These seedlings are often dwarfed and more or less sterile, like hybrids. I possessed some long-styled plants of Primula veris, which during four successive generations had been produced from illegitimate unions between long-styled plants; they were, moreover, in some degree inter-related, and had been subjected all the time to similar conditions in pots in the greenhouse. As long as they were cultivated in this manner, they grew well and were healthy and fertile. Their fertility even increased in the later generations, as if they were becoming habituated to illegitimate fertilisation. Plants of the first illegitimate generation when taken from the greenhouse and planted in moderately good soil out of doors grew well and were healthy; but when those of the two last illegitimate generations were thus treated they became excessively sterile and dwarfed, and remained so during the following year, by which time they ought to have become accustomed to growing out of doors, so that they must have possessed a weak const.i.tution.
Under these circ.u.mstances, it seemed advisable to ascertain what would be the effect of legitimately crossing long-styled plants of the fourth illegitimate generation with pollen taken from non-related short-styled plants, growing under different conditions. Accordingly several flowers on plants of the fourth illegitimate generation (i.e., great-great-grandchildren of plants which had been legitimately fertilised), growing vigorously in pots in the greenhouse, were legitimately fertilised with pollen from an almost wild short-styled cowslip, and these flowers yielded some fine capsules. Thirty other flowers on the same illegitimate plants were fertilised with their own pollen, and these yielded seventeen capsules, containing on an average thirty-two seeds. This is a high degree of fertility; higher, I believe, than that which generally obtains with illegitimately fertilised long-styled plants growing out of doors, and higher than that of the previous illegitimate generations, although their flowers were fertilised with pollen taken from a distinct plant of the same form.
These two lots of seeds were sown (for they will not germinate well when placed on bare sand) on the opposite sides of four pots, and the seedlings were thinned, so that an equal number were left on the two sides. For some time there was no marked difference in height between the two lots; and in Pot 3, Table 6/93, the self-fertilised plants were rather the tallest. But by the time that they had thrown up young flower-stems, the legitimately crossed plants revealed much the finest, and had greener and larger leaves. The breadth of the largest leaf on each plant was measured, and those on the crossed plants were on an average a quarter of an inch (exactly .28 of an inch) broader than those on the self-fertilised plants. The plants, from being too much crowded, produced poor and short flower-stems. The two finest on each side were measured; the eight on the legitimately crossed plants averaged 4.08, and the eight on the illegitimately self-fertilised plants averaged 2.93 inches in height; or as 100 to 72.
These plants after they had flowered were turned out of their pots, and planted in fairly good soil in the open ground. In the following year (1870), when in full flower, the two tallest flower-stems on each side were again measured, as shown in Table 6/93, which likewise gives the number of flower-stems produced on both sides of all the pots.
TABLE 6/93. Primula veris.
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Height: Legitimately crossed Plants.
Column 3: Number of Flower-stems produced: Legitimately crossed Plants.
Column 4: Height: Illegitimately crossed Plants.
Column 5: Number of Flower-stems produced: Illegitimately crossed Plants.
Pot 1 : 9 : 16 : 2 1/8 : 3.
Pot 1 : 8 : : 3 4/8.
Pot 2 : 7 : 16 : 6 : 3.
Pot 2 : 6 4/8 : : 5 4/8.
Pot 3 : 6 : 16 : 3 : 4.
Pot 3 : 6 2/8 : : 0 4/8.
Pot 4 : 7 3/8 : 14 : 2 5/8 : 5.
Pot 4 : 6 1/8 : : 2 4/8.
Total : 56.26 : 62 : 25.75 : 15.
The average height of the eight tallest flower-stems on the crossed plants is here 7.03 inches, and that of the eight tallest flower-stems on the self-fertilised plants 3.21 inches; or as 100 to 46. We see, also, that the crossed plants bore sixty-two flower-stems; that is, above four times as many as those (namely fifteen) borne by the self-fertilised plants. The flowers were left exposed to the visits of insects, and as many plants of both forms grew close by, they must have been legitimately and naturally fertilised. Under these circ.u.mstances the crossed plants produced 324 capsules, whilst the self-fertilised produced only 16; and these were all produced by a single plant in Pot 2, which was much finer than any other self-fertilised plant. Judging by the number of capsules produced, the fertility of an equal number of crossed and self-fertilised plants was as 100 to 5.
In the succeeding year (1871) I did not count all the flower-stems on these plants, but only those which produced capsules containing good seeds. The season was unfavourable, and the crossed plants produced only forty such flower-stems, bearing 168 good capsules, whilst the self-fertilised plants produced only two such flower-stems, bearing only 6 capsules, half of which were very poor ones. So that the fertility of the two lots, judging by the number of capsules, was as 100 to 3.5.
