ILLEGITIMATE OFFSPRING OF HETEROSTYLED PLANTS.
Illegitimate offspring from all three forms of Lythrum salicaria.
Their dwarfed stature and sterility, some utterly barren, some fertile.
Oxalis, transmission of form to the legitimate and illegitimate seedlings.
Primula Sinensis, Illegitimate offspring in some degree dwarfed and infertile.
Equal-styled varieties of P. Sinensis, auricula, farinosa, and elatior.
P. vulgaris, red-flowered variety, Illegitimate seedlings sterile.
P. veris, Illegitimate plants raised during several successive generations, their dwarfed stature and sterility.
Equal-styled varieties of P. veris.
Transmission of form by Pulmonaria and Polygonum.
Concluding remarks.
Close parallelism between illegitimate fertilisation and hybridism.
We have hitherto treated of the fertility of the flowers of heterostyled plants, when legitimately and illegitimately fertilised. The present chapter will be devoted to the character of their offspring or seedlings. Those raised from legitimately fertilised seeds will be here called LEGITIMATE SEEDLINGS or PLANTS, and those from illegitimately fertilised seeds, ILLEGITIMATE SEEDLINGS or PLANTS. They differ chiefly in their degree of fertility, and in their powers of growth or vigour. I will begin with trimorphic plants, and I must remind the reader that each of the three forms can be fertilised in six different ways; so that all three together can be fertilised in eighteen different ways. For instance, a long-styled form can be fertilised legitimately by the longest stamens of the mid-styled and short-styled forms, and illegitimately by its own- form and mid-length and shortest stamens, also by the mid-length stamens of the mid-styled and by the shortest stamens of the short-styled form; so that the long-styled can be fertilised legitimately in two ways and illegitimately in four ways. The same holds good with respect to the mid-styled and short-styled forms. Therefore with trimorphic species six of the eighteen unions yield legitimate offspring, and twelve yield illegitimate offspring.
I will give the results of my experiments in detail, partly because the observations are extremely troublesome, and will not probably soon be repeated-- thus, I was compelled to count under the microscope above 20,000 seeds of Lythrum salicaria--but chiefly because light is thus indirectly thrown on the important subject of hybridism.
Lythrum salicaria.
Of the twelve illegitimate unions two were completely barren, so that no seeds were obtained, and of course no seedlings could be raised. Seedlings were, however, raised from seven of the ten remaining illegitimate unions. Such illegitimate seedlings when in flower were generally allowed to be freely and legitimately fertilised, through the agency of bees, by other illegitimate plants belonging to the two other forms growing close by. This is the fairest plan, and was usually followed; but in several cases (which will always be stated) illegitimate plants were fertilised with pollen taken from legitimate plants belonging to the other two forms; and this, as might have been expected, increased their fertility. Lythrum salicaria is much affected in its fertility by the nature of the season; and to avoid error from this source, as far as possible, my observations were continued during several years. Some few experiments were tried in 1863. The summer of 1864 was too hot and dry, and, though the plants were copiously watered, some few apparently suffered in their fertility, whilst others were not in the least affected. The years 1865 and, especially, 1866, were highly favourable. Only a few observations were made during 1867. The results are arranged in cla.s.ses according to the parentage of the plants. In each case the average number of seeds per capsule is given, generally taken from ten capsules, which, according to my experience, is a nearly sufficient number. The maximum number of seeds in any one capsule is also given; and this is a useful point of comparison with the normal standard--that is, with the number of seeds produced by legitimate plants legitimately fertilised. I will give likewise in each case the minimum number. When the maximum and minimum differ greatly, if no remark is made on the subject, it may be understood that the extremes are so closely connected by intermediate figures that the average is a fair one. Large capsules were always selected for counting, in order to avoid over-estimating the infertility of the several illegitimate plants.
In order to judge of the degree of inferiority in fertility of the several illegitimate plants, the following statement of the average and of the maximum number of seeds produced by ordinary or legitimate plants, when legitimately fertilised, some artificially and some naturally, will serve as a standard of comparison, and may in each case be referred to. But I give under each experiment the percentage of seeds produced by the illegitimate plants, in comparison with the standard legitimate number of the same form. For instance, ten capsules from the illegitimate long-styled plant (Number 10), which was legitimately and naturally fertilised by other illegitimate plants, contained on an average 44.2 seeds; whereas the capsules on legitimate long-styled plants, legitimately and naturally fertilised by other legitimate plants, contained on an average 93 seeds. Therefore this illegitimate plant yielded only 47 per cent of the full and normal complement of seeds.
STANDARD NUMBER OF SEEDS PRODUCED BY LEGITIMATE PLANTS OF THE THREE FORMS, WHEN LEGITIMATELY FERTILISED.
Long-styled form: Average number of seeds in each capsule, 93; Maximum number observed out of twenty-three capsules, 159.
Mid-styled form: Average number of seeds, 130; Maximum number observed out of thirty-one capsules, 151.
Short-styled form: Average number of seeds, 83.5; but we may, for the sake of brevity, say 83; Maximum number observed out of twenty-five capsules, 112.
CLa.s.sES 1 AND 2. ILLEGITIMATE PLANTS RAISED FROM LONG-STYLED PARENTS FERTILISED WITH POLLEN FROM THE MID-LENGTH OR THE SHORTEST STAMENS OF OTHER PLANTS OF THE SAME FORM.
From this union I raised at different times three lots of illegitimate seedlings, amounting altogether to 56 plants. I must premise that, from not foreseeing the result, I did not keep a memorandum whether the eight plants of the first lot were the product of the mid-length or shortest stamens of the same form; but I have good reason to believe that they were the product of the latter. These eight plants were much more dwarfed, and much more sterile than those in the other two lots. The latter were raised from a long-styled plant growing quite isolated, and fertilised by the agency of bees with its own pollen; and it is almost certain, from the relative position of the organs of fructification, that the stigma under these circ.u.mstances would receive pollen from the mid-length stamens.
All the fifty-six plants in these three lots proved long-styled; now, if the parent-plants had been legitimately fertilised by pollen from the longest stamens of the mid-styled and short-styled forms, only about one-third of the seedlings would have been long-styled, the other two-thirds being mid-styled and short-styled. In some other trimorphic and dimorphic genera we shall find the same curious fact, namely, that the long-styled form, fertilised illegitimately by its own-form pollen, produces almost exclusively long-styled seedlings. (5/1.
Hildebrand first called attention to this fact in the case of Primula Sinensis ("Botanische Zeitung" January 1, 1864 page 5); but his results were not nearly so uniform as mine.)
The eight plants of the first lot were of low stature: three which I measured attained, when fully grown, the heights of only 28, 29, and 47 inches; whilst legitimate plants growing close by were double this height, one being 77 inches.
They all betrayed in their general appearance a weak const.i.tution; they flowered rather later in the season, and at a later age than ordinary plants. Some did not flower every year; and one plant, behaving in an unprecedented manner, did not flower until three years old. In the two other lots none of the plants grew quite to their full and proper height, as could at once be seen by comparing them with the adjoining rows of legitimate plants. In several plants in all three lots, many of the anthers were either shrivelled or contained brown and tough, or pulpy matter, without any good pollen-grains, and they never shed their contents; they were in the state designated by Gartner as contabescent, which term I will for the future use. (5/2. "Beitrage zur Kenntniss der Befruchtung" 1844 page 116.) In one flower all the anthers were contabescent excepting two which appeared to the naked eye sound; but under the microscope about two-thirds of the pollen-grains were seen to be small and shrivelled. In another plant, in which all the anthers appeared sound, many of the pollen- grains were shrivelled and of unequal sizes. I counted the seeds produced by seven plants (1 to 7) in the first lot of eight plants, probably the product of parents fertilised by their own-form shortest stamens, and the seeds produced by three plants in the other two lots, almost certainly the product of parents fertilised by their own-form mid-length stamens.
[PLANT 1.
This long-styled plant was allowed during 1863 to be freely and legitimately fertilised by an adjoining illegitimate mid-styled plant, but it did not yield a single seed-capsule. It was then removed and planted in a remote place close to a brother long-styled plant Number 2, so that it must have been freely though illegitimately fertilised; under these circ.u.mstances it did not yield during 1864 and 1865 a single capsule. I should here state that a legitimate or ordinary long-styled plant, when growing isolated, and freely though illegitimately fertilised by insects with its own pollen, yielded an immense number of capsules, which contained on an average 21.5 seeds.
PLANT 2.
This long-styled plant, after flowering during 1863 close to an illegitimate mid-styled plant, produced less than twenty capsules, which contained on an average between four and five seeds. When subsequently growing in company with Number 1, by which it will have been illegitimately fertilised, it yielded in 1866 not a single capsule, but in 1865 it yielded twenty-two capsules: the best of these, fifteen in number, were examined; eight contained no seed, and the remaining seven contained on an average only three seeds, and these seeds were so small and shrivelled that I doubt whether they would have germinated.
PLANTS 3 AND 4.
These two long-styled plants, after being freely and legitimately fertilised during 1863 by the same illegitimate mid-styled plant as in the last case, were as miserably sterile as Number 2.
PLANT 5.
This long-styled plant, after flowering in 1863 close to an illegitimate mid- styled plant, yielded only four capsules, which altogether included only five seeds. During 1864, 1865, and 1866, it was surrounded either by illegitimate or legitimate plants of the other two forms; but it did not yield a single capsule.
It was a superfluous experiment, but I likewise artificially fertilised in a legitimate manner twelve flowers; but not one of these produced a capsule; so that this plant was almost absolutely barren.
PLANT 6.
This long-styled plant, after flowering during the favourable year of 1866, surrounded by illegitimate plants of the other two forms, did not produce a single capsule.
PLANT 7.
This long-styled plant was the most fertile of the eight plants of the first lot. During 1865 it was surrounded by illegitimate plants of various parentage, many of which were highly fertile, and must thus have been legitimately fertilised. It produced a good many capsules, ten of which yielded an average of 36.1 seeds, with a maximum of 47 and a minimum of 22; so that this plant produced 39 per cent of the full number of seeds. During 1864 it was surrounded by legitimate and illegitimate plants of the other two forms; and nine capsules (one poor one being rejected) yielded an average of 41.9 seeds, with a maximum of 56 and a minimum of 28; so that, under these favourable circ.u.mstances, this plant, the most fertile of the first lot, did not yield, when legitimately fertilised, quite 45 per cent of the full complement of seeds.]
In the second lot of plants in the present cla.s.s, descended from the long-styled form, almost certainly fertilised with pollen from its own mid-length stamens, the plants, as already stated, were not nearly so dwarfed or so sterile as in the first lot. All produced plenty of capsules. I counted the number of seeds in only three plants, namely Numbers 8, 9, and 10.
[PLANT 8.
This plant was allowed to be freely fertilised in 1864 by legitimate and illegitimate plants of the other two forms, and ten capsules yielded on an average 41.1 seeds, with a maximum of 73 and a minimum of 11. Hence this plant produced only 44 per cent of the full complement of seeds.
PLANT 9.
This long-styled plant was allowed in 1865 to be freely fertilised by illegitimate plants of the other two forms, most of which were moderately fertile. Fifteen capsules yielded on an average 57.1 seeds, with a maximum of 86 and a minimum of 23. Hence the plant yielded 61 per cent of the full complement of seeds.
PLANT 10.
This long-styled plant was freely fertilised at the same time and in the same manner as the last. Ten capsules yielded an average of 44.2 seeds, with a maximum of 69 and a minimum of 25; hence this plant yielded 47 per cent of the full complement of seeds.]
The nineteen long-styled plants of the third lot, of the same parentage as the last lot, were treated differently; for they flowered during 1867 by themselves so that they must have been illegitimately fertilised by one another. It has already been stated that a legitimate long-styled plant, growing by itself and visited by insects, yielded an average of 21.5 seeds per capsule, with a maximum of 35; but, to judge fairly of its fertility, it ought to have been observed during successive seasons. We may also infer from a.n.a.logy that, if several legitimate long-styled plants were to fertilise one another, the average number of seeds would be increased; but how much increased I do not know; hence I have no perfectly fair standard of comparison by which to judge of the fertility of the three following plants of the present lot, the seeds of which I counted.
[PLANT 11.
This long-styled plant produced a large crop of capsules, and in this respect was one of the most fertile of the whole lot of nineteen plants. But the average from ten capsules was only 35.9 seeds, with a maximum of 60 and a minimum of 8.
PLANT 12.
This long-styled plant produced very few capsules; and ten yielded an average of only 15.4 seeds, with a maximum of 30 and a minimum of 4.
PLANT 13.
This plant offers an anomalous case; it flowered profusely, yet produced very few capsules; but these contained numerous seeds. Ten capsules yielded an average of 71.9 seeds, with a maximum of 95 and a minimum of 29. Considering that this plant was illegitimate and illegitimately fertilised by its brother long-styled seedlings, the average and the maximum are so remarkably high that I cannot at all understand the case. We should remember that the average for a legitimate plant legitimately fertilised is 93 seeds.]
CLa.s.s 3. ILLEGITIMATE PLANTS RAISED FROM A SHORT-STYLED PARENT FERTILISED WITH POLLEN FROM OWN-FORM MID-LENGTH STAMENS.
I raised from this union nine plants, of which eight were short-styled and one long-styled; so that there seems to be a strong tendency in this form to reproduce, when self-fertilised, the parent-form; but the tendency is not so strong as with the long-styled. These nine plants never attained the full height of legitimate plants growing close to them. The anthers were contabescent in many of the flowers on several plants.