Part 40 (1/2)

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.