In considering the great difference in height and the wonderful difference in fertility between the two sets of plants, we should bear in mind that this is the result of two distinct agencies. The self-fertilised plants were the product of illegitimate fertilisation during five successive generations, in all of which, excepting the last, the plants had been fertilised with pollen taken from a distinct individual belonging to the same form, but which was more or less closely related. The plants had also been subjected in each generation to closely similar conditions. This treatment alone, as I know from other observations, would have greatly reduced the size and fertility of the offspring. On the other hand, the crossed plants were the offspring of long-styled plants of the fourth illegitimate generation legitimately crossed with pollen from a short-styled plant, which, as well as its progenitors, had been exposed to very different conditions; and this latter circ.u.mstance alone would have given great vigour to the offspring, as we may infer from the several a.n.a.logous cases already given. How much proportional weight ought to be attributed to these two agencies,--the one tending to injure the self-fertilised offspring, and the other to benefit the crossed offspring,--cannot be determined. But we shall immediately see that the greater part of the benefit, as far as increased fertility is concerned, must be attributed to the cross having been made with a fresh stock.
Primula veris.
EQUAL-STYLED AND RED-FLOWERED VAR.
I have described in my paper "On the Illegitimate Unions of Dimorphic and Trimorphic Plants" this remarkable variety, which was sent to me from Edinburgh by Mr. J. Scott. It possessed a pistil proper to the long-styled form, and stamens proper to the short-styled form; so that it had lost the heterostyled or dimorphic character common to most of the species of the genus, and may be compared with an hermaphrodite form of a bis.e.xual animal. Consequently the pollen and stigma of the same flower are adapted for complete mutual fertilisation, instead of its being necessary that pollen should be brought from one form to another, as in the common cowslip. From the stigma and anthers standing nearly on the same level, the flowers are perfectly self-fertile when insects are excluded. Owing to the fortunate existence of this variety, it is possible to fertilise its flowers in a legitimate manner with their own pollen, and to cross other flowers in a legitimate manner with pollen from another variety or fresh stock. Thus the offspring from both unions can be compared quite fairly, free from any doubt from the injurious effects of an illegitimate union.
The plants on which I experimented had been raised during two successive generations from spontaneously self-fertilised seeds produced by plants under a net; and as the variety is highly self-fertile, its progenitors in Edinburgh may have been self-fertilised during some previous generations. Several flowers on two of my plants were legitimately crossed with pollen from a short-styled common cowslip growing almost wild in my orchard; so that the cross was between plants which had been subjected to considerably different conditions. Several other flowers on the same two plants were allowed to fertilise themselves under a net; and this union, as already explained, is a legitimate one.
The crossed and self-fertilised seeds thus obtained were sown thickly on the opposite sides of three pots, and the seedlings thinned, so that an equal number were left on the two sides. The seedlings during the first year were nearly equal in height, excepting in Pot 3, Table 6/94, in which the self-fertilised plants had a decided advantage. In the autumn the plants were bedded out, in their pots; owing to this circ.u.mstance, and to many plants growing in each pot, they did not flourish, and none were very productive in seeds. But the conditions were perfectly equal and fair for both sides. In the following spring I record in my notes that in two of the pots the crossed plants are "incomparably the finest in general appearance," and in all three pots they flowered before the self-fertilised. When in full flower the tallest flower-stem on each side of each pot was measured, and the number of the flower-stems on both sides counted, as shown in Table 6/94. The plants were left uncovered, and as other plants were growing close by, the flowers no doubt were crossed by insects. When the capsules were ripe they were gathered and counted, and the result is likewise shown in Table 6/94.
TABLE 6/94. Primula veris (equal-styled, red-flowered variety).
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Height of tallest flower-stem: crossed Plants.
Column 3: Number of Flower-stems: crossed Plants.
Column 4: Number of good capsules: crossed Plants.
Column 5: Height of tallest flower-stem: self-fertilised Plants.
Column 6: Number of Flower-stems: self-fertilised Plants.
Column 7: Number of good capsules: self-fertilised Plants.
Pot 1 : 10 : 14 : 163 : 6 4/8 : 6 : 6.
Pot 2 : 8 4/8 : 12 : * : 5 : 2 : 0.
*Several, not counted.
Pot 3 : 7 4/8 : 7 : 43 : 10 4/8 : 5 : 26.
Totals : 26.0 : 33 : 206 : 22.0 : 13 : 32.
The average height of the three tallest flower-stems on the crossed plants is 8.66 inches, and that of the three on the self-fertilised plants 7.33 inches; or as 100 to 85.
All the crossed plants together produced thirty-three flower-stems, whilst the self-fertilised bore only thirteen. The number of the capsules were counted only on the plants in Pots 1 and 3, for the self-fertilised plants in Pot 2 produced none; therefore those on the crossed plants on the opposite side were not counted. Capsules not containing any good seeds were rejected. The crossed plants in the above two pots produced 206, and the self-fertilised in the same pots only 32 capsules; or as 100 to 15. Judging from the previous generations, the extreme unproductiveness of the self-fertilised plants in this experiment was wholly due to their having been subjected to unfavourable conditions, and to severe compet.i.tion with the crossed plants; for had they grown separately in good soil, it is almost certain that they would have produced a large number of capsules. The seeds were counted in twenty capsules from the crossed plants, and they averaged 24.75; whilst in twenty capsules from the self-fertilised plants the average was 17.65; or as 100 to 71. Moreover, the seeds from the self-fertilised plants were not nearly so fine as those from the crossed plants. If we consider together the number of capsules produced and the average number of contained seeds, the fertility of the crossed plants to the self-fertilised plants was as 100 to 11. We thus see what a great effect, as far as fertility is concerned, was produced by a cross between the two varieties, which had been long exposed to different conditions, in comparison with self-fertilisation; the fertilisation having been in both cases of the legitimate order.
Primula sinensis.
As the Chinese primrose is a heterostyled or dimorphic plant, like the common cowslip, it might have been expected that the flowers of both forms when illegitimately fertilised with their own pollen or with that from flowers on another plant of the same form, would have yielded less seed than the legitimately crossed flowers; and that the seedlings raised from illegitimately self-fertilised seeds would have been somewhat dwarfed and less fertile, in comparison with the seedlings from legitimately crossed seeds. This holds good in relation to the fertility of the flowers; but to my surprise there was no difference in growth between the offspring from a legitimate union between two distinct plants, and from an illegitimate union whether between the flowers on the same plant, or between distinct plants of the same form. But I have shown, in the paper before referred to, that in England this plant is in an abnormal condition, such as, judging from a.n.a.logous cases, would tend to render a cross between two individuals of no benefit to the offspring. Our plants have been commonly raised from self-fertilised seeds; and the seedlings have generally been subjected to nearly uniform conditions in pots in greenhouses. Moreover, many of the plants are now varying and changing their character, so as to become in a greater or less degree equal-styled, and in consequence highly self-fertile. From the a.n.a.logy of Primula veris there can hardly be a doubt that if a plant of Primula sinensis could have been procured direct from China, and if it had been crossed with one of our English varieties, the offspring would have shown wonderful superiority in height and fertility (though probably not in the beauty of their flowers) over our ordinary plants.
My first experiment consisted in fertilising many flowers on long-styled and short-styled plants with their own pollen, and other flowers on the same plants with pollen taken from distinct plants belonging to the same form; so that all the unions were illegitimate. There was no uniform and marked difference in the number of seeds obtained from these two modes of self-fertilisation, both of which were illegitimate. The two lots of seeds from both forms were sown thickly on opposite sides of four pots, and numerous plants thus raised. But there was no difference in their growth, excepting in one pot, in which the offspring from the illegitimate union of two long-styled plants exceeded in a decided manner in height the offspring of flowers on the same plants fertilised with their own pollen. But in all four pots the plants raised from the union of distinct plants belonging to the same form, flowered before the offspring from the self-fertilised flowers.
Some long-styled and short-styled plants were now raised from purchased seeds, and flowers on both forms were legitimately crossed with pollen from a distinct plant; and other flowers on both forms were illegitimately fertilised with pollen from the flowers on the same plant. The seeds were sown on opposite sides of Pots 1 to 4 in Table 6/95; a single plant being left on each side. Several flowers on the illegitimate long-styled and short-styled plants described in the last paragraph, were also legitimately and illegitimately fertilised in the manner just described, and their seeds were sown in Pots 5 to 8 in the same table. As the two sets of seedlings did not differ in any essential manner, their measurements are given in a single table. I should add that the legitimate unions in both cases yielded, as might have been expected, many more seeds than the illegitimate unions. The seedlings whilst half-grown presented no difference in height on the two sides of the several pots. When fully grown they were measured to the tips of their longest leaves, and the result is given in Table 6/95.
TABLE 6/95. Primula sinensis.
Heights of plants measured in inches.
Column 1: Number (Name) of Pot.
Column 2: Plants from legitimately Crossed seeds.
Column 3: Plants from illegitimately Self-fertilised seeds.