a monstrous hermaphrodite in a somewhat similar manner as the two s.e.xual forms are combined in the same flower of an equal- styled variety of a heterostyled species.
FINAL REMARKS.
The existence of plants which have been rendered heterostyled is a highly remarkable phenomenon, as the two or three forms of the same undoubted species differ not only in important points of structure, but in the nature of their reproductive powers. As far as structure is concerned, the two s.e.xes of many animals and of some plants differ to an extreme degree; and in both kingdoms the same species may consist of males, females, and hermaphrodites. Certain hermaphrodite cirripedes are aided in their reproduction by a whole cl.u.s.ter of what I have called complemental males, which differ wonderfully from the ordinary hermaphrodite form. With ants we have males and females, and two or three castes of sterile females or workers. With Termites there are, as Fritz Muller has shown, both winged and wingless males and females, besides the workers. But in none of these cases is there any reason to believe that the several males or several females of the same species differ in their s.e.xual powers, except in the atrophied condition of the reproductive organs in the workers of social insects. Many hermaphrodite animals must unite for reproduction, but the necessity of such union apparently depends solely on their structure. On the other hand, with heterostyled dimorphic species there are two females and two sets of males, and with trimorphic species three females and three sets of males, which differ essentially in their s.e.xual powers. We shall, perhaps, best perceive the complex and extraordinary nature of the marriage arrangements of a trimorphic plant by the following ill.u.s.tration. Let us suppose that the individuals of the same species of ant always lived in triple communities; and that in one of these, a large-sized female (differing also in other characters) lived with six middle-sized and six small-sized males; in the second community a middle-sized female lived with six large- and six small-sized males; and in the third, a small-sized female lived with six large- and six middle-sized males. Each of these three females, though enabled to unite with any male, would be nearly sterile with her own two sets of males, and likewise with two other sets of males of the same size with her own which lived in the other two communities; but she would be fully fertile when paired with a male of her own size. Hence the thirty-six males, distributed by half-dozens in the three communities, would be divided into three sets of a dozen each; and these sets, as well as the three females, would differ from one another in their reproductive powers in exactly the same manner as do the distinct species of the same genus. But it is a still more remarkable fact that young ants raised from any one of the three female ants, illegitimately fertilised by a male of a different size would resemble in a whole series of relations the hybrid offspring from a cross between two distinct species of ants. They would be dwarfed in stature, and more or less, or even utterly barren. Naturalists are so much accustomed to behold great diversities of structure a.s.sociated with the two s.e.xes, that they feel no surprise at almost any amount of difference; but differences in s.e.xual nature have been thought to be the very touchstone of specific distinction. We now see that such s.e.xual differences--the greater or less power of fertilising and being fertilised--may characterise the co-existing individuals of the same species, in the same manner as they characterise and have kept separate those groups of individuals, produced during the lapse of ages, which we rank and denominate as distinct species.
CHAPTER VII.
POLYGAMOUS, DIOECIOUS, AND GYNO-DIOECIOUS PLANTS.
The conversion in various ways of hermaphrodite into dioecious plants.
Heterostyled plants rendered dioecious.
Rubiaceae.
Verbenaceae.
Polygamous and sub-dioecious plants.
Euonymus.
Fragaria.
The two sub-forms of both s.e.xes of Rhamnus and Epigaea.
Ilex.
Gyno-dioecious plants.
Thymus, difference in fertility of the hermaphrodite and female individuals.
Satureia